COVER: The jumping gall wasp, Neuroterus saltatorius (Edwards) (Hymenoptera:

Cynipidae), was first discovered in Canada in 1986 in the Victoria area. The wasp forms

galls on Garry oak, Quercus garryana Douglas, British Columbia’s only native oak, and

has since spread to cover much of the host’s distribution (primarily southeastern Vancouver

Island). The insect’s native range is the western United States, where it has several hosts in

the white oak Family. The common name is derived from the bouncing action of tiny galls

(1-1.5 mm) that fall from scorched leaves in summer, each containing an agamic female

larva. Adult illustrated on cover (body length about 1mm) and text, by Stephanie Sopow.

The Journal of the Entomological Society of British Columbia is published annually in

December by the Society.

Designed and typeset by David Raworth and David Holden.

Printed by Reprographics, Simon Fraser University, Burnaby, BC, Canada.

Printed on Recycled Paper.

J. ENTOMOL. SOC. BRIT. COLUMBIA 97, DECEMBER 2000 1

Journal

of the

Entomological Society

of British Columbia

Volume 97 Issued December 2000 ISSN #0071-0733

Directors of the Entomological Society of British Columbia 2000-2001....................:.06265:2

Allison, J.D., R.L. McIntosh, J.H. Borden and L.M. Humble. A new parasitoid (Diptera:

Tachinidae) of Acanthocinus princeps (Coleoptera: Cerambycidae) in North America.....3

Progar, R.A., M.T. AliNiazee and J.L. Olsen. The economic and environmental impact of an

IPM program on hazelnuts MOTE COIN cae eeeue seen aw irctle ci-gnc erence na ere eee ea

Li, S.Y. and I.S. Otvos. Enhancement of the activity of a nuclear sSReHOHE virus by an

optical brightener in the eastern hemlock looper, Lambdina fiscellaria fiscellaria

ber OplerasGCOMCIIGAC) 2.2. cs sere ina oak bascesbeb as einesas seve naies Ueda tieeulelubas gegevens vanes 19

Mayer, D.F., E.R. Miliczky, B.F. ue and C.A. Johansen. The bee fauna eu:

poidea) of southeastern WashingtON............cececceese cece eee eee a eessecesseeeseeeeneeens eS

Dodds, K.J., D.W. Ross and G.E. Daterman. A comparison of traps and trap trees for

capturing Douglas-fir beetle, Dendroctonus pseudotsugae (Coleoptera: Scolytidae).......33

Garland, J.A. The eae of Canada emai recent acquisitions pe in British

Columbia and Yukon.. aR elu rearesnthass Maude ianaet eae vac aeee tens eaten Oo

Miller, D.R. and B.S. eee connec of a-pinene and myrcene on attraction of

mountain pine beetle, Dendroctonus ponderosae (Coleoptera: Scolytidae) to pheromones

FM StANGS GHWESIEIN WHILE DINE... .4..6sc00hs...sevesesencstsceossesvessevseesu'ed s outings od sia asesserteevescosetene 4]

Kenner, R.D. Somatochlora kennedyi (Odonata: Corduliidae): a new species for British

Columbia, with notes on geographic variation in size and wing venation.................... 47

Scudder, G.G.E. Heteroptera (Hemiptera: Prosorrhyncha) new to Canada. Part 1.............. 51

Miller, D.R. and J.H. Borden. Pheromone interruption of pine engraver, Jps pini, by

pheromones of mountain pine beetle, Dendroctonus ponderosae (Coleoptera:

COIN MIGIAG eran pen asec ae Hace nner ue aot ce ace ilan tent Said Na ins vita inc aioic a Rane etme Bet eee 57

Kenner, R.D. Gyrinus cavatus and G. minutus (Coleoptera: Gyrinidae) in British Columbia

with comments on their nearctic distribUtions..............cccccceeseeseesese ee eee cess sees ceeesseesesOT

Morewood, W.D. Occurrence and inheritance of a colour pattern dimorphism in adults of

Hyalophora euryalus (Lepidoptera: Saturniidae)................cccc ese e eee eeeeseeseeeeee teens 73

Bomford, M.K., R.S. Vernon and P. Pats. Aphid (Homoptera: Aphididae) accumulation and

distribution near fences designed for cabbage fly (Diptera: Anthomyiidae) exclusion .....79

McGregor, R.R., D.R. Gillespie, D.M.J. Quiring and D. Higginson. Parasitism of the eggs of

Lygus shulli and Lygus elisus (Heteroptera: Miridae) by Anaphes iole (Hymenoptera:

IV RVLAN AUDI LAC Ni Za beach mete Seren ister cients cle: cee atan Peete at cv av voce ens eaeancvanelhims txt satu nv ce eaee Se: 89

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

(EHemuptera: Prosotrhyncha) i Canada....<...5ccc scsi cc ccc cee cessecceescsnsseseoscneesrenssesens oS

Kuhnholz, S., J.-H. Borden and R.L. McIntosh. The ambrosia beetle, Gnathotrichus retusus

(Coleoptera: Scolytidae) breeding in red alder, A/nus rubra (Betulaceae)................ 103

OMMGE 1 OiCON PRIB OTORS - ori oa eb atenccavetassdeatentnoniuiagacciesee anew eouaecameacael OS

J. ENTOMOL. SOC. BRIT. COLUMBIA 97, DECEMBER, 2000

DIRECTORS OF THE ENTOMOLOGICAL SOCIETY OF

BRITISH COLUMBIA FOR 2000-2001

President

Rob Cannings

Royal British Columbia Museum, Victoria

President-Elect

Lorraine Maclauchlan

BC Ministry of Forests, Kamloops

Past-President

Neville Winchester

University of Victoria, Victoria

Secretary / Treasurer

Robb Bennett

BC Ministry of Forests, 7380 Puckle Rd., Saanichton BC V8M 1W4

Editorial Committee (Journal)

Dave Raworth (Editor) Peter Belton Ken Naumann

Ward Strong (also, Editor Web Page) Lorraine Maclauchlan

H.R. MacCarthy (Editor Emeritus)

Editor (Boreus)

Phil Jones

Directors

Rene Alfaro (1st) Keith Deglow (1st) Tracey Hueppelsheuser (1st)

Hugh Barclay (2nd) Rob Cannings (2nd)

Honorary Auditor

Neville Winchester

Regional Director of National Society

Terry Shore

Canadian Forest Service, Victoria

Web page: http://www.harbour.com/commorgs/ESBC/

J. ENTOMOL. SOC. BRIT. COLUMBIA 97, DECEMBER 2000 3

A new parasitoid (Diptera: Tachinidae) of Acanthocinus

princeps (Coleoptera: Cerambycidae) in North America

JEREMY D. ALLISON’, RORY L. McINTOSH’, JOHN H. BORDEN

CENTRE FOR ENVIRONMENTAL BIOLOGY,

DEPARTMENT OF BIOLOGICAL SCIENCES, SIMON FRASER UNIVERSITY,

8888 UNIVERSITY DRIVE, BURNABY, BC, V5A 1S6

LELAND M. HUMBLE

NATURAL RESOURCES CANADA, CANADIAN FOREST SERVICE, 506 WEST

BURNSIDE ROAD, VICTORIA, BC, V8Z 1MS5

ABSTRACT

An undescribed species of Bi/laea Robineau-Desvoidy was reared from field-collected

larvae of Acanthocinus princeps (Walker) maintained on artificial diet in the

laboratory. Billaea is a novel larval parasitoid for 4. princeps with natural parasitism

levels of ca. 28%.

Key words: Acanthocinus, Billaea, larval parasitoid, Monochamus

INTRODUCTION

Cerambycid beetles are host to many parasitoids in the orders Hymenoptera (Linsley

1961; Krombein et al. 1979; Woolwine et al. 1996) and Diptera (Linsley 1961; Arnaud

1978; Campadelli and Gardenghi 1991; Tsankov and Georgiev 1991). Twelve species of

Tachinidae, including Zelia vertebrata (Say), Lixophaga variabilis (Coquillet), Ptilodexia

canescens (Walker) and Chetogena floridensis (Townsend) and eight species of Billaea

Robineau-Desvoidy (Table 1), have been confirmed as larval or pupal parasitoids of

Cerambycidae (Arnaud 1978; Campadelli and Gardenghi 1991; Tsankov and Georgiev

1991). We report the occurrence of a ninth, undescribed, species of Bil/laea reared from

cerambycid larvae from British Columbia.

MATERIALS AND METHODS

In October and November of 1998 a total of 539 cerambycid larvae of the genera

Acanthocinus and Monochamus were collected from beneath the bark of burned ponderosa

pine, Pinus ponderosae P. Laws. ex C. Laws., about 17 km north of Lytton, B.C. on the

Izman Forest Service Road of the Lillooet Forest District. We did not differentiate between

larvae of the two genera. Each larva was immediately placed in artificial media [diet

number three in Payne ef al. (1975)] in a separate 60 mL glass jar. Temperature was

maintained at 30°C through 17 November 1998, and then at 10, 15 and 6°C from 17

November 1998 to 8 January 1999, 8 January — 28 February 1999, and 28 February — 28

March 1999, respectively, to simulate diapause conditions. The photoregime was 14:10

L:D and the relative humidity was ca. 55%. To ensure consistency in the quality of food, all

larvae were transferred to clean jars containing fresh diet at monthly intervals.

' Author to whom correspondence should be addressed.

* Current address: Forest Ecosystems Branch, Saskatchewan Environment & Resource

Management, Box 3003, Prince Albert, Saskatchewan, S6V 6G1, Canada.

4 J. ENTOMOL. SOC. BRIT. COLUMBIA 97, DECEMBER 2000

RESULTS AND DISCUSSION

When the rearing programme was terminated in May 1999, 176 larvae were still alive,

and 104 adult beetles, mostly Acanthocinus princeps (Walker) with a few Monochamus

scutellatus (Say) (exact counts not kept) had eclosed. Of the 259 larvae that died, 151 had

been parasitized by an undescribed Billaea Robineau-Desvoidy species (D. M. Wood! and

J. E. O’Hara' pers. comm.). The cause of mortality is unknown for the remaining 108

larvae. A total of 56 of the parasitoid larvae were reared to adulthood. Parasitism by Billaea

n. sp. exceeded 28% of the larvae originally collected, indicating that this tachinid is a

significant source of mortality. This level of parasitism is much higher than the 0.6-7.5%

levels of parasitism of M. scutellatus by B. monohammi (Townsend) (Soper and Olsen

1963), but is similar to levels of parasitism of Saperda scalaris L. by B. triangulifera

Zetterstedt (Campadelli and Gardenghi 1991) and S. populnea L. by B. irrorata (Meigen)

(Tsankov and Georgiev 1991) (28% and 9-19%, respectively).

Table 1

Cerambycid host records for the genus Billaea Robineau-Desvoidy (Diptera:

Tachinidae) in North America and Europe. Nine species are currently placed in

Billaea in North America (Wood 1987). ;

Location Parasitoid Host Reference

North B. rutilans (F.) Enapholodes atomarius (Drury) Fattig 1949

America B. monohammi (Townsend)!* Monochamus scutellatus (Say) Aldrich 1932

M. notatus (Drury) Soper and Olson 1963

M. titillator (F.) Savely 1939

B. nipigonensis Curran?’ Rhagium inquisitor (L.)" Thomas 1955

B. sp. prob. satisfacta (West) R. inquisitor Soper and Olson 1963

B. trivittata (Curran) M. notatus Thomas 1955

M. scutellatus Thomas 1955

M. titillator Savely 1939

R. inquisitor* Thomas 1955

B. interrupta (Curran) Acanthocinus obseletus (Curran) Linsley and

Chemsak 1995

Europe B. triangulifera Zetterstedt | Saperda scalaris L. Campadelli and

Gardenghi 1991

B. irrorata (Meigen) Saperda populnea (L.) Tsankov and

Georgiev 1991

' reported as Theresia (Savely 1939; Fattig 1949)

* reported as Eutheresia (Soper and Olson 1963)

: reported as Eutheresia (Thomas 1955)

* reported as Stenocorus inquisitor (Thomas 1955)

* reported as Eutheresia (Linsley and Chemsak 1995)

The remains of two larvae from which Billaea n. sp. larvae had emerged were retained

for identification. In both instances, the host larva was not completely consumed and the

parasitoid larva had emerged from the posterior abdomen. The larval characteristics of both

specimens were consistent with those of the genus Acanthocinus (Craighead 1923; and

Duffy 1953), confirming A. princeps as a host of Billaea n. sp. This is not the first record of

parasitism by a Billaea sp. of a host in the genus Acanthocinus (Table 1) although it is a

novel host record for A. princeps. Voucher specimens of adult Billaea n. sp. have been

' Systematic Entomology Section, Eastern Cereal and Oilseed Research Centre, Agriculture

and Agri-Food Canada, Ottawa, Ontario, K1A 0C6, Canada.

J. ENTOMOL. SOC. BRIT. COLUMBIA 97, DECEMBER 2000 5

deposited in the Canadian National Collection’ (n=6) and at the Pacific Forestry Centre

(n=50), Victoria, BC. The two specimens of A. princeps from which Billaea n. sp. emerged

have also been deposited at the Pacific Forestry Centre.

ACKNOWLEDGEMENTS

We thank Tristan Mennel, Mark Sidney, Peter Katinic, Leslie Chong, Monty Wood and

James O’Hara for assistance. This research was supported by the Natural Sciences and

Engineering Research Council of Canada; the Science Council of British Columbia; Forest

Renewal British Columbia; Ainsworth Lumber Co. Ltd.; B.C. Hydro and Power Authority;

Bugbusters Pest Management Inc.; Canadian Forest Products Ltd.; Crestbrook Forest

Industries Ltd.; Donohue Forest Products Inc.; Gorman Bros. Ltd.; International Forest

Products Ltd.; Lignum Ltd.; Manning Diversified Forest Products Ltd.; Phero Tech Inc.;

Riverside Forest Products Ltd.; Slocan Forest Products Ltd.; TimberWest Ltd.; Tolko

Industries Ltd.; Weldwood of Canada Ltd.; West Fraser Mills Ltd.; Western Forest

Products Ltd.; and Weyerhaeuser Canada Ltd.

REFERENCES

Aldrich, J.M. 1932. Records of Dipterous insects of the family Tachinidae reared by the late George

Dimmock, with descriptions of one new species and notes on the genus Anetia Robineau-Desvoidy.

Proceedings of the United States National Museum 80: 1-8.

Arnaud, P.H. Jr. 1978. A Host-Parasite Catalog of North American Tachinidae (Diptera). Science and

Education Administration: United States Department of Agriculture, Washington, D.C.

Campadelli, G. and G. Gardenghi. 1991. Biological notes on Billaea triangulifera Zett. (Dipt. Tachinidae),

a parasitoid of Saperda scalaris L. (Col. Cerambycidae) (abstr.). Bolletino Dell’ Istituto Di entomologia

della Universitata Degli Studi Di Bologna 45: 181-189.

Craighead, F.C. 1923. North American cerambycid larvae. Bulletin of the Department of Agriculture,

Canada. 27:1-239.

Duffy, E.A.J. 1953. A Monograph of the Immature Stages of British and Imported Timber Beetles

(Cerambycidae). British Museum (Natural History), London.

Fattig, P.W. 1949. The Larvaevoridae (Tachinidae) or parasitic flies of Georgia. Bulletin of the Emory

University of Georgia Museum 8:1-40.

Krombein, K.V., P.D. Hurd, Jr., D.R. Smith and B.D. Burks (Eds.). 1979. Catalog of the Hymenoptera in

America north of Mexico. Vol. 1. Smithsonian Institution Press, Washington, D.C.

Linsley, E.G. 1961. The Cerambycidae of North America. Part I: Introduction. University of California

Press. Berkeley and Los Angeles. Publication Number 18.

Linsley, E.G. and J.A. Chemsak. 1995. The Cerambycidae of North America, Part VII, No. 2: Taxonomy

and Classification of the Subfamily Lamiinae, Tribes Acanthocinini through Hemilophini. University of

California Press. Berkeley, Los Angeles and London. Publication Number 114.

Payne, J.A., H. Lowman and R.R. Pate. 1975. Artificial diets for rearing the tilehorned Prionus. Annals of

the Entomological Society of America 68: 680-682.

Savely, H.E. Jr. 1939. Ecological relations of certain animals in dead pine and oak logs. Ecological

Monographs 9: 323-385.

Soper, R.S. and R.E. Olson. 1963. Survey of biota associated with Monochamus (Coleoptera:

Cerambycidae) in Maine. The Canadian Entomologist 95: 83-95.

Thomas, J.B. 1955. Notes on insects and other arthropods in red and white pine logging slash. The

Canadian Entomologist 87: 338-343.

Tsankov, G. and G. Georgiev. 1991. Records on parasitoids of smaller poplar borer, Saperda populnea

(Coleoptera: Cerambycidae) along the Danube in Bulgaria. Entomophaga 36: 493-498.

Wood, D.M. 1987. Tachinidae. p. 1193-1269 In: J.F. McAlpine (Ed.), Manual of the Nearctic Diptera Vol.

2. Research Branch Agriculture Canada Monograph 28. Ottawa, Ontario.

Woolwine, A.E., J.D. Culin and C.S. Gorsuch. 1996. Parasitoids of larval Oberea myops (Coleoptera:

Cerambycidae). Journal of Agricultural Entomology 13: 227-229.

J. ENTOMOL. SOC. BRIT. COLUMBIA 97, DECEMBER 2000 7

The economic and environmental impact of an IPM program

on hazelnuts in Oregon

'R.A. PROGAR, M.T. ALINIAZEE AND J.L. OLSEN

OREGON STATE UNIVERSITY, CORVALLIS, OR 97331-2907, USA

ABSTRACT

An integrated pest management (IPM) program based on monitoring, parasite releases,

and economic thresholds was implemented in the hazelnut industry in the early 1980's. To

assess the economic and environmental benefits of the IPM program, growers were

surveyed in 1981 to determine insecticide use in 1980, prior to the inception of the

program, and in 1998 to quantify insecticide use in 1997, after the program had been

adopted throughout the growing region. Survey respondents encompassed 23% and 20%

of the hazelnut producing acreage in 1980 and 1997 respectively. Data indicate that the

total number of annual spray applications was reduced by about 50%, resulting in an

annual industry savings of over a half-million dollars.

Key words: IPM efficacy, pesticide use pattern, environmental impact, economic impact

INTRODUCTION

Integrated pest management utilizes alternate strategies in making pest control decisions by

emphasizing increased information and by integrating cultural, biological and chemical control

methods. It often results in environmental benefits through the decreased use of pesticides and

associated reduction of environmental contamination. There are numerous examples of the

development of IPM programs (Trumble et a/.1997), and many studies that evaluate the

economic benefits of IPM programs (Trumble and Alvarado-Rodriguez 1993; Trumble et a/.

1994; White and Wetzstein 1995; Headley and Hoy 1986), yet few document both the

economic and environmental savings that result from a successful IPM program on a regional

scale. Concerns over the impact of pesticide residues on food and in the environment

(Pimentel et al. 1993) are causing industry-wide regulation of insecticide use and changing the

way exposure to insecticides is assessed in the environment as set forth in the US EPA’s Food

Quality Protection Act of 1996. These concerns are causing the reduction or elimination of

insecticides and changing our perspective of IPM from spray-based management to an

ecosystem perspective by focusing on predators and parasitoids, and alternative methods of

pest control.

Economics and insecticide use patterns are fundamental to IPM practices and should be

used to measure program success. Studies suggest that it is conceivable to reduce pesticide use

in the US by 35-50% without a significant loss of crop yield (Office of Technology

Assessment 1979; National Academy of Sciences 1989). In our study, we summarize the

reduction in insecticide use and economic impact of a program to control the major insect

pests of hazelnuts (Corylus avellana L.).

The list of insects and mites associated with hazelnut trees is long, representing almost all

of the major insect and mite groups. In Oregon, over 150 species have been found on hazelnut

‘Correspondence to: Robert Progar, FHP USDA FS 1249 S. Vinnell Way, Suite 200,

Boise, ID 83709 Phone: 208-373-4226 Fax: 208-303-4111 email:rprogar@fs.fed.us

8 J. ENTOMOL. SOC. BRIT. COLUMBIA 97, DECEMBER 2000

trees; most are harmless, over half are beneficial, only two-dozen or so species are injurious,

and of those only six or so are considered important pests (AliNiazee 1998). Although there

are numerous potential insect and mite pests on hazelnuts in Oregon, only four have warranted

consistent insecticide application (AliNiazee 1994). It should be noted that pest incidence and

importance change with time and orchard management practices.

Prior to the development of an integrated pest management program, insecticide use was

widespread. This practice resulted in resistance by the filbert aphid, Myzocallis coryli (Goetze)

(Homoptera: Aphidae) reoccurring every | or 2 years (AliNiazee and Messing 1995; Katundu

and AliNiazee 1990), outbreaks of secondary pests, and rapid resurgence of primary pests.

These outbreaks required repeated application of insecticides that further aggravated pest

conditions. As a result, by the early 1970's, as many as five different insecticide applications

were applied each season to control hazelnut insect pests (AliNiazee 1977).

Research conducted during the 1970's led to the formation of an integrated pest

management program on hazelnuts in Oregon (AliNiazee 1977). In 1982, the USDA funded a

4-year project to develop an IPM program in Oregon hazelnut orchards. This program entailed

the establishment of economic injury levels for hazelnut pests (Fisher 1984; Calkin et al. 1984;

Calkin and Fisher 1985), and design and implementation of a scouting and monitoring

program which remains in use by hazelnut growers (Olsen et al. 2000). These efforts resulted

in establishing levels of tolerance (1%) and economic damage for the primary pest of

hazelnuts, the filbertworm, and initiated pheromone trapping as a viable method of monitoring

populations and timing spray applications. Before IPM, light trapping was used to determine

adult emergence and time of spray application. However, there were no existing ways to

measure population levels, therefore sprays were applied based on the presence of filbertworm

moths in trap catches. In addition, sprays were applied to control other perceived insect pests

based on their presence or simply by the calendar because there were no established economic

levels of concern. Lack of knowledge of a pest’s status strongly contributed to the overuse of

insecticides on hazelnuts. As a direct result of the IPM program growers monitor their own

orchards or employ field scouts to assess population levels. This level of current information

enables growers to determine the need, timing and location of spray applications.

Concurrent with the establishment of the IPM program, a parasitic wasp (Messing and

AliNiazee 1989) was released as a biological control of the filbert aphid. The success of this

classical biological control program aided in the implementation of the IPM program in

the1980's (AliNiazee 1991; AliNiazee and Messing 1995), and nearly eliminated all

insecticide sprays applied against aphids. By allowing early-season beneficial insects to

become established it also has indirectly reduced the application of insecticides on other insect

pests in hazelnut orchards.

In this paper we present data from a survey of hazelnut growers conducted in 1981 prior to

the inception of an IPM program on hazelnuts, and contrast it with data from a similar survey

conducted in 1998, after adoption of the program by hazelnut growers. Our objectives in this

study were to evaluate the economic and environmental impacts of an industry-wide IPM

program to control the primary hazelnut insect pests.

METHODS

A survey of hazelnut growers was conducted in 1981 to assess grower pesticide use

patterns in 1980, 1.e., prior to the initiation of an IPM program in hazelnut orchards in Oregon

(Progar and AliNiazee 1999). In 1998, a similar survey of Oregon hazelnut growers was

conducted to determine changes in insecticide use patterns resulting from adoption of hazelnut

IPM (i.e., in 1997). The number of hazelnut growers in Oregon has declined from over 1,000

to about 800, while the total area has increased, indicating a trend toward larger orchard size

J. ENTOMOL. SOC. BRIT. COLUMBIA 97, DECEMBER 2000 9

or grower-managed area (Rowley 1997). Table 1 summarizes the hazelnut area for each

survey. The surveys represent 23.5% (171 responses) and 20% (80 responses) of total

hazelnut-bearing area in 1980 and 1997, respectively. All insecticide quantities are expressed

in kg of active ingredient (a.i.) because not all growers use the same pesticide formulations.

Table 1

Survey summary data of hazelnut orchard area in Oregon.

1980 1997

Number of growers 1,063 826

Total hazelnut hectares 10,316 2124

Bearing hectares 8,741 11,412

Non-bearing hectares 1,574 708

Hectares represented in the survey 2,383 2,501

Bearing hectares (survey) 2,058 229i

Non-bearing hectares (survey) B25 210

% total hectares represented by the survey 200) 20.07

The U.S. Bureau of Labor Statistics Consumer Price Index (CPI) regional index (Bureau of

Labor Statistics Data 1998) was used to compare pest control costs between 1980 and 1997.

The 1980 index value of 247 was compared with the 1997 index value of 469 to express 1980

dollars in 1997 values.

Costs for different pesticides increased disproportionately, e.g.,a kg of Sevin® (carbaryl)

increased in cost by 56% from 1980 to 1997; whereas Guthion® (azinphos-methyl) increased

in cost by 157%. Many of the insecticides used in 1980 were no longer registered for use on

hazelnuts in 1997, and newer, more efficient compounds were used in 1997 that were not

available in 1980. Therefore, direct comparison of costs associated with specific insecticides

cannot be made, however total pesticide costs can be compared. The cost to apply an

insecticide treatment to a hectare of hazelnuts has increased from $20.34 (unadjusted dollars

US) in 1980 to $50.06 (Seavert and Olsen 1999) in 1997.

RESULTS AND DISCUSSION

Filbertworm, Cydia latiferreana Walsingham (Lepidoptera: Tortricidae)

Filbertworm is the primary insect pest in hazelnut orchards. Because there is an industry

standard of less than a 1% tolerance for filbertworm infestation, the percentage of hazelnut

orchard area treated remains about the same before and after the IPM program (Table 2).

However, the composition of the insecticides used to control filbertworm has changed and the

amount of insecticide active ingredient (a.1.) has declined dramatically (Table 3).

Table 2

Primary pests in Oregon hazelnut orchards and percent of growers and orchard area using

insecticides to control them.

% growers % ha % growers % ha

Pest (1980) (1980) (1997) ~ (1997)

Filbertworm 88.2 95.8 87.5 94.0

Filbert leafroller 38.1 57.1 16.2 28.6

Obliquebanded leafroller 5.9 6.4 2.5 aS

Filbert aphid 48.8 69.0 5.0 6.3

J. ENTOMOL. SOC. BRIT. COLUMBIA 97, DECEMBER 2000

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J. ENTOMOL. SOC. BRIT. COLUMBIA 97, DECEMBER 2000 1

In 1980, an estimated 39,916 kg (a.i.) of insecticides were applied to control filbertworm

on approximately 96% of the hazelnut orchard area by 88% of the growers (Tables 2 and 3). In

1997, only an estimated 1,453 kg of insecticide (a.i.) were applied to control filbertworm by

87% of the growers on 94% of the hazelnut area, indicating higher efficiencies and

effectiveness of insecticide application. The most common insecticide used in 1997 was

Asana® (esfenvalerate, a pyrethroid), with a small fraction of Guthion® (azinophos-methy!)

and Lorsban® (chlorpyrifos) (Table 3). Asana® was applied in amounts 10 to 20-fold less a.1.

because it was more effective than insecticides applied 16 years earlier and can be applied in

smaller amounts. Although the same portion of growers are treating the same percentage of

area for filbertworm in 1997 as in 1980, the change from organophosphates (OP) and

carbamate to esfenvalerate resulted in a large decline in the amount of insecticide a.i. applied

and an enormous benefit to the environment. Only an estimated 5% of the growers used OP’s

on 0.21% of the hazelnut area in 1997 as opposed to an estimated 24% of the growers on 27%

of the area in 1980 (Table 3).

There is currently an effective IPM system in place in commercial hazelnut orchards that

incorporates an online degree-day model, and scouting and monitoring for adult filbertworm

moths. The decrease in total pesticide use may be attributed to more efficient monitoring with

pheromone trapping rather than the previous method of light-trapping, better timing and

targeting of insecticide applications, more efficient spray equipment, and the shift from

carbamate and organo-phosphate insecticides to synthetic pyrethroids.

Although the quantity of insecticide used to control filbertworm declined by an estimated

38,463 kg (a.1.) from 1980 to 1997, the estimated cost of control was $1,275,832 in 1997 vs.

$1,066,658 in 1980 (converted to 1997 dollars), a 20% increase in expense to control

filbertworm (Table 7).

Filbert Leafroller (European Leafroller), Archips rosanus (L.) (Lepidoptera: Tortricidae)

In 1980, an estimated 10,440 kg (a.1.) of insecticide were applied by 38% of the growers on

57% of the hazelnut area to control filbert leafroller (Tables 2 and 4). In 1997, 2,998 kg of

insecticide (a.i.) were applied by 16% of the growers on 29% of the hazelnut area. Lorsban®

was the primary insecticide applied followed by a small percentage of Guthion®. The area

treated in 1997 was about half that of 1980 and less than a third the amount of pesticide a.i.

was used to control leafroller, resulting in an estimated annual reduction of 7,445 kg of

insecticide a.i. during the 16-year period (Table 4).

The adoption of an IPM program on hazelnuts has significantly reduced the use of

insecticides to control filbert leafroller. The emergence of filbert leafroller is now predicted by

degree-day modeling, and there are more accurate methods of monitoring to assess levels of

economic injury. Also of importance are secondary effects attributed to the effective biological

control of the filbert aphid. The elimination of the early-season treatments for the filbert aphid

may enable the establishment of populations of beneficial insects that prey on filbert

leafrollers. Few leafrollers have been observed in abandoned hazelnut orchards, in contrast to

managed orchards where leafroller populations are continually building.

The estimated cost of insecticide treatment for the control of filbert leafroller was $213,688

in 1980 ($406,007 1997 dollars) . In 1997 the estimated cost was $274,190 (Table 4). This is

a decrease in cost of $131,817 (Table 7), corresponding to a reduction of more than 7,445 kg

of insecticide (a.1.), and a decrease of nearly 25% in the area treated with insecticide.

Obliquebanded Leafroller, Choristoneura rosaceana (Harris), (Lepidoptera: Tortricidae)

Obliquebanded leafroller (OBLR) populations occasionally increase to levels of economic

injury. However, first generation OBLR populations are managed when sprays are applied to

control filbert leafroller since they are present concurrently. In 1980, 1,862 kg of insecticide

J. ENTOMOL. SOC. BRIT. COLUMBIA 97, DECEMBER 2000

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J. ENTOMOL. SOC. BRIT. COLUMBIA 97, DECEMBER 2000 15

(a.i.) were applied by 6% of the growers to 5% of the hazelnut area to control OBLR (Tables 2

and 5). In 1997, 153 kg of insecticide (a.1.) were applied by 2.5% of the growers to 2.5% of

the hazelnut area. The insecticides used were Asana® and Guthion®.

The pest status of the obliquebanded leafroller has declined in hazelnut orchards during the

16-year period between 1980 and 1997. However some growers (2.5%) still apply insecticides

to control this pest. Cost adjustments between 1980 and 1997 show that an annual saving of

over $26,000 was achieved by adopting IPM practices in hazelnut orchards (Tables 5 and 7).

As observed with control of filbert leafroller, a decrease of greater than 50% in the total

hectares treated occurred as a result of the establishment of an IPM program to manage pests

in hazelnut orchards.

Filbert Aphid, Myzocallis coryli (Goeze) (Homoptera: Aphidae)

The most dramatic change in insecticide use patterns in hazelnut orchards has occurred in

the control of the filbert aphid: In 1980, 6,809 kg a.1. of insecticide were applied by 49% of the

growers on 69% of the hazelnut area to control this pest (Tables 2 and 6). In 1997, 440 kg of

insecticide (a.i.) were applied by 5% of the growers to 6% of the hazelnut area. A 15-fold

reduction in the volume of insecticides, and a 10-fold reduction in the area treated occurred

during the 16-year interval between surveys.

The filbert aphid was a serious pest of hazelnuts; reproducing parthenogenetically, it has 6-

8 generations each year (AliNiazee and Messing 1995). It was an ideal candidate for the

development of resistance that occurred every 1-3 years (Katundu and AliNiazee 1990).

Therefore, finding and establishing an effective biological control was highly desirable. From

the results of natural enemy surveys, it was concluded that filbert aphid was a suitable

candidate for a classical biological control program based on the introduction of a host-

specific parasitoid. During the 1984-1985 seasons, Trioxys pallidus Haliday (Hymenoptera:

Aphidiidae), an effective parasitoid from Europe was introduced by Messing and AliNiazee

(1989) to control the aphid. The parasitoid readily established; studies conducted in 1987 and

1988 showed that 7: pallidus had an average level of parasitism of 25 -50% (AliNiazee and

Messing 1995). This biological control program is noted as one of the most successful

introductions of a biological control agent on record (AliNiazee and Messing 1995), and it

resulted in an important reduction in the use of insecticides on hazelnuts. An estimated 5% of

the hazelnut growers are currently using insecticides on 6% of the filbert acreage to control

filbert aphid - a reduction in the use of insecticides by 90% of the growers on 91% of the

hazelnut area. This translates to a vast environmental benefit in terms of the total reduction of

pesticides used in Oregon hazelnut orchards.

Not only has the establishment of the 7. pallidus wasp had a favorable impact on the

environment, but it has resulted in large economic savings as well. The reduction in pesticide

use on hazelnuts has directly increased the profitability of growing hazelnuts in Oregon. The

total area treated for filbert aphid was reduced from 69% to 6%, a reduction of 91%. This

reduction of insecticide use on filbert aphid has resulted in an annual savings of nearly one-

half-million dollars (Tables 6 and 7).

In summary, the insecticide use pattern on hazelnuts in Oregon has changed dramatically

due to the establishment of a successful IPM program. A key component of this program was

the successful release of a parasitic wasp as a biological control agent. Additionally, more

effective sampling and monitoring methods for filbert leafroller and obliquebanded leafroller

and the establishment of economic levels of injury have reduced the use of insecticides to

control infestations. The adoption of an effective IPM system and effective biological control

agent of a single pest have beneficially influenced the entire pest management strategy for

hazelnuts; reducing grower costs by large amounts each year, and significantly reducing

16 J. ENTOMOL. SOC. BRIT. COLUMBIA 97, DECEMBER 2000

environmental pollution associated with the production of an agricultural commodity in an

environmentally sensitive area.

Table 7

Costs to control hazelnut pests using the CPI ratio of 469/247 (1.9) to express 1980 costs

as 1997 values.

Insect pest Estimated cost to Value in 1997 $ Estimated cost to Estimated change

control in 1980 control in 1997

Filbertworm 560,783 1,065,488 275,832 +210,344

Filbert leafroller 212,490 403,731 274,190 -129,541

OBLR 31,304 59,478 33,305 -26,143

Filbert aphid 323,374 614,411 48,666 -565,745

Total $1,127,951 $2,143,108 $1,632,023 -$511,085

ACKNOWLEDGEMENTS

Drs. Russell Messing and Glen Fisher have been instrumental in establishing the hazelnut

IPM program and conducting the 1981 survey. We are thankful for their assistance in this

project.

REFERENCES

AliNiazee, M.T. 1977. Insect pest management in Oregon filberts. Proceedings of the Nut Growers Society of

Oregon, Washington and British Columbia 62: 37-40.

AliNiazee, M. T. 1991. Biological control of the filbert aphid, Myzocallis coryli in hazelnut orchards.

Proceedings of the Nut Growers Society of Oregon, Washington and British Columbia 76: 46-53.

AliNiazee, M.T. 1994. Insect pest management in hazelnut orchards of North America. Acta Horticulture 351:

543-549.

AliNiazee, M.T. 1998. Ecology and management of hazelnuts. Annual Review of Entomology 43: 359-419.

AliNiazee, M.T. and R.H. Messing. 1995. Filbert Aphid, pp. 123-129. In: J.R. Nechols, L.A. Andres, J.W.

Beardsley, R.D. Goeden, and C.G. Jackson (Eds.), Biological control in the Western United States.

University of California, Division of Agriculture and Natural Resources Publication 3361.

Bureau of Labor Statistics Data. 1998. Consumer Price Index-All Urban Consumers- Portland-Salem, OR.

Internet: http://146.142.4.24/cgi-bin/surveymost?r9

Calkin, J. and G. C. Fisher. 1985. Three years of filbert IPM. In: Abstracts of Reports From the 59" Annual

Western Orchard Pest and Disease Management Conference. Imperial Hotel, Portland, Oregon. January 16-

18, 1985.

Calkin, J., M. T. AliNiazee and G. C. Fisher. 1984. Hazelnut integrated pest management. Proceedings of the

International Congress on Hazelnut. Avellino, Italy, September, 22-24, 1983. pp. 477-484.

Fisher, G. C. 1984. Integrated pest management-Filberts. Proceedings of the Nut Growers Society of Oregon,

Washington and British Columbia 69: 92-95.

Headley, J.C. and M.A. Hoy. 1986. The economics of integrated mite management in almonds. California

Agriculture Jan.-Feb.: 28-30.

Katundu, J.M. and M.T. AliNiazee. 1990. Variable resistance of filbert aphid (Homoptera: Aphididae) to

insecticides in the Willamette Valley, Oregon. Journal of Economic Entomology 83: 41-47.

Messing R. H. and M. T. AliNiazee. 1989. Introduction and establishment of 7rioxys pallidus in Oregon for the

control of the filbert aphid. Entomophaga 34: 153-163.

National Academy of Sciences, 1989. Alternative Agriculture. National Academy Press, Washington, DC, 448

Pp.

Office of Technology Assessment. 1979. Pest Management Series. Volume II. Working Papers. Office of

Technology Assessment, Washington, DC, 169 pp.

Olsen, J.L., G.C. Fisher and J.W. Pscheidt. 2000. Oregon State University Pest Management Guide. Publication

No. EM8328.

J. ENTOMOL. SOC. BRIT. COLUMBIA 97, DECEMBER 2000 17

Pimentel, D., L. McLaughlin, D. Zepp, B. Lakitan, T. Kraus, P. Kleinman, F. Vancini, W.J. Roach, E. Graap,

W.S. Keeton and G. Selig. 1993. Environmental and economic effect of reducing pesticide use in agriculture.

Agriculture, Ecosystems and the Environment 46: 273-288.

Progar, R.A. and M.T. AliNiazee. 1999. Insecticide use patterns in Oregon: past and present. Proceedings of the

Nut Growers Society of Oregon, Washington and British Columbia 84: 43-46.

Rowley, H. K. 1997. Hazelnut tree report. Internet address:http://www.oda.state.or.us/oass/hzltre97.htm

Seavert, C.F. and J.L. Olsen. 1999. Enterprise Budget: Hazelnut, Willamette Valley Region. Oregon State

University Extension Service Publication EM8556.

Trumble, J.T. and B. Alvarado-Rodriguez. 1993. Development and economic evaluation of an IPM program for

fresh market tomato production in Mexico. Agriculture, Ecosystems and the Environment 43: 267-284.

Trumble, J.T., W.G. Carson and K.K. White. 1994. Economic analysis of a Bacillus thuringiensis-based

integrated pest-management program in fresh market tomatoes. Journal of Economic Entomology 87: 1463-

1469.

Trumble, J.T., W.G. Carson and G.S. Kund. 1997. Economics and environmental impact of a sustainable

integrated pest management program in celery. Journal of Economic Entomology 90: 139-146.

White, F.C. and M.E. Wetzstein. 1995. Market effects of cotton integrated pest management. American Journal

of Agricultural Economics 77: 602-612.

J. ENTOMOL. SOC. BRIT. COLUMBIA 97, DECEMBER 2000 19

Enhancement of the activity of a nuclear polyhedrosis virus

by an optical brightener in the eastern hemlock looper,

Lambdina fiscellaria fiscellaria (Lepidoptera: Geometridae)

S. Y. LI

NATURAL RESOURCES CANADA, CANADIAN FOREST SERVICE, ATLANTIC

FORESTRY CENTRE, P. O. BOX 960, CORNER BROOK, NEWFOUNDLAND, CANADA

A2H 6J3

I.S. OTVOS

NATURAL RESOURCES CANADA, CANADIAN FOREST SERVICE, PACIFIC

FORESTRY CENTRE, 506 WEST BURNSIDE ROAD, VICTORIA, BRITISH COLUMBIA,

CANADA V8Z 1M5

ABSTRACT

The pathogenicity of a nuclear polyhedrosis virus originally isolated from Lambdina

fiscellaria lugubrosa (Hulst) was compared between treatments with and without the

optical brightener Blankophor P167 against L. f fiscellaria (Guenée) in the laboratory.

The brightener significantly enhanced viral activity by 7.5-fold in terms of LDso, and

by 22.9-fold in terms of LDo;. With the addition of the brightener, the virus killed L. f

fiscellaria larvae 1.5- to 1.8-fold faster than without the brightener.

Key words: nuclear polyhedrosis virus, Lambdina fiscellaria fiscellaria, optical

brightener, virus enhancement

INTRODUCTION

The eastern hemlock looper (EHL), Lambdina fiscellaria fiscellaria (Guenée)

(Lepidoptera: Geometridae), is one of the most destructive defoliators of balsam fir, Abies

balsamea (L.), in eastern Canada (Hudak and Raske 1995). Outbreaks of EHL occur

periodically, and each outbreak lasts several years before populations collapse abruptly.

Viral diseases are suspected to play an important role in the collapse of outbreaks of this

insect, although other field studies suggested that fungi are important mortality factors

(Otvos et al. 1973). Cunningham (1970) isolated a multicapsid nuclear polyhedrosis virus

(LAMNPV) from an outbreak population of EHL. Previous laboratory studies revealed that

larvae of EHL were susceptible to LA/MNPV and to other NPV viruses originally isolated

from the western hemlock looper, L. f /ugubrosa (Hulst) or from the western oak looper, L.

f somniaria (Hulst) (Cunningham 1970). However, no satisfactory results have been

obtained in a field spray trial with LAMNPV against EHL (Cunningham and Kaupp 1995).

Optical brighteners act as whiteners, ultraviolet absorbers and protectants, thus they are

widely used in detergent, paper, plastics, and organic coatings industries (Villaume 1958).

Recently, optical brighteners were shown not only to protect insect viruses from the

denaturing effect of UV radiation (Shapiro 1992), but also to enhance biological activity of

several insect viruses against their respective hosts (Shapiro and Robertson 1992; Li and

Otvos 1999a). Enhancement of viral activity by optical brighteners varied greatly from one

virus-host system to another. Whether optical brighteners could enhance NPV activity

against EHL was not determined previously. Here we report the results of laboratory

20 J. ENTOMOL. SOC. BRIT. COLUMBIA 97, DECEMBER 2000

experiments on EHL larvae exposed to various dosages of a nuclear polyhedrosis virus

with and without Blankphor P167, an optical brightener that significantly enhanced activity

of another NPV against the western spruce budworm (Li and Otvos 1999a). The objectives

of this study were to determine 1) effects of the brightener on larval mortality of EHL

(enhancement of viral activity), 2) effects of the brightener on the time-to-death of larvae

killed by the virus.

MATERIALS AND METHODS

Experimental insects. Larvae were obtained from a laboratory colony of L. f

fiscellaria. The colony originated from larvae collected from the field in Québec and

Newfoundland, and had been reared in the laboratory for one generation on artificial diet

and natural foliage before the experiment (Li and Otvos 1999b). Following a 3 month

diapause in darkness at 2.0 + 0.5°C and 100% R.H., the eggs of EHL were moved to a

rearing room under conditions of 20 + 1°C, 55-60% R.H., and a photoperiod of 16:8 (L:D)

h. Eggs were checked twice a day, and newly hatched larvae were transferred into 20-ml

creamer cups (five larvae/cup) that contained a modified spruce budworm artificial diet

without formalin (Robertson 1979). Larvae in the cups with diet were kept at the above

rearing conditions for about 2 weeks before being transferred onto flushing young (1 m-

high) grand fir trees, Abies grandis (Dougl. ex D. Don) housed in a cage (0.8 x 1.0 x 1.5

m) at a rate of about 100 larvae per tree. The detailed larval rearing techniques were

reported by Li and Otvos (1999b). Fourth-instar larvae, < 24-h-old, were removed from

the trees in the cages, and placed in 24-well tissue culture plates (one larva/well) without

food for 16-20 h before bioassays. Fourth-instar larvae were chosen for the tests because

they are large enough to consume a virus-contaminated artificial diet pellet within 24 h (see

below), and because their susceptibility to virus is not significantly different from those of

younger larvae (Cunningham 1970).

Virus inocula. A multicapsid nuclear polyhedrosis virus originally isolated from L.f

lugubrosa (LAIMNPV) was found to infect EHL larvae (Cunningham 1970). LAIMNPV was

purified by repeated centrifugation (3,000 - 8,000 rpm for 30 min each time at 15 °C) and

resuspension in sterile distilled water. Stock suspensions of LAMNPV were quantified by

counting polyhedral inclusion bodies (PIB) using Wigley’s method (1980) and stored at 2

°C before use. Inocula were diluted in distilled water or in the final concentration of 1%

(wt/wt) of optical brightener Blankophor P167 (Bayer Corp., Pittsburgh, PA) to the desired

concentrations in the bioassays. The 1% concentration was tested because it was an

optimal concentration for enhancing viral activity (Argauer and Shapiro 1997; Li and

Otvos 1999a).

Bioassays. Five viral concentrations from 39 to 5,000 PIB/ul and six from 39 to

10,000 PIB/ul were used in the treatments with and without brightener, respectively. In

addition, one control (distilled water alone) was made for the treatment without brightener,

and 1% Blankophor P167 without virus was used as a control for the treatment with

brightener. One ul of each viral dilution or control was applied onto a small pellet [4.4 mg

+ 0.1 (SE), n = 20] of artificial diet inside each well of a 24-well tissue culture plate. The

diet pellets were large enough to fully absorb | ul of liquid and allowed larvae to ingest

known amount of virus. Immediately after the virus was added to the pellets, one fourth-

instar larva, fasted for 16-20 h, was placed into each of the wells. Larvae were confined in

the wells by covering the tissue culture plates with lids and were allowed to feed on the

treated diet plug for 24 h under conditions of darkness, 20 + 1°C, and 55-60% R.H.

Preliminary tests indicated that higher proportion of larvae consumed the entire diet plug

within 24 h in the dark than in the light. Twenty-four larvae were tested for each replicate,

J. ENTOMOL. SOC. BRIT. COLUMBIA 97, DECEMBER 2000 21

and three replicates were made for each dilution or control. Those larvae that consumed

the entire pellet of diet were transferred to untreated fresh one-year-old foliage of grand fir

in a 170-ml fluted food cup (Sweetheart Cup Co. Inc., Chicago, IL) (five per cup) and

placed at 25 + 1°C, 55-60% RH, and 16:8 (L:D) h. ere that did not consume the entire

pellet of diet were discarded.

Data analysis. Mortality was checked twice per week, and foliage changes Tests

were terminated 31 d after treatment, by which time larvae had either died or pupated. The

cumulative mortality by 31 d was analyzed using probit analysis (LeOra Software 1994) to

estimate the lethal doses of LDs) and LDy;. Differences in LDs) or LDo; between treatments

with and without Blankophor P167 were compared for significance (P < 0.05) using the

lethal-dose ratio test (Robertson and Preisler 1992).

To determine the effect of the optical brightener on the time to death (LTs 9 or

LT»5;), the data on larval mortality over time was analyzed with a complementary log-log

model (Preisler and Robertson 1989; Robertson and Preisler 1992). The LTs59 and LT9; for

both treatments were estimated at the concentration of 5,000 PIB/ul of LAMNPV. The

lethal-dose ratio test (Robertson and Preisler 1992) was used to determine significant

differences (P < 0.05) in LTs) or LT»; between the two treatments.

RESULTS AND DISCUSSION

Effects of the optical brightener on larval mortality of EHL. Larval mortality was

low in both controls [4.2% (n = 72) for distilled water alone, and 4.3% (n = 70) for 1%

optical brightener alone], indicating that the larvae tested were healthy and that optical

brightener was not toxic to EHL larvae. The LDs5) and LDos for the treatment with

LAIMNPV plus 1% Blankophor P167 were significantly (P < 0.05) lower than those for the

treatment with LAIMNPV alone (Table 1), indicating that the brightener enhanced

LAIMNPY activity against L. f fiscellaria. About 174 PIB per larva were required to kill

50% of the test larvae when virus was used alone, while only 23 PIB per larva were needed

to kill the same percentage of larvae when 1% brightener was added to the LAIMNPV

suspension. To increase larval mortality from 50 to 95%, 42.8 times as much virus was

required for the treatment using L/[MNPV alone (i.e., an increase from 173.9 to 7443.3

PIB per larva). In contrast, only 14.1 times as much virus was needed for the treatment

with LAIMNPV plus 1% brightener to increase larval mortality from 50 to 95% (i.e., an

increase from 23.1 to 326.6 PIB per larva). In terms of LDso, the addition of the brightener

enhanced viral activity by 7.5 times. In terms of LDos, the brightener enhanced LAMNPV

activity by 22.9 times (Table 1). The 7.5- to 22.9-fold enhancement of viral activity in this

study was much lower than the 90- to 1,500-fold increases previously reported when

stilbene brighteners were added to LAMNPV against Lymantria dispar (L.) (Shapiro and

Robertson 1992; Argauer and Shapiro 1997), but was comparable to the 1.8- to 13-fold

increase in the virus-host system of C(AMNPV and Choristoneura occidentalis Freeman (L1

and Otvos 1999a).

Several bioassays have been developed to study insect viruses in the laboratory. In this

study, we used a bioassay in which the same known amount of virus was consumed by each

test larva. This technique may have some advantages over surface-contamination or

foliage dipping bioassays where larvae ingested unknown amount of virus. With the

addition of optical brighteners to the virus, feeding behavior of the larvae may have

changed and they may not have consumed the same amount of virus. Thus, the bioassay

used in this study may give more reliable results on the effects of optical brighteners on

viral activity.

J. ENTOMOL. SOC. BRIT. COLUMBIA 97, DECEMBER 2000

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BLAST

J. ENTOMOL. SOC. BRIT. COLUMBIA 97, DECEMBER 2000 D3

Effects of the optical brightener on time to death of EHL larvae killed by

LfIMNPV. The times to death (LTs9 or LT9s) for the treatment with L//MNPV plus 1%

Blankophor P167 were significantly (P < 0.05) shorter than those for the treatment with

LAIMNPV alone (Table 2), indicating that larvae died faster when the brightener was added

to LAIMNPV suspensions. At the concentration of 5,000 PIB per larva of LA-AMNPV, 13.6

days were required to kill 50% of the larvae when virus was used alone, while only 9.1

days were needed to kill the same percentage of larvae when 1% brightener was added to

LAIMNPYV suspensions. To increase the lethal time from LTs9 to LT95, 2.3-fold as long was

required for the treatment using L//MNPV alone (i.e., an increase from 13.6 to 31.8 days).

In contrast, only 1.9-fold as long was needed for the treatment with LAMNPV plus 1%

brightener to increase the lethal time from LTs9 to LT95 (i.e., an increase from 9.1 to 17.3

days). In terms of LTso, larvae in the treatment with virus plus brightener died 1.5 times

faster than those in the treatment with virus alone. In terms of LT95, the brightener reduced

the time to death by 1.8 times (Table 2).

The addition of 1% Blankophor P167 not only increased LAIMNPV activity against L. f

fiscellaria, but also hastened larval death in the laboratory. Although several insect viruses

have been operationally used in pest management programs (Cunningham and Kaupp

1995), one of the drawbacks is the slow killing of pests by viruses. In most cases, foliage

protection is the main aim of pest management strategies. Thus, the slow-action of insect

viruses may be a serious disadvantage in pest control programs because severe damage

may occur before pests are killed. Hastening larval death by the addition of optical

brighteners to insect viruses may be of significance in crop protection, because viruses may

kill pests before serious damage occurs. Hastening larval death may also lead to earlier

and greater horizontal transmission of the virus, which enhances the development of

secondary infection and possibly terminates the outbreak of the pest (Otvos ef al. 1989).

More research is needed to test the efficacy of virus with the addition of optical brighteners

in the field under natural conditions.

ACKNOWLEDGEMENTS

We wish to thank the following individuals: A. Van de Raadt (Natural Resources

Canada, Canadian Forest Service, Victoria, British Columbia) for technical assistance, D.

Moranville (Société de protection des foréts contre les insectes et maladies, Québec City,

Québec) and H. Crummey (Newfoundland Department of Forest Resources and Agrifoods)

for supplying EHL larvae and eggs, and W. Bowers (Natural Resources Canada, Canadian

Forest Service, Corner Brook, Newfoundland) for constructive comments on an early draft

of the manuscript. This research was funded by Forest Renewal BC and Natural Resources

Canada.

REFERENCES

Argauer, R. and M. Shapiro. 1997. Fluorescence and relative activities of stilbene optical brighteners as

enhancers for the gypsy moth (Lepidoptera: Lymantriidae) baculovirus. Journal of Economic

Entomology 90: 416-420.

Cunningham, J.C. 1970. Pathogenicity tests of nuclear polyhedrosis viruses infecting the eastern hemlock

looper, Lambdina fiscellaria fiscellaria (Lepidoptera: Geometridae). The Canadian Entomologist 102:

1534-1539.

Cunningham, J.C. and W.J. Kaupp. 1995. Insect viruses. pp. 327-340. In: J.A. Armstrong and W.G.H. Ives

(Eds.), Forest Insect Pests in Canada. Natural Resources Canada, Canadian Forest Service, Ottawa.

Hudak, J. and A.G. Raske. 1995. Forest insect pests in the Newfoundland and Labrador region. pp. 1-9. In:

J.A. Armstrong and W.G.H. Ives (Eds.), Forest Insect Pests in Canada. Natural Resources Canada,

Canadian Forest Service, Ottawa.

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LeOra Software. 1994. Polo-PC. A User's Guide to Probit or Logit Analysis. LeOra Software, Berkeley. 28

Pp.

Li, S.Y. and I.S. Otvos. 1999a. Optical brighteners enhance activity of a nuclear polyhedrosis virus against

western spruce budworm (Lepidoptera: Tortricidae). Journal of Economic Entomology 92: 335-339.

Li, S.Y. and I.S. Otvos. 1999b. Laboratory rearing of the eastern hemlock looper (Lepidoptera: Geometridae)

on artificial diet and grand fir foliage. Journal of the Entomological Society of British Columbia 96: 25-

ay.

Otvos, I.S., J.C. Cunningham and W.J. Kaupp. 1989. Aerial application of two baculoviruses against the

western spruce budworm, Choristoneura occidentalis Freeman (Lepidoptera: Tortricidae), in British

Columbia. The Canadian Entomologist 121: 209-217.

Otvos, I.S., D.M. MacLeod and D. Tyrrell. 1973. Two species of Entomophthora pathogenic to the eastern

hemlock looper (Lepidoptera: Geometridae) in Newfoundland. The Canadian Entomologist 105: 1435-

1441.

Preisler, H.K. and J.L. Robertson. 1989. Analysis of time-dose-mortality data. Journal of Economic

‘ Entomology 82: 1534-1542.

Robertson, J.L. 1979. Rearing the Western Spruce Budworm. Canada -- United States Spruce Budworms

Program, USDA Forest Service, Washington, D.C. 18 pp.

Robertson, J.L. and H.K. Preisler. 1992. Pesticide Bioassays with Arthropods. CRC Press, Boca Raton,

Florida. 127 pp.

Shapiro, M. 1992. Use of optical brighteners as radiation protectants for gypsy moth (Lepidoptera:

Lymantriidae) nuclear polyhedrosis virus. Journal of Economic Entomology 85: 1682-1686.

Shapiro, M. and J.L Robertson. 1992. Enhancement of gypsy moth (Lepidoptera: Lymantriidae) baculovirus

activity by optical brighteners. Journal of Economic Entomology 85: 1120-1124.

Villaume, R.L. 1958. Optical bleaches in soaps and detergents. Journal of American Oil Chemistry Society.

35: 558-566.

Wigley, P.J. 1980. Counting micro-organisms. pp. 29-35. In: J. Kalmakoff and J.F. Longworth (Eds.),

Microbial Control of Pests. New Zealand Department of Science and Industrial Research Bulletin 228.

Wellington, New Zealand.

J. ENTOMOL. SOC. BRIT. COLUMBIA 97, DECEMBER 2000 25

The bee fauna (Hymenoptera: Apoidea) of

southeastern Washington

D.F. MAYER, E.R. MILICZKY, B.F. FINNIGAN, C.A. JOHANSEN

WASHINGTON STATE UNIVERSITY, IAREC, 24106 NORTH BUNN ROAD

PROSSER, WA 99350

ABSTRACT

A survey of the species composition, distribution, and host plants of bees (Hymenoptera:

Apoidea) was conducted in the Snake River area, the Colton area, and the Moscow

Mountain area of southeastern Washington. Nineteen genera and 100 species occurred in

the three areas. The number of species found in each family were: 1 Colletidae; 11

Halictidae; 31 Megachilidae; 27 Adrenidae; 15 Anthophoridae and 15 Apidae. Location

and flowers visited are listed for each species.

Key words: bees, Hymenoptera, Apoidea, bee fauna, Washington State

INTRODUCTION

There are no published faunal studies of bees in Washington despite their importance in

pollination and their high priority with respect to conservation of biodiversity (Williams e¢ al.

1993). Most information on the native bee species occurring in Washington is difficult to

access because it is In various systematic works dealing with particular taxa (families, genera).

Bee studies in Washington have, in many cases, been concerned with the role of bees as

pollinators of commercial crops. Menke (1952) listed a number of genera of Apoidea

associated with apple (Malus =< domestica Borkh). The alkali bee (Nomia melanderi

Cockerell) and the alfalfa leafcutter bee (Megachile rotundata (Fabricius)) are important

pollinators managed by alfalfa growers and have been studied extensively in Washington

(Menke 1954; Johansen ef a/. 1978; Eves et al. 1980). Bumblebees have been studied for

pollination of red clover seed (Johansen 1960; Dade and Johansen 1962) and cranberries

(Johansen 1967; Macfarlane et al. 1994). Other studies of native bees in Washington examined

and related behaviors of Anthophora urbana urbana Cresson (Mayer and Johansen 1976),

Andrena vicina Smith (Miliczky and Osgood 1995), and Melissodes microsticta Cockerell

(Miliczky 2000).

Bee diversity on the Columbia Plateau is expected to be high. Stephen ef al. (1969)

estimated that 879 species occurred in northwestern North America, and Washington, Oregon,

and Idaho formed the core of this region. Bees are thought to reach their greatest diversity in

number of species in warm temperate, xeric regions (Linsley 1958; Michener 1979).

Mountainous areas of moderate rainfall, varied floras, and soils suitable for ground nesting

forms also support rich bee faunas (Linsley 1958). Washington offers large expanses of both

types of habitat, especially the part of the state east of the Cascade Mountains. Here we

document the species composition, distribution, and host plants of the native bees of

southeastern Washington in the first comprehensive study of Washington’s bee fauna.

MATERIALS AND METHODS

We sampled pollinator communities in three ecological regions of southeastern

Washington: 1) The Snake River area is 15.3 ha located 21-26 km southwest of Pullman, WA

26 J. ENTOMOL. SOC. BRIT. COLUMBIA 97, DECEMBER 2000

along the Snake River Road 5-8 km below the Wawawai railroad siding (T.36N-R.43E of

quadrangle 57). The elevation ranged from 183-207 m and the area is Upper Sonoran Life

Zone (St. John 1937). 2) The Colton area is a 11.3 ha original Palouse prairie vegetation area

21 km south of Pullman, WA (T.37N-R45E of quadrangle 57). The elevation ranged from

808-853 m and the area is Arid Transition Life Zone (St. John 1937). 3) Moscow Mountain is

about 24 km northeast of Pullman, WA (T.39N-R5W of quadrangle 58) The sample area was

on a 22.6 ha slope at the 1450 m level in the Canadian Zone (St. John 1937).

In each area we observed and collected bees from 15 June 1962 to 22 October 1962 and

from 30 March 1963 to 20 June 1963. One collector made weekly trips to each site for a

minimum of 8 h per day during the spring, summer and fall; a total of 34 collecting trips. The

majority of collecting and observations were from 0730 h to 1730 h though we did

occasionally collect and observe from dawn to dusk. Whenever possible, bee species

identifiable in the field (Bombus, and some Andrea and Anthophora) were released to maintain

the populations. Unidentified flowers were first given a site number. The bees and time of

visitation were recorded in reference to this number and then plant specimens were collected

for identification.

We used the direct searching method and insect nets to capture bees on flowers or in flight.

We used the taxonomic system of Michener ef.a/. (1994) and used LaBerge (1956a, 1956b,

1961), Stephen (1954), Stephen (1957), Stephen er al. (1969) and Thorp et al. (1983) to

identify collected specimens. Most of the Andrena, Mellissodes, Diadasia and Colletes were

determined by Wallace E. LaBerge. Voucher specimens for 89 species were deposited at the

Insect Museum at Washington State University, Pullman, WA.

Plant names are those used by Hitchcock (1955), except for the majority of Compositae

which are from St. John (1937). Flower specimens were compared with determined material in

the Herbarium at Washington State University.

RESULTS

Six-hundred-and-seven bees were collected. The diversity of bee species was greatest at

the Snake River site (18 genera, 64 species) followed by Moscow Mountain (14 genera, 54

species) and the Colton site (14 genera, 37 species) (Table 1). Six families of bees were found

(Colletidae, Halictidae, Megachilidae, Andrenidae, Anthophoride and Apidae). Nineteen

genera and 100 species occurred in the three areas. One genus of Colletidae, 4 genera of

Halictidae, 5 genera of Megachilidae, 2 genera of Andrenidae, 6 genera of Anthophoride and 2

genera of Apidae (the honey bee (Apis melifera L.) was not included) occurred in the study

areas. One species of Colletidae, 11 species of Halictidae, 31 species of Megachilidae, 27

species of Andrenidae, 15 species of Anthophoridae and 15 species of Apidae occurred in the

Study areas.

Twenty-one families of plants (80 species) were identified as sources of nectar or pollen

(usually both) for the visiting Apoidea (Table 1). The effect of elevation on host plant

distrubtion and phenology was reflected in the distrubtion and capture dates of bee species at

all three locations. Osmia nanula Cockerell:was found in the middle of May at the lowest

elevation but one month later at the mountain area. Halictus ligatus Say, H. tripartitus

Cockerell, Andrena prunorum Cockerell and A. opaciventris Cockerell were also captured

later in the season at higher elevations. This same phenomenon was also shown by the six most

common Bombus species. Queens of all six species were observed by mid-April at the Snake

River area, about 2 weeks later at the Colton area, except for B. occidentalis Greene, and at the

Moscow mountain area about 4 weeks later.

J. ENTOMOL. SOC. BRIT. COLUMBIA 97, DECEMBER 2000 27

Table 1

List of bee species collected from three specific areas in southeastern Washington.

S = Snake River area, C = Colton area, M = Moscow Mtn. area.

Cockerell

Family and Species Area Flowers visited

COLLETIDAE (1 sp.)

Colletinae

Colletes californicus Provancher M Mertensia paniculata

HALICTIDAE (11 spp.)

Halictinae

Agapostemon cockerelli Crawford M Rudbeckia occidentalis, Cirsium arvense,

Cirsium vulgare

Agapostemon texanus Cresson sc Convolvulus sp., Medicago sativa,

Helianthus annus, Haplopappus sp.

Agapostemon virescens (Fabricius) S,M,C C. vulgare, Vicia sp., Helianthus annus,

Rosa sp., Gaillardia aristata, Epilobium

angustifolium, Gentiana calycosa,

Compositae

Halictus farinosus Smith 5 Lomatium spp., Malus x domestica,

Brassica campestris, Sisvmbrium

attissimum, Helianthus annus, Solidago sp.

Halictus ligatus Say S, M,C Helianthus annus, Cirsium arvense,

Solidago sp., Hapiopappus sp.

Halictus rubicundus (Christ) S, M Lomatium spp., Trifolium repens,

Taraxacum officinate, Ranunculus sp..

Cirsium arvense

Halictus tripartitus Cockerell SVL © Lomatium spp., Rosa sp., Compositae,

Solidago sp., Trifolium repens, Ranunculus

sp., Taraxacum officinale, Collinsia

parviflora, Cirsium arvense

Lasioglossum spp. sen. Ss. Ss, M,C Lomatium spp., Malus x domestica,

Balsamorhiza sagittata, Potentilla

gracilis, Helianthella uniflora,

Trifolium repens

Lasioglossum spp. (Dialictus) 5 Lomatium spp.,Malus « domestica,

Gaillardia aristata

Lasioglossum spp. (Evylaeus) S Lomatium spp., Prunus avium, Malus ~

domestica, Prunus virginiana, Taraxacum

officinale

Rophitinae

Dufourea sp. M Phacelia heterophylla

MEGACHILIDAE (31 spp.)

Megachilinae

Anthidium emarginatum (Say) S Phacelia heterophylla

Anthidium utahense Swenk S Vicia villosa

Hoplitis albifrons argentifrons M Lupinus polyphyllus, Phacelia heterophylla

(Cresson)

Hoplitis fulgida fulgida (Cresson) M Ranunculus sp., Delphinium nuttalliana,

Physocarpus malvaceus, Phacelia

heterophylla

Hoplitis hypocrita (Cockerell) S,M Balsamorhiza sagitatta, Lomatium spp..

Penstemon attenuatus

Megachile brevis Say S Solidago sp.

Megachile gemula Cresson M Physocarpus malvaceus

Megachile melanophaea calogaster M none

Megachile parallela Smith

Megachile perihirta Cockerell

Megachile pugnata Say

Osmia atrocyanea atrocyanea

Cockerell

Osmia brevis Cresson

Osmia bruneri Cockerell or cobaltina

Cresson

Osmia californica Cresson

Osmia calla Cockerell

Osmia coloradensis Cresson

Osmia juxta juxta Cresson

Osmia kincaidii Cockerell

Osmia lignaria Say

Osmia montana Cresson

Osmia nanula Cockerell

Osmia nr. nanula Cockerell

Osmia nemoris Sandhouse

Osmia nifoata Cockerell

Osmia nigrifrons Cresson

Osmia pentstemonis Cockerell

Osmia pikei Cockerell

Osmia subaustralis Cockerell

Stelis nr. foederalis Smith

Stelis subcaerulea Cresson

ANDRENIDAE (27 spp.)

Andrenina

Andrena amphibola ( Viereck)

Andrena angustitarsata Viereck

Andrena auricoma (Smith)

Andrena caerulea Smith

Andrena candida Smith

Andrena chlorogaster Viereck

Andrena crataegi Robertson

Andrena cressonii Robertson

Andrena helianthi Robertson

Andrena hemileuca Viereck

Andrena merriami Cockerell

Andrena microchlora Cockerell

Andrena miserabilis Cresson

Andrena nigrocaerulea Cockerell

Andrena nivalis Smith

Se eae

S26

PEZPPPNREZEPYEY

J, ENTOMOL. SOC. BRIT. COLUMBIA 97, DECEMBER 2000

Helianthus annus

Vicia villosa, Xanthium sp., Compositae,

Solidago sp., Gaillardia aristata, Senecio

serra, Cirsium vulgare, Cirsium arvense

Erigeron speciosus

Malus x domestica, Balsamorhiza

sagittata, Lupinus polyphyllus, Vicia

villosa

Vicia villosa, Trifolium repens, Phacelia

heterophylla

Penstemon lanatum

Lomatium spp., Ribes aureum,

Balsamorhiza sagittata, Gaillardia aristata

Vicia villosa

Trifolium repens, Arnica cordifolia

Epilobium angustifolium

Phacelia heterophylla, Collinsia parviflora

Pyrus scopulina, Arnica cordifolia,

Phacelia heterophylla

Rosa sp., Gaillardia aristata

Geranium viscosissimum, Ranunculus sp.

Vicia villosa, Trifolium repens

Balsamorhiza sagittata, Arnica cordifolia

Pyrus scopulina

Balsamorhiza sagittata, Vicia villosa

Penstemon albertinus

Balsamorhiza sagittata

Gaillardia aristata

none

Eriophyllum lanatum, Achillea millefolium

Agastache urticifolia

Lomatium spp.,Malus < domestica, Prunus

virginina, Pyrus scopulina, Rosa sp..,

Ranunculus sp., Physocarpus malvaceus,

Rubus parviflorus

Potentilla sp., Achillea millefolium,

Physocarpus malvaceus

Ranunculus sp., Prunus virginiana

Lomatium spp.,Prunus avium,

Balsamorhiza sagittata

Physcarpus malvaceus, Potentilla sp.

Physcarpus malvaceus

Lomatium spp., Balsamorhiza sagitata,

Rosa sp., Prunus virginiana, Geranium

viscosissimum

Helianthus annus, Solidago canadensis

Pyrus scopulina

Lomatium spp., Prunus avium

Lomatium spp., Malus x domestica, Ribes

aureum

Physcarpus malvaceus

Geranium viscosissimum

Physcarpus malvaceus

J. ENTOMOL. SOC. BRIT. COLUMBIA 97, DECEMBER 2000

Andrena pallidifovea (Viereck)

Andrena perarmata Cockerell

Andrena pertristis carliniformis

Viereck & Cockerell

Andrena prunorum Cockerell

Andrena subsaustralis Cockerell

Anarena subtilis Smith

Andrena topozana Cockerell

Andrena trizonata Ashmead

Andrena vicina Smith

'Andrena sp. E new sp.

Panurginae

Perdita lingualis Cockerell

Perdita wyomingensis sculleni

Timberlake

ANTHOPHORIDAE (15 spp.)

Anthophorinae

Anthophora bomboides Kirby

Anthophora pacifica Cresson

Anthophora ursina Cresson

Diadasia enavata Cresson

Diadasia nigrifrons (Cresson)

Habropoda cineraria (Smith)

Melissodes agilis Cresson

Mellisodes lupina Cresson

Melissodes metenua Cockerell

Mellisodes rivalis Cresson

Mellisodes robustior Cockerell

Synhalonia actuosa (Cresson)

Synhalonia edwardsii (Cresson)

Synhalonia frater (Cresson)

Xylocopinae

Ceratina acantha Provancher

APIDAE (15 spp.)

Bombinae

Bombus appositus Cresson

Bombus bifarius Cresson

ovis €

SMEG

Eriophyllum lanatum

Lomatium spp.

Lomatium spp., Sisymbrium attissimum,

Philadelphus lewisii, Holodiscus discolor,

Geranium. viscosissimum, Physcarpus

malvaceus

Balsamorhiza sagittata

Rosa sp.

Cirsium arvense

Phycarpus malvaceus

Phycarpus malvaceus, Rosa sp., Geranium

viscosissimum, Holodiscus discolor, Rubus

parviflorus

Balsamorhiza sagittata, Lomatium spp.

Rosa sp., Geranium viscosissimum,

Helianthus annus

Holodiscus discolor, Achillea millefolium

none

Lomatium spp., Prunus armeniaca, Malus

x domestica, Syringa sp., Balsamorhiza

sagitatta, Ribes aureum

Vicia villosa

Helianthus annus

Sidalcea oregana

Physcarpus armeniaca, Malus =< domestica,

Rosa sp., Ribes aureum

Helianthus annus, Gaillardia aristata

Helianthus annus

Haplopappus liatriformis

_ Cirsium vulgare

Helianthus annus

Balsamorhiza sagitatta, Malus ~

domestica, Prunus virginiana, Lupinus sp..,

Vicia villosa

Vicia villosa, Lupinus polyphyllus,

Dipsacus sylvestris

Balsamorhiza sagitatta, Malus ~

domestica, Syringa sp., Trifolium repens,

Penstemon attenuatus, Brodiaea douglasii

Lomatium spp., Rosa sp., Penstemon

triphyllus, Eriophyllum lanatum, Helianthus

annus, Geranium viscosissimum, Cirsium

lanceolatum

Phacelia sp., Balsamorhiza sagittata, Vicia

villosa, Agastache urticifolia,

Brodiaea douglasii

Anaphalis margaritacea, Epilobium

Bombus californicus F. Smith

Bombus centralis Cresson

Bombus fervidus (Fabricius)

Bombus flavirons Cresson

Bombus griseocollis (Degeer)

Bombus mixtus Cresson

Bombus nevadensis Cresson

Bombus occidentalis Greene

Bombus rufocinctus Cresson

Bombus vagans Smith

Psithyrus insularis (F. Smith)

SVG

S, M,-C

5; M,C

S,M

@-=

J. ENTOMOL. SOC. BRIT. COLUMBIA 97, DECEMBER 2000

angustifolium, Rudbeckia occidentalis,

Collinsia parviflora, Cirsium arvense,

Phacelia spp., Sisyrinchium albus, Lupinus

polyphyllus, Vicia villosa, Penstemon spp.

Vicia villosa, Sisyrincyhium albus

Epilobium angustifolium, Rosa sp., Rubus

parviflorus, Malus x domestica, Geranium

viscosissimum, Anaphalis margaritacea,

Rudbeckia occidentalis, Collinsia

parviflora, Sisyrinchium albus,

Balsamorhiza sagittata, Lupinus

polyphyllus, Trifolium repens, Mertensia

paniculata, Dipsacus sylvestris, Vicia

villosa, Agastache urticifolia, Penstemon

spp., Brodiaea douglasii

Epilobium angustifolium, Rosa sp., Malus

x domestica, Geranium viscosissimum,

Anaphalis margaritacea, Rudbeckia

occidentalis, Sisyrinchium albus, Medicgo

sativa, Balsamorhiza sagittata, Cirsium

lanceolatum, Lupinus polyphyllus,

Dipsacus sylvestris, Vicia villosa,

Agastache urticifolia, Brodiaea douglasii

Epilobium angustifolium, Cirsium arvense,

Sisyrinchium albus, Helianthus annus,

Dipsacus sylvestris, Vicia villosa,

Agastache urticifolia, Penstemon spp..,

Castilleja sp.

Epilobium angustifolium, Rosa sp.,

Solidago sp., Phacelia sp., Sisyrinchium

albus, Medicgo sativa, Balsamorhiza

sagittata, Helianthus annus, Lupinus

polyphyllus, Vicia villosa, Penstemon spp.

Epilobium angustifolium, Rudbeckia

occidentalis, Collinsia parviflora, Phacelia

sp., Sisyrinchium albus, Lupinus

polyphyllus, Arnica cordifolia, Mertensia

paniculata

Malus x domestica, Solidago sp., Phacelia

sp., Medicgo sativa, Balsamorhiza

sagittata, Cirsium lanceolatum, Trifoluim.

repens, Dipsacus sylvestris, Vicia villosa,

Agastache urticifolia, Astragulus sp.,

Penstemon spp., Brodiaea douglasii

Epilobium angustifolium, Rosa sp., Rubus

parviflorus, Malus x domestica, Phacelia sp..,

Sisyrinchium albus, Medicgo _ sativa,

Balsqamorhiza sagittata, Cirsium

lanceolatum, Lupinus polyphyllus, Trifoluim

repens, Aconitium columbianum, Vicia

villosa, Penstemon sp., Brodiaea douglasii

Epilobium angustifolium, Geranium

viscosissimum, Phacelia sp., Sisyrinchium

albus, Brodiaea douglasii

Sisyrinchium albus

Epilobium angustifolium, Dipsacus

J. ENTOMOL. SOC. BRIT. COLUMBIA 97, DECEMBER 2000 3]

sylvestris, Agastache urticifolia

Psithyrus suckleyi (Greene) S, M Epilobium angustifolium, Sisyrinchium

albus, Agastache urticifolia, Brodiaea

douglasii, Senecio viscosissimum

Psithyrus variabilis (Cresson) M Epilobium angustifolium

' Species E in the collection of Dan Mayer; yet to be described.

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Washington state (Hymenoptera:Apidae). Pan-Pacific Entomologist. 76: 184-196.

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Washington, with new parasite records for A. (M.) regularis Malloch and a review of Me/andrena biology.

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St. John, H. 1937. Flora of southeastern Washington and of adjacent Idaho. Washington State College Student's

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Stephen, W.P. 1954. A revision of the bee genus Co//etes in America north of Mexico (Hymenoptera:Colletidae).

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Stephen, W.P. 1957. Bumble bees of western America. Oregon Agricultural Experiment Station Technical

Bulletin 40. 163 pp.

Stephen, W.P., G.E. Bohart and P.F. Toricho. 1969. The biology and external morphology of bees with a

synopsis of the genera of northwestern America. Agricultural Experiment Station, Oregon State University,

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Thorp, R.W., D.S. Horning and L.L. Dunning. 1983. Bumble bees and cuckoo bumble bees of California

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23.99 pp: ;

Williams, P.H., R.I. Vane-Wright and C.J. Humphries. 1993. Measuring biodiversity for choosing conservation

areas. pp. 309-328. In: J. LaSalle and I.D. Gauld (Eds.) Hymenoptera and Biodiversity. C.A.B.

International, Wallingford, U.K.

Points

eas

ih wh)

? Hi

i =

188

J. ENTOMOL. SOC. BRIT. COLUMBIA 97, DECEMBER 2000 33

A comparison of traps and trap trees for capturing

Douglas-fir beetle, Dendroctonus pseudotsugae

(Coleoptera: Scolytidae)

KEVIN J. DODDS, DARRELL W. ROSS

DEPARTMENT OF FOREST SCIENCE, OREGON STATE UNIVERSITY,

CORVALLIS OR 97331

GARY E. DATERMAN

USDA FOREST SERVICE, PACIFIC NORTHWEST RESEARCH STATION, 3200

JEFFERSON WAY, CORVALLIS, OR 97331

ABSTRACT

We compared pheromone-baited traps and trap trees for managing Douglas-fir beetle

(DFB), Dendroctonus pseudotsugae Hopkins populations. Pheromone-baited traps

caught significantly more DFB than did trap trees. More male DFB were caught in

pheromone-baited traps than in trap trees, while significantly higher numbers of

females were caught in the trap trees. Additional benefits of pheromone-baited traps

include, easy deployment, less mortality of some beneficial insects, and low cost.

Key words: Dendroctonus pseudotsugae, Scolytidae, pheromones, trapping, trap trees

INTRODUCTION

The Douglas-fir beetle (DFB), Dendroctonus pseudotsugae Hopkins (Coleoptera:

Scolytidae) is found throughout the range of Douglas-fir, Pseudotsugae menziesii (Mirbel).

Although endemic populations of DFB usually inhabit dead, dying, downed, or injured

trees, epidemic populations may also attack and kill large numbers of apparently healthy

trees. Tree mortality caused by these beetles can lead to severe economic losses and

interfere with management objectives in the infested area.

Pheromones of DFB are well known (Pitman and Vité 1970; Kinzer et al. 1971;

Furniss et al. 1972; Rudinsky et a/. 1974; Libbey et a/. 1983) and several have been

implemented in management strategies. Aerial application of the DFB anti-aggregation

pheromone, 3-methylcyclohex-2-en-l-one (MCH), can effectively prevent the infestation

of windthrown trees (McGregor ef al. 1984). Strategies incorporating pheromone-baited

traps and MCH (Ross and Daterman 1994), or MCH alone (Ross and Daterman 199Sa),

have significantly reduced DFB infestations in live trees in high-risk stands. Aggregation

pheromones have been used to create trap trees in areas where DFB population levels are

high (Knopf and Pitman 1972; Pitman 1973; Ringold ef a/. 1975). Trap trees concentrate

DFB in selected trees that are subsequently harvested, thereby removing beetles from the

local population. Aggregation and anti-aggregation pheromones can be used to selectively

create tree snags, an important wildlife habitat component (Ross and Niwa 1997).

Pheromone-baited traps may be an alternative to trap trees in some situations (Ross and

Daterman 1995b). While trap trees have been used for a number of years in operational

programs (Patterson 1992), pheromone-baited traps have been used only to a limited extent

by managers. This study was designed to compare the efficacy of trap trees and

pheromone-baited traps in managing DFB.

34 J. ENTOMOL. SOC. BRIT. COLUMBIA 97, DECEMBER 2000

MATERIALS AND METHODS

Field research was conducted in the Nezperce National Forest in central Idaho. The

study area was a mixed-conifer stand composed primarily of Douglas-fir, with ponderosa

pine (Pinus ponderosa Laws.) and grand fir (Abies grandis Lindl.) present at lower

densities. Elevation of the study area ranged from 1524 to 1584 m and it was bisected by a

forest road, with a recent clearcut on one side and a mature mixed-conifer stand on the

other.

On 28 April 1997, before the onset of DFB flight, pheromone-baited traps were placed

in the clearcut area adjacent to the Douglas-fir stand. Seven 16-unit multiple funnel traps

(Lindgren 1983) were baited with 400 mg of frontalin (1,5-dimethyl-6,8-dioxabicyclo

[3.2.1] octane) and 200 mg of seudenol (3-methylcyclohex-2-en-1-ol) in polyvinylchloride

(PVC) formulations, and 15 ml of ethanol in a plastic pouch formulation. Release rates and

chemical descriptions can be found in Ross and Daterman (1997). Traps were positioned

in a line approximately 75 m apart. A piece of dichlorvos-impregnated plastic was added

to each collection cup to kill captured insects. Captured insects were collected weekly

from 15 May to 26 August. Samples were sorted to remove DFB and three primary bark

beetle predators, Thanasimus undatulus (Say) (Coleoptera: Cleridae), Temnochila

chlorodia (Mannerheim) (Coleoptera: Trogositidae), and Enoclerus sphegeus Fabricius

(Coleoptera: Cleridae). All DFB in the samples were counted and sexed. Beetles captured

in each trap were summed over the trapping period to determine the total number of

beetles removed from the population by each trap.

When the traps were deployed, seven trees in the Douglas-fir stand adjacent to the clear

cut were baited with pheromones to initiate DFB attack. These trees were spaced about 75

m apart in a line roughly parallel to the trap line. The line of trees and trap line were 150-

200 m apart. A commercially available tree bait (Phero Tech Inc., Delta, BC, Canada)

containing frontalin and a-pinene was stapled to each trap tree at a height of 2-3 m. In

addition to the commercial tree bait, frontalin (20 mg) and seudenol (10 mg) in PVC

formulations were attached to the tree boles. Mean diameter at breast height (dbh) of trap

trees was 66 cm (SE + 2.5), and mean height was 36.3 m (SE + 1.2).

Trap trees were sampled on 28 July 1997, after the DFB flight had ended. Each tree

was climbed to determine height at the top of the infestation, circumference at the top of

the infestation, and to remove bark samples to estimate attack densities. In addition, height

at the base of the infestation and circumference at the base of the infestation were

measured. An axe was used to cut through the bark to determine if DFB galleries were

present. This was continued until no DFB galleries were found at the top or bottom of trap

trees. The average of the circumference at the base and top of the infestation was used

along with length of the infested bole to estimate the amount of infested bark area for each

tree based on the equation for the surface area of a cylinder. The areas surrounding trap

trees were surveyed to determine if there were any spill-over attacks on adjacent trees.

At three heights along the infested tree bole, four 100 cm? circular bark samples were

removed with an electric drill and hole saw. Sample heights were near the top, middle, and

bottom of the infested portion of the bole. Samples were placed in plastic bags and stored

in an ice chest until transported to the lab. In the lab, attack sites were determined for each

sample. Attack sites were distinguished from ventilation holes or exit holes by their angle

and the presence of packed frass.

To determine attack sites per tree, mean number of attack sites per cm? was multiplied

by the surface area of the infested tree bole. Because DFB is monogamous, each attack

site represents one pair of beetles that entered the trap tree. The total number of attack sites

was multiplied by two to determine total number of DFB caught in each tree.

J. ENTOMOL. SOC. BRIT. COLUMBIA 97, DECEMBER 2000 35

Catches of traps and trap trees were compared using a t-test. A square root transformation

was used to meet assumptions of equal variances. All tests were performed with the

statistical software JMP (ver 3.1.5, SAS Institute Inc., Cary, NC)

RESULTS

Mean infested tree bole surface area was 29.8 m* (SE + 4.0) ranging from 19.2 to 48.9

m’. Mean number of attack sites per tree was 3,320.8 (SE + 607.0). Mean attack densities

were 90 per m’ and did not differ significantly by height (P = 0.26). No trees adjacent to

trap trees were attacked by DFB.

The mean of the total number of beetles caught per trap over the season was 13,740.6

(SE + 2813.5). In comparison, trap trees captured on average 6641.6 (SE + 1213.9) beetles.

Significantly more beetles were captured in the traps than in the trap trees (P = 0.04).

Significantly more males were captured in traps than in trap trees (P = 0.04), assuming a

1:1 sex ratio in trap trees. In comparison, significantly more females were captured in trap

trees than in traps (P = 0.009). Mean percent male beetles caught in traps was 80.8 (SE +

0.66).

DISCUSSION

Pheromone-baited traps are used extensively to study the biology and behavior of many

bark beetle species. In addition, pheromone-baited traps have been implemented in

strategies to manage or monitor some pest species, or both (Lindgren and Borden 1983;

Billings 1985; Shore and McLean 1985). However, trap trees have been used more

commonly in the past to manage DFB populations than pheromone-baited traps. We could

find no published data comparing the efficacy of trap trees and pheromone-baited traps in

the management of DFB.

In our study, pheromone-baited traps were more effective at capturing DFB than trap

trees. More beetles were removed from the population with pheromone-baited traps than

trap trees. Because of damage to pheromone-baited traps, total trap catches were likely

higher than our final results indicate. Throughout the study, ten trap collections were lost

due to trap damage. Four of these occurred on 11 June when DFB activity was high. The

average trap catch for the two undisturbed traps on that date was 1,307 beetles. We do not

know exactly when the traps were damaged. If they were damaged immediately after they

were last emptied then they likely caught few beetles. However, if they were damaged just

before they were visited, then they may have caught as many as 5,228 additional beetles

that were not included in our estimate of the total catch. In operational programs, damage

to traps might be reduced by suspending them in non-host trees at a height where wildlife

and livestock could not disturb them. However, deploying and maintaining suspended

traps takes more time and, therefore, is more costly than for traps that are placed at ground

level.

Although our estimate of captured beetles in traps is higher than in trap trees, it is

possible that traps have an even greater impact on local beetle populations than suggested

by a simple comparison of numbers of captured beetles. Because the brood sex ratio is 1:1]

(Bedard 1937; Vité and Rudinsky 1957) and DFB is predominantly monogamous, removal

of one beetle could actually represent the removal of a mated pair. Since we do not know

what proportion of beetles collected in traps would have mated with one another if they

had not been captured, we cannot determine the actual impact of trapping on local beetle

populations. At one extreme, assuming that no beetles in the traps would have mated with

each other, then the traps actually could have removed twice as many mated pairs from the

population as indicated by the number of captured beetles.

36 J. ENTOMOL. SOC. BRIT. COLUMBIA 97, DECEMBER 2000

There is evidence from laboratory studies that suggests some male DFB may mate with

more than one female (Vité and Rudinsky 1957). However, there are no published data to

indicate how often this occurs under natural conditions. If DFB males mate more than

once under natural conditions, the removal of a single male beetle would not be equivalent

to removal of a mated pair. Courtship in DFB is initially aggressive (Ryker 1984) and

beetles may suffer significant damage during the mating process and gallery construction.

Consequently, it is likely that many re-emerging male beetles are damaged and incapable

of prolonged flights to locate new host trees and female beetles. With extended time

searching for host trees and female beetles, DFB males would be exposed to higher levels

of predation and other mortality factors. Until research is conducted to determine the

sexual behavior of DFB under field conditions, we cannot be certain of the impact of

removal of males from local breeding populations.

One possible reason that traps caught more DFB is that they continuously remove

beetles from the population for the entire season. In comparison, trap trees have a finite

capacity for trapping beetles. Once trees are fully colonized, MCH is released by adult

DFB to deter other beetles from colonizing the tree. Consequently, beetles arriving at trap

trees after they are fully colonized will attack nearby host trees if they are present, or they

will disperse in search of suitable habitat.

Pheromone-baited traps removed a significantly higher number of male beetles from

the population than trap trees. In comparison, trap trees removed a significantly higher

number of female beetles than traps. It is possible that by manipulating trap lure

components, a higher number of females could be captured. For example, addition of

ethanol to the trap lure increases both total number of beetles and the proportion of females

captured (Ross and Daterman 1995c). However, this may not be important, because DFB

broods have a 1:1 sex ratio and the beetle is monogamous. Consequently, as discussed

above, removing a male or a female theoretically removes a mating pair of beetles from

the local population.

While a higher number of DFB are removed from local populations using traps

compared to trap trees, impacts on beneficial insects are likely less. For example, when

trap trees are harvested, beneficial insects inhabiting those trees are also removed from the

local population. Beneficial insects, including predators and parasitoids, have been shown

to cause high levels of mortality to several bark beetle species (Linit and Stephen 1983;

Weslien 1994; Schroeder and Weslien 1994; Schroeder 1996) and some may have a

regulating effect on populations (Reeve 1997; Turchin ef a/. 1999). Depending on timing

of DFB infestation and removal of trap trees, beneficial insects including Coeloides

brunneri Vierick (Hymenoptera: Braconidae), Medetera aldrichii Wheeler (Diptera:

Dolichopidae), Thanasimus undatulus, Enoclerus sphegeus, Temnochila chlorodia, and

possibly others could still be developing within or inhabiting host trees. Removal of these

species may significantly impact natural controls in subsequent bark beetle generations.

While traps catch several predaceous beetle species, the impact on local populations is

unknown. Many 7. undatulus are often captured in traps. This beetle preys on DFB, but

laboratory studies suggest that it prefers smaller species of Scolytus and Pseudohylesinus

(Schmitz 1978). To minimize the possible impact of removing predators from the

population, trap modifications can be employed to prevent their capture or provide for

their escape (Ross and Daterman 1998). Additionally, traps do not capture parasitoids

because they are not attracted to pheromones.

In addition to catching higher numbers of bark beetles, traps have several other

advantages. First, traps are easily deployed and can be placed almost anywhere there is the

threat of tree mortality. Traps, unlike trap trees, can be located in non-host stands or

openings to minimize attacks on nearby host trees. Pheromones and traps are relatively

J. ENTOMOL. SOC. BRIT. COLUMBIA 97, DECEMBER 2000 BH,

inexpensive and traps can be used for several to many years depending upon their method

of construction. Also, by using traps, no trees need be sacrificed.

Pheromone-baited traps are effective at capturing large numbers of DFB, thus

removing beetles from the breeding population in local areas. Natural resource managers

should consider substituting traps for trap trees in their management plans for DFB. By

doing this, more beetles may be removed from local populations, while valuable trees need

not be sacrificed.

ACKNOWLEDGEMENTS

The authors thank Tiffany Neal for technical assistance in the field. We also thank

personnel on the Salmon River Ranger District, Nez Perce National Forest, particularly

Jennifer Nelson and Cynthia Onthank, for locating the research site, providing maps and

access to the site, and for technical assistance in the field. This research was supported by

funds from the USDA Forest Service, Forest Health Protection, Special Technology

Development Program. Mention of a proprietary product does not constitute an

endorsement or recommendation for its use by USDA or Oregon State University.

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Shore, T.L. and J.A. McLean. 1985. A survey for the ambrosia beetles 7rypodendron lineatum and

Gnathotrichus retusus (Coleoptera: Scolytidae) in a sawmill using pheromone-baited traps. The

Canadian Entomologist 117: 49-55.

Turchin, P., A.D. Taylor, and J. D. Reeve. 1999. Dynamical role of predators in population cycles of a

forest insect: an experimental test. Science 285:1068- 1071.

Vité, J.P. and J.A. Rudinsky. 1957. Contribution toward a study of Douglas-fir beetle development.

Forest. Science 3: 156-167.

Weslien, J. 1994. Interactions within and between species at different densities of the bark beetle /ps

typographus (Coleoptera: Scolytidae) and its predator Thanasimus formicarius (Coleoptera: Cleridae).

Entomologia Experimentalis et Applicata 71: 133-143.

J. ENTOMOL. SOC. BRIT. COLUMBIA 97, DECEMBER 2000 39

The Chrysopidae of Canada (Neuroptera): recent

acquisitions chiefly in British Columbia and Yukon

J. A. GARLAND

1011 CARLING AVENUE, OTTAWA, ONTARIO, CANADA K1Y 4E7

ABSTRACT

Chrysopidae collected since 1980 chiefly in British Columbia and Yukon, Canada, and

some late additions collected before 1980, are reported. Nineta gravida (Banks) is

reported for the first time in the last 90 years. This is the first supplement to the inventory

of Chrysopidae in Canada.

Key words: Neuroptera, Chrysopidae, Canada

INTRODUCTION

The chrysopid fauna of Canada, as presently understood (Garland 1984, 1985), has been

fully inventoried up to 1980 (Garland 1982). Since then, newly collected specimens in British

Columbia and the Yukon, and some older-dated specimens not previously seen, have become

available. The purpose of publishing these specimen label data is to supplement the already

extensive inventory of label data on the Canadian chrysopid fauna, thereby extending it to the

year 2000. Materials and methods appropriate to this study have been documented elsewhere

(Garland 2000). All specimens reported here are deposited in the Spencer Entomological

Museum, Department of Zoology, University of British Columbia. Acronyms used below:

BC, British Columbia; SK, Saskatchewan; and YK, Yukon Territory.

FAMILY CHRYSOPIDAE Schneider, 1851

SUBFAMILY CHRYSOPINAE Schneider, 1851

A total of 57 specimens belonging to 10 recent species of Chrysopinae are reported here, as

follows:

Chrysopa chi Fitch

BC: 1 ¢, Gavin L[ake], 25.v.1987 (R. Reich); 1 3, id, 3.vi.1987 (R. Reich).

Chrysopa coloradensis Banks

BC: 1 &, Penticton, 23.vili.1983 (R.J. Cannings).

Chrysopa nigricornis Burmeister

BC: 1 &, Galiano I[sland], Spanish Hills, 26.vi.1987 (G.G.E. Scudder); 1 3, Osoyoos,

15.vii.1990 (G.G.E. Scudder); 4 do, id, 16.vii.1990 (G.G.E. Scudder); 1 3, Osoyoos, East

Bench, 30.viii.1997 (G.G.E. Scudder); 1 3, Penticton, 23.viii.1983 (R.J. Cannings); 1 3,

Vanc[ouver], 4. 6 [vi]. [19]25, Permanent Loan from Vancouver City Museum.

Chrysopa oculata Say

YK: 1 2, Alaska Hwy, 29.vi.1974 (G.G.E. Scudder); 2 2 2, McCabe Cr[eek], 8 km S[outh],

30.vi.1985 (E. Krebs & J.J. Robinson); 1 2, Pelly Crossing, 30.vi.1985 (E. Krebs & J.J.

Robinson); 1 2, Tatchun L[ake], 29.vi.1985 (E. Krebs & J.J. Robinson).

BC: 1 2, Dutch Cr[eek], 1 km N{orth], 31.vili.1998 (G.G.E. Scudder); 1 2, F[or]t Nelson,

1367 [collected in alcohol], 2.vili. 1982 (G.G.E. Scudder); | 2, Galiano I[{sland], North end,

25.v1.1989 (G.G.E. Scudder); 1 ¢, Moyie, 1.ix.1998 (G.G.E. Scudder); 1 2, Osoyoos L[ake],

Haynes Ecol[ogical] Res[erve], 27.vi.1981 (S.G. Cannings); | 2, Sparwood, 20.vi.1982

(G.G.E. Scudder).

40 : J. ENTOMOL. SOC. BRIT. COLUMBIA 97, DECEMBER 2000

SK: 1 o, Regina, 18.vi.1943 (P. Larkin).

Chrysopa pleuralis Banks

BC: 1 &, Osoyoos, M[oun]t Kobau R[oa]d, km 17.1, 1720 m, IDFdk1,6VK:F/2DYd4/2RO,

sweeping K3, 28.vil.1997 (G.G.E. Scudder).

Chrysoperla carnea (Stephens)

BC: 1 3, Alaska Hwy, km 670, Racing River, |.viii.1982 (G.G.E. Scudder); 1 2, Alaska Hwy,

32.1 km E[ast of] Steamboat, 1.vili.1982 (G.G.E. Scudder); 1 ¢, Charlie L[ake], 25 km

Wlest], 5.vii.1982 (G.G.E. Scudder); 3 oo, 10 22, Flor]t Nelson, 1367 [collected in

alcohol], 2.vili.1982 (G.G.E. Scudder); 1 2, Galiano I[sland], Spanish Hills, 26.vi.1987

(G.G.E. Scudder); 1 2, Nanaimo, 7.v.1987 (G.G.E. Scudder); 1 3, Sonora I[sland], Owen

Bay, 26.vii.1986 (G.G.E. Scudder); 1 3, 1 2, W[est] Vancouver, 12.viii.[19]34 (G.H.

Larnder), Permanent Loan from Vancouver City Museum; 5 33,2 2 2, Wood Creek, 2 km

‘N[orth of] Toad River, 1361 [collected in alcohol], 1.viii.1982 (G.G.E. Scudder).

Meleoma dolicharthra (Navas)

BC: 1 3, Galiano I[sland], Spanish Hills, 26.vi.1987 (G.G.E. Scudder).

Meleoma emuncta (Fitch)

BC: 1 ¢, Penticton, 23.vili.1983 (R.J. Cannings).

Meleoma signoretti Fitch

BC: 1 ¢ [left FW & abdomen missing], W[est] Vancouver, 12.viil.[19]34 (G.H. Larnder),

Permanent Loan from Vancouver City Museum.

Nineta gravida (Banks)

BC: | 3, Gabriola Island, 19.vii.1999 (R.D. Kenner & G.S. Kenner).

DISCUSSION

For all ten species, the provincial and territorial distributions reported here were already

known (Garland 1982, 1985; cf Penny et al. 1997). However, for some of the species, the

present data include important new locality records, e.g., Chrysopa pleuralis in the South

Okanagan just west of Osoyoos; for Chrysoperla carnea, all the specimens were yellow-green,

characteristic of summer generations of this insect, which is thought to be bivoltine about the

latitude of Whitehorse, Yukon and southward (Garland 1989); and, for Nineta gravida, the

specimen reported here marks the first collecting record of the species for British Columbia in

over 90 years, since 1908 in fact (Garland 2000).

ACKNOWLEDGEMENTS

Dr. G. G. E. Scudder, Professor Emeritus, University of British Columbia, kindly made the

specimens mentioned here available for study.

REFERENCES

Garland, J.A. 1982. The Taxonomy of Chrysopidae of Canada and Alaska (Insecta: Neuroptera). Ph. D. Thesis,

McGill University, Canada. 1: 418 pp. 2: 132 figs., 24 maps.

Garland, J.A. 1984. Catalogue of Chrysopidae of Canada and Alaska (Neuroptera). Neuroptera International

3:93-94. [Legal deposit, 31 December 1984]

Garland, J.A. 1985. Identification of Chrysopidae in Canada, with bionomic notes (Neuroptera). The Canadian

Entomologist 117:737-762, 1278 (errata).

Garland, J.A. 1989. Phénologie de l’espéce holarctique Chrysoperla carnea (Stephens) (Neuroptera:

Chrysopidae) dans la partie septentrionale de |!’ Amérique du Nord. Neuroptera International 5:181-183.

Garland, J.A. 2000. Rediscovery of Nineta gravida (Banks, 1911) in British Columbia, and review of the genus

Nineta Navas, 1912 in Canada (Neuroptera: Chrysopidae). Journal of Neuropterology 3: [in press].

Penny, N.D., P.A. Adams, and L.A. Stange. 1997. Species catalog of the Neuroptera, Megaloptera, and

Raphidioptera of America north of Mexico. Proceedings of the California Academy of Sciences 50:39-114.

J. ENTOMOL. SOC. BRIT. COLUMBIA 97, DECEMBER 2000 4

Comparison of a-pinene and myrcene on attraction of

mountain pine beetle, Dendroctonus ponderosae (Coleoptera:

Scolytidae) to pheromones in stands of western white pine

DANIEL R. MILLER ! AnD B. STAFFAN LINDGREN ”

PHERO TECH INC., 7572 PROGRESS WAY, DELTA, BC V4G 1E9

ABSTRACT

Multiple-funnel traps baited with exo-brevicomin and a mixture of cis- and trans-

verbenol were used to test the relative attractiveness of myrcene and (-)-a-pinene to the

mountain pine beetle, Dendroctonus ponderosae Hopkins, in a stand of western white

pine, Pinus monticola Dougl. Traps baited with myrcene caught significantly more D.

ponderosae than traps baited with (-)-a-pinene, irrespective of the presence of exo-

brevicomin. exo-Brevicomin was attractive to Thanasimus undatulus (Say)

(Coleoptera: Cleridae) whereas 7rypodendron lineatum (Olivier) (Coleoptera:

Scolytidae) was attracted to (-)-a-pinene. Our results support the use of myrcene in

commercial trap lures and tree baits for D. ponderosae in stands of western white pine

in British Columbia.

Key words: Scolytidae, Dendroctonus ponderosae, kairomones, Pinus monticola,

Trypodendron lineatum, Cleridae, Thanasimus undatulus

INTRODUCTION

The mountain pine beetle, Dendroctonus ponderosae Hopkins (Coleoptera: Scolytidae),

has killed over 500 million lodgepole, Pinus contorta var. latifolia Engelm., ponderosa, P.

ponderosa P. Laws. and western white pines, P. monticola Dougl. (Pinaceae) in British

Columbia over the past 80 years (Unger 1993). The current integrated pest management

program for D. ponderosae in BC (Maclauchlan and Brooks 1994) is cost-effective, with

positive economic, social and environmental impacts (Miller et a/. 1993).

Semiochemicals play an important role in several tactics within the program

(Maclauchlan and Brooks 1994). Population levels and flight periods of D. ponderosae are

monitored with multiple-funnel traps baited with commercial lures consisting of the

pheromones, exo-brevicomin and cis- and trans-verbenol, and the kairomone, myrcene

(Stock 1984; Maclauchlan and Brooks 1994). The spread of infestations has been curtailed

by the application of commercial tree baits consisting of the same semiochemicals (Borden

and Lacey 1985; Borden et al. 1986) or simply the pheromones, exo-brevicomin and cis-

and ¢rans-verbenol (Borden et al. 1993).

Semiochemical blends for these commercial lures and baits were developed in stands

of lodgepole and ponderosa pine rather than western white pine, and discrepancies exist

concerning the most appropriate kairomone. The host compound a-pinene was more

effective than myrcene in enhancing attraction of D. ponderosae to trans-verbenol in

stands of western white pine in Idaho (Pitman 1971). Myrcene was more effective than a-

pinene in increasing attraction of D. ponderosae to pheromones in stands of ponderosa

' Current address: USDA Forest Service, Southern Research Station, 320 Green Street,

Athens, GA 30602

* Current address: College of Science and Management, University of Northern British

Columbia, Prince George, BC V2N 4Z9

42 J. ENTOMOL. SOC. BRIT. COLUMBIA 97, DECEMBER 2000

pine (Billings et al. 1976) and lodgepole pine (Borden ef al. 1983; Conn ef al. 1983). In

lodgepole pine stands, catches of D. ponderosae in pheromone-baited traps exhibited a

dose-dependent increase to both myrcene and 3-carene whereas a-pinene had no effect

(Miller and Borden 2000).

Our objective was to verify the effectiveness of myrcene, relative to a-pinene, in

commercial lures for D. ponderosae in stands of western white pine. Specifically we

attempted to compare the response of beetles to pheromones in traps baited with o-pinene

to those baited with myrcene. Our expectation was that myrcene would be as effective as

a-pinene in attracting D. ponderosae.

MATERIALS AND METHODS

Semiochemical-Releasing Devices. Phero Tech Inc. (Delta, British Columbia) supplied

polyethylene bubble-cap lures containing a 13:87 mixture of frans- and cis-verbenol [both

chemical purities 98%; both enantiomeric compositions 83:17 (-):(+)], (+)-exo-brevicomin

polyurethane flex lures (chemical purity >98%), and separate closed, low-density

polyethylene bottles (15 mL) containing either a-pinene [chemical purity >98%:;

enantiomeric composition > 99% (-)] or B-myrcene (chemical purity > 98%). The

verbenols were released at a combined rate of approximately 1.74 mg/d at 24 °C

(determined by weight loss) whereas oa-pinene and myrcene were released at

approximately 413 mg/d and 281 mg/d at 24-28 °, respectively (determined by weight

loss). exo-Brevicomin was released at approximately 0.1 mg/d at 24 °C (determined by

collection of volatiles) (Phero Tech Inc.).

Experiments. Two experiments were conducted in a mature stand of western white pine

with approximately 15% of live trees infested by D. ponderosae near Barriere, British

Columbia (51°10’N, 120°8’W). In both experiments, forty 8-unit multiple-funnel traps

(Lindgren 1983) (Phero Tech Inc.) were set 10-15 m apart, and > 2m from any tree, along

two parallel transect lines spaced approximately 20 m apart. Each trap was suspended

between trees by rope such that the top funnel of each trap was 1.3—1.5 m above ground.

In Experiment 1, the effect of a-pinene and myrcene on the attraction of D. ponderosae to

traps baited with the verbenol mix was determined, with and without exo-brevicomin. All

traps, baited with the verbenol mix, were set on 1 August 1990. The following treatments

were randomly assigned to 10 traps each: (1) a-pinene; (2) myrcene; (3) a-pinene and exo-

brevicomin; and (4) myrcene and exo-brevicomin. Experiment | was terminated on 25

August 1990.

Experiment 2 tested the interaction between a-pinene and myrcene on the attraction of

D. ponderosae to traps baited with exo-brevicomin and the verbenol mix. All traps, baited

with the verbenol mix and exo-brevicomin, were set on 25 August 1990. The following

treatments were randomly assigned to 10 traps each: (1) no kairomone control; (2) a-

pinene; (3) myrcene; and (4) a-pinene and myrcene. Experiment 2 was terminated on 12

September 1990.

Catches of D. ponderosae and serendipitous catches of Trypodendron lineatum

(Olivier) (Coleoptera:Scolytidae) and Thanasimus undatulus (Say) (Coleoptera: Cleridae)

were tallied for each treatment. Sexes of D. ponderosae captured in Experiment 2 were

determined by dissection and examination of genitalia. Voucher specimens were deposited

at the Entomology Museum, Simon Fraser University, Burnaby, BC.

Statistical Analyses. Trap catch data were analysed by 2-way ANOVA using the

SYSTAT statistical package version 8.0 (SPSS 1998). The model factors in Experiment |

were exo-brevicomin, monoterpenes (a-pinene or myrcene), and the interaction between

exo-brevicomin and monoterpenes. In Experiment 2, the model factors were a-pinene,

J. ENTOMOL. SOC. BRIT. COLUMBIA 97, DECEMBER 2000 43

myrcene and the interaction of a-pinene and myrcene. Catches of D. ponderosae were

transformed by In(Y) to remove heteroscedasticity whereas catches of Thanasimus

undatulus and Trypodendron lineatum were transformed by In(Y+1) due to zero catches in

some treatments. Sex ratio data, expressed as percentage of males in catches, from

Experiment 2 were transformed by arcsine(Y). Fisher’s least significant difference (LSD)

multiple range tests were performed when P < 0.05.

RESULTS AND DISCUSSION

Our results clearly support the retention of myrcene in commercial lures for D.

ponderosae in stands of western white pine. Catches of D. ponderosae were significantly

higher in traps baited with myrcene than in traps baited with a-pinene (Figs. 1,2). exo-

Brevicomin did not affect the preference of beetles for myrcene over a-pinene (F) 36 =

0.605, P = 0.442) in Experiment | nor was there any interaction between a-pinene and

myrcene on catches of D. ponderosae (F\ 35 = 0.018, P = 0.894) in Experiment 2. There

was no significant difference in sex ratio among the treatments in Experiment 2 (F319 =

2.232, P = 0.121) with the mean (+ SE) percentage of males in catches at 55 + 3 %.

eB+M ; b

0 600 1200 1800

Mean (+SE) number of beetles

Figure 1. Effect of exo-brevicomin (eB), a-pinene (P) and myrcene (M) on the attraction

of Dendroctonus ponderosae to verbenol-baited multiple-funnel traps from 1 August to 25

August 1990 (7 = 10). Means followed by different letters are significantly different at P <

0.05 (LSD test).

44 J. ENTOMOL. SOC. BRIT. COLUMBIA 97, DECEMBER 2000

0 40 80 120

Mean (+SE) number of beetles

Figure 2. Effect of a-pinene (P) and myrcene (M) on the attraction of Dendroctonus

ponderosae to multiple-funnel traps baited with verbenols and exo-brevicomin from 25

August to 12 September 1990 (n = 10). Means followed by different letters are

significantly different at P < 0.05 (LSD test); control (c).

Our results are inconsistent with those of Pitman (1971) who demonstrated that a-

pinene was more effective than myrcene in attracting D. ponderosae in stands of western

white pine and are surprising since a-pinene is the most common monoterpene in the resin

of western white pine, which has low amounts of myrcene (Mirov 1961). The relative

proportion of myrcene is higher in the resin of lodgepole and ponderosa pines with

amounts of myrcene greater than or equal to amounts of a-pinene (Mirov 1961; Shrimpton

1973). Geographic variation in semiochemical responses, similar to that in /ps pini (Say)

(Miller e¢ al. 1997), may explain some of these results.

Finally, research by Pitman (1971), Billings et al. (1976), Borden et al. (1983) and

Conn ef al. (1983) were conducted before the importance of the enantiomeric composition

of a-pinene was widely recognised. It is likely, but not certain, that they used either (+)- or

(-)-a-pinene due to the high costs associated with (+)-a-pinene. We used (-)-a-pinene in

our trials since it is the predominant enantiomer in the resin of western white pine phloem

tissue (Mirov 1961).

In Experiment 1, catches of Trypodendron lineatum were lowest in traps baited with

myrcene alone, and highest in traps baited with either a-pinene or exo-brevicomin (Table

1). a-Pinene significantly increases the attraction of 7. /ineatum to ethanol and the

J. ENTOMOL. SOC. BRIT. COLUMBIA 97, DECEMBER 2000 45

pheromone lineatin (Borden ef a/. 1982; Schroeder and Lindeléw 1989). No 7. lineatum

were caught in Experiment 2.

Table 1

Mean (+ SE) catches of Trypodendron lineatum (Scolytidae) and Thanasimus undatulus

(Cleridae) in verbenol-baited multiple-funnel traps from 1 August to 25 August 1990 “.

Treatment Trypodendron lineatum Thanasimus undatulus

(-)-a-Pinene je ee eeelta

Myrcene Dae lea ltla

(-)-a-Pinene + (+)-exo-brevicomin 2 26 282 C

Myrcene + (+)-exo-brevicomin 9+2b 1623 b

Means within the same column followed by the same letter are not significantly

different, P < 0.05 (LSD multiple comparison test).

The predator, Thanasimus undatulus, showed a preference for traps baited with exo-

brevicomin in combination with myrcene or a-pinene, particularly the latter (Table 1). As

might be expected for a generalist predator, similar results with 7. undatulus have been

reported with the following bark beetle pheromones: frontalin, exo- and endo-brevicomin,

ipsdienol, ipsenol, and cis-verbenol (Kline ef al. 1974; Dyer 1975; Chatelain and Schenk

1984; Miller et al. 1987; Miller and Borden 1990; Miller et a/, 1991; Miller et a/. 1997;

Poland and Borden 1997). Usually, 7. undatulus are not attracted to host tree compounds

(Furniss and Schmitz 1971; Miller and Borden 1990) although Macias-Samano et al.

(1998) demonstrated attraction of 7. undatulus to host blends from grand fir, Abies grandis

(Doug]l.) Lindl. No 7. undatulus were caught in Experiment 2.

ACKNOWLEDGEMENTS

We thank J.H. Borden, K.O. Britton, B. Sullivan and an anonymous reviewer for their

reviews of the manuscript. L.E. Maclauchlan, J.E. Macias-Samano and D. Piggin provided

technical and field assistance. This research was supported by the Science Council of

British Columbia.

REFERENCES

Billings, R.F., R.l. Gara and B.F. Hrutfiord. 1976. Influence of ponderosa pine resin volatiles on the

response of Dendroctonus ponderosae to synthetic trans-verbenol. Environmental Entomology 5: 171-

179:

Borden, J.H. and T.E. Lacey. 1985. Semiochemical-based manipulation of the mountain pine beetle,

Dendroctonus ponderosae Hopkins: a component of lodgepole pine silviculture in the Merritt Timber

Supply Area of British Columbia. Zeitschrift fiir angewandte Entomologie 99: 139-145.

Borden, J.H., C.J. King, S. Lindgren, L. Chong, D.R. Gray, A.C. Oehlschlager, K.N. Slessor and H.D.

Pierce, Jr. 1982. Variation in response of T7rypodendron lineatum from two continents to

semiochemicals and trap form. Environmental Entomology 11: 403-408.

Borden, J.H., J.E. Conn, L.M. Friskie, B.E. Scott and L.J. Chong. 1983. Semiochemicals for the mountain

pine beetle, Dendroctonus ponderosae (Coleoptera: Scolytidae), in British Columbia: baited-tree

studies. Canadian Journal of Forest Research 13: 325-333.

Borden, J.H., L.J. Chong and T.E. Lacey. 1986. Pre-logging baiting with semiochemicals for the mountain

pine beetle, Dendroctonus ponderosae, in high hazard stands of lodgepole pine. Forestry Chronicles

62: 20-23.

Borden, J.H., L.J. Chong, B.S. Lindgren, E.J. Begin, T.M. Ebata, L.E. Maclauchlan and R.S. Hodgkinson.

1993. A simplified tree bait for the mountain pine beetle. Canadian Journal of Forest Research 23:

1108-1113.

46 J. ENTOMOL. SOC. BRIT. COLUMBIA 97, DECEMBER 2000

Chatelain, M.P. and J.A. Schenk. 1984. Evaluation of frontalin and exo-brevicomin as kairomones to

control mountain pine beetle (Coleoptera: Scolytidae) in lodgepole pine. Environmental Entomology

13: 1666-1674.

Conn, J.E., J.-H. Borden, B.E. Scott, L.M. Friskie, H.D. Pierce, Jr. and A.C. Oehlschlager. 1983.

Semiochemicals for the mountain pine beetle, Dendroctonus ponderosae (Coleoptera: Scolytidae) in

British Columbia: field trapping studies. Canadian Journal of Forest Research 13: 320-324.

Dyer, E.D.A. 1975. Frontalin attractant in stands infested by the spruce beetle, Dendroctonus rufipennis

(Coleoptera: Scolytidae). The Canadian Entomologist 107: 979-988.

Furniss, M.M. and R.F. Schmitz. 1971. Comparative attraction of Douglas-fir beetles to frontalin and tree

volatiles. U.S. Department of Agriculture Forest Service Research Paper INT-96.

Kline, L.N., R.F. Schmitz, J.A. Rudinsky and M.M. Furniss. 1974. Repression of spruce beetle (Coleoptera

) attraction by methylcyclohexenone in Idaho. The Canadian Entomologist 106: 485-491.

Lindgren, B.S. 1983. A multiple-funnel trap for scolytid beetles. The Canadian Entomologist 115: 299-

302.

‘Macias-Samano, J.E., J.H. Borden, R. Gries, H.D. Pierce, Jr., G. Gries and G.G.S. King. 1998. Primary

attraction of the fir engraver, Sco/lytus ventralis. Journal of Chemical Ecology 24: 1049-1075.

Maclauchlan, L.E. and J.E. Brooks. 1994. Strategies and tactics for managing the mountain pine beetle,

Dendroctonus ponderosae. Kamloops Forest Region, British Columbia Forest Service. 60 pp.

Miller, D.R. and J.H. Borden. 1990. B-Phellandrene: kairomone for pine engraver, /ps pini (Say)

(Coleoptera: Scolytidae). Journal of Chemical Ecology 16: 2519-2531.

Miller, D.R. and J.H. Borden. 2000. Dose-dependent and species-specific responses of pine bark beetles

(Coleoptera: Scolytidae) to monoterpenes in association with pheromones. The Canadian Entomologist

132: 183-195.

Miller, D.R., J.H. Borden, G.G.S. King and K.N. Slessor. 1991. Ipsenol: an aggregation pheromone for /ps

latidens (LeConte) (Coleoptera: Scolytidae). Journal of Chemical Ecology 17: 1517-1527.

Miller, D.R., J.A. Carlson and M. Stemeroff. 1993. Socioeconomic analysis of mountain pine beetle

management in British Columbia. British Columbia Forest Service. 67 pp.

Miller, D.R., K.E. Gibson, K.F. Raffa, S.J. Seybold, S.A. Teale and D.L. Wood. 1997. Geographic

variation in response of pine engraver, /ps pini, and associated species to pheromone, lanierone.

Journal of Chemical Ecology 23: 2013-2031.

Miller, M.C., J.C. Moser, M. McGregor, J.C. Gregoire, M. Baisier, D.L. Dahlsten and R.A, Werner. 1987.

Potential for biological control of native North American Dendroctonus beetles (Coleoptera:

Scolytidae). Annals of the Entomological Society of America 80: 417-428.

Mirov, N.T. 1961. Composition of gum turpentines of pines. U.S. Department of Agriculture Forest

Service Technical Bulletin No. 1239.

Pitman, G.B. 1971. trans-Verbenol and alpha-pinene: their utility in manipulation of the mountain pine

beetle. Journal of Economic Entomology 64: 426-430.

Poland, T.M. and J.H. Borden. 1997. Attraction of a bark beetle predator, Thanasimus undatulus

(Coleoptera: Cleridae), to pheromones of the spruce beetle and two secondary bark beetles

(Coleoptera: Scolytidae). Journal of the Entomological Society of British Columbia 94: 35-41.

Schroeder, L.M. and A. Lindelow. 1989. Attraction of scolytids and associated beetles by different

absolute amounts and proportions of a-pinene and ethanol. Journal of Chemical Ecology 15: 807-817.

Shrimpton, D.M. 1973. Extractives associated with wound response of lodgepole pine attacked by the

mountain pine beetle and associated microorganisms. Canadian Journal of Botany 51: 527-534.

Stock, A.J. 1984. Use of pheromone baited Lindgren funnel traps for monitoring mountain pine beetle

flights. British Columbia Forest Service. 11 pp.

SPSS Inc. 1998. SYSTAT 8.0 Statistics. Chicago, IL. 1086 pp.

Unger, L. 1993. Mountain pine beetle. Forestry Canada Forest Pest Leaflet 76.

J. ENTOMOL. SOC. BRIT. COLUMBIA 97, DECEMBER 2000

Somatochlora kennedyi (Odonata: Corduliidae):

a new species for British Columbia, with notes on

geographic variation in size and wing venation

REX D. KENNER

5560 LINSCOTT COURT, RICHMOND, BC V7C 2W9

ABSTRACT

The first confirmed record for Somatochlora kennedyi Walker in British Columbia is

reported. Specimens of this species from the northern Yukon are smaller than those

from elsewhere in its range and have a reduced number of cells in certain parts of the

wings. The reduced number of cells may cause some keys to yield ambiguous results.

Keywords: Somatochlora, Corduliidae, Odonata, British Columbia, geographic

variation

INTRODUCTION

In the summer of 1997, the Conservation Data Centre (Ministry of Environment,

Lands and Parks) sponsored two expeditions to northeastern British Columbia to survey

the odonate fauna of that region. During the first, Leah Ramsay and I made a number of

interesting discoveries among which was the capture of a female Somatochlora kennedyi

Walker near Fort Nelson. This is the first confirmed record for this species in British

Columbia. While comparing this specimen to named S. kennedyi specimens in the

collection of the Spencer Entomological Museum (SEM), certain anomolies were noted

in the size and wing venation of specimens from the Yukon. That information is

presented here.

Somatochlora kennedyi is a southern boreal species with a distribution which extends

from the northern Yukon east to Newfoundland, south to Minnesota, Wisconsin,

Michigan; and east to Ohio, New York and Massachusetts (Walker and Corbet 1975;

Cannings et al. 1991; Bick and Mauffray 2000). The distribution is poorly known,

especially in the west; Cannings and Cannings (1994) noted that it is “unusual for a

southern boreal species to be unknown in British Columbia and Alberta”. The records

for the Yukon, Manitoba and the Northwest Territories are from late June and July

(Walker and Corbet 1975; Cannings ef al. 1991). Somatochlora kennedyi has been

reported from “sedge/rush and polygon sedge fens” and “deep sedge/moss marsh” in the

Yukon (Cannings and Cannings 1994) and “a shallow pond in a swampy wood” in New

Brunswick (Walker 1925).

MATERIAL EXAMINED

All specimens of S. kennedyi examined are deposited in the SEM, Department of

Zoology, University of British Columbia. All specimens were dried in acetone and are

stored in clear envelopes. The collection data for the BC specimen are | 2, Andy Bailly

Lake, S of Fort Nelson, 25 June 1997, R. D. Kenner. The details for the 20 Yukon

specimens are given in Cannings ef a/. (1991): 1 3, 1 2, Loon Lake (60° 02’ N 127° 35’

W) and 15 3,3 ¢, Old Crow area (2 separate sites).

48 J. ENTOMOL. SOC. BRIT. COLUMBIA 97, DECEMBER 2000

RESULTS AND DISCUSSION

The specimen from Andy Bailly Lake is a young female, caught while it was resting

on the road beside our vehicle. We saw no other individuals. Due to its teneral nature,

the abdomen partially collapsed during treatment with acetone for preservation. |

determined it to be S. kennedyi using the keys in Walker and Corbet (1975) and by

comparison with named specimens in the collection of the SEM. The specimen was also

examined by R. A. Cannings and S. G. Cannings who have previous experience with this

species and they confirmed the determination. The occurrence of S. kennedyi in BC was

expected (Cannings and Stuart 1977), especially since it has been collected in the

southeastern Yukon not far from the BC-Yukon border (Cannings et al. 1991).

Although this is the first confirmed record for BC, it may not be the first time S.

_kennedyi has been collected in BC. There is in the SEM, a previously unidentified final

stadium larval specimen which keys out as S. kennedyi. It was collected in sweeps of the

“moss/rush/sedge” in the fen at the south end of Eddontenajon Lake on 17 June 1987 by

S. G. Cannings. Separating the larvae of S. kennedyi from those of S. franklini (Selys)

depends on differences in the arrangement of setae on the dorsum of the abdominal

segments (Walker 1925; Walker and Corbet 1975). Some of these setae may break off

during capture and storage and it is difficult for me to be completely certain of the

identification without named material for direct comparison.

In keys for adult female Somatochlora sp. (Walker 1925; Walker and Corbet 1975)

the number of cells “in the fork of R” is one of several characters used for separating S.

kennedyi and S. franklini;, S. kennedyi has 11—20 cells and S. franklini has 6—9 cells. This

character is also used in the key in Needham and Westfall (1955). The data in Table |

show that the number of cells in the fork of R; is not a reliable character for separating S.

kennedyi and S. franklini in the northern Yukon. A more useful character appears to be

the colour of the lateral lobes of the postclypeus (brown in S. kennedyi but black in S.

franklini).

Table 1

Latitudinal variation in morphological characteristics of Somatochlora kennedyi from

British Columbia and the Yukon.

Andy Loon L. Old Crow Area Walker and Corbet

Bailly L. 60°02’N 67°35°N (1975)

58°49’N

1 9 lo 1 9 [536 3 9 3 9

Total 43.5 47 AQ’. 41-45 2 7 A0-43, ilo 47-55

length (42.5) (42)

(mm)

Hind wing 3029.5 33 -25.5-28 265-28 29/5-32 29-865

length (26.7), (O75)

(mm)

# cells in 12/14 12/1365 ais Al5 5-15 Ale gels 11-18

R, fore (10.8) (9.2)

wing

# cells in 12/14. 13/13. 18/19 9-15 9.17) 9 2-19 12-20

R> hind (11.9) (10)

win

Numbers in parentheses are mean values.

J. ENTOMOL. SOC. BRIT. COLUMBIA 97, DECEMBER 2000

The data in Table 1 also show that both total length and hind wing length for the

specimens from the Old Crow area are smaller than the lower limit given in Walker and

Corbet (1975). The partial collapse of the abdomen of the BC specimen may have

contributed to its apparent small size.

The literature contains a number of references to geographical variations in size.

Walker (1925) briefly discusses geographical variations in Somatochlora spp. and

reports that S. franklini is “larger towards the southern limit of its range” and S.

albicincta (Burmeister) is smaller in the far north and on the Labrador coast. Although

Tennessen (1977) states that a decrease in size with latitude is common in North

American odonates, there are a number of references which show either increases or

decreases in size with increasing latitude for both odonates and non-odonate insects (see,

for examples, citations in Stewart 1982 and Corbet 1999). Cannings (1982) showed that

the larvae of Sympetrum illotum Hagen increase in size with increasing latitude. It 1s

clear that factors other than latitude need to be taken into account in developing an

understanding of the observed size variations.

ACKNOWLEDGEMENTS

I thank S. G. Cannings for the opportunity and the Ministry of Environment, Lands

and Parks for financial support during the field work . | thank Leah Ramsay for being a

great field companion, R. A. Cannings and S. G. Cannings for examining the specimen

and G. G. E. Scudder and K. Needham for allowing me unlimited access to the

collections of the SEM and space to work.

REFERENCES

Bick, G. H. and W. Mauffray. 2000. Distribution summary of North American Anisoptera.

http://www.afn.org/~iori/nalist.html. Last updated 10 September 2000.

Cannings, R. A. 1982. The larvae of the Tarnetrum subgenus of Sympetrum, with a description of the

larva of Sympetrum nigrocreatum Calvert (Odonata: Libellulidae). Advances in Odonatology |: 9-

14.

Cannings, R. A. and K. M. Stuart. 1977. The Dragonflies of British Columbia. British Columbia

Provincial Museum, Handbook No. 35, Victoria, 254 pp.

Cannings, S. G. and R. A. Cannings. 1994. The Odonata of the northern cordilleran peatlands of North

America. Memoirs of the Entomological Society of Canada 169: 89-110.

Cannings, S. G., R. A. Cannings and R. J. Cannings. 1991. Distribution of the dragonflies (Insecta:

Odonata) of the Yukon Territory, Canada with notes on ecology and behaviour. Contributions to

Natural Science, Number 13, Royal British Columbia Museum, Victoria, B.C., 27 pp.

Corbet, P. S. 1999. Dragonflies: Behavior and Ecology of Odonata, Cornell University Press, Ithaca,

N.Y., 829 pp.

Needham, J. G. and M. J. Westfall. 1955. A Manual of the Dragonflies of North America (Anisoptera)

including the Greater Antilles and the Provinces of the Mexican Border. University of California

Press, Berkeley, 615 pp.

Stewart, W. E. 1982. An analysis of geographic variation of the adults of the Australian genus

Diphlebia Selys (Odonata:Amphipterygidae). Australian Journal of Zoology, 30: 435-460.

Tennessen, K. J. 1977. Rediscovery of Epitheca costalis (Odonata: Corduliidae). Annals of the

Entomological Society of America 70: 267-273.

Walker, E. M. 1925. The North American Dragonflies of the Genus Somatochlora. University of

Toronto Studies, Biological Series, 26: 202 pp.

Walker, E. M. and P. S. Corbet. 1975. The Odonata of Canada and Alaska, Volume 3. University of

Toronto Press, Toronto, 307 pp.

J. ENTOMOL. SOC. BRIT. COLUMBIA 97, DECEMBER 2000 5]

Heteroptera (Hemiptera: Prosorrhyncha) new to Canada.

Part 1

G.G.E. SCUDDER

DEPARTMENT OF ZOOLOGY, UNIVERSITY OF BRITISH COLUMBIA,

VANCOUVER, BC V6T 124

ABSTRACT

The occurrence of 34 species of true bugs newly recognized in Canada is documented.

INTRODUCTION

In preparing the recently published checklist of the Hemiptera of Canada and Alaska

(Maw ef al. 2000), various collections across Canada were examined to document the

distribution of the species. In the process, 34 species new to Canada were identified. The

locality and collection records for these new discoveries are published herein, so as to

validate the inclusion of the species in the above mentioned checklist. Additional species

will be published in Part 2 when all determinations have been confirmed. The records of

new Miridae to Canada are being published elsewhere (Schwartz and Scudder 2000).

The arrangement in the list below is according to the checklist (Maw ef a/. 2000), and

in general follows the catalogue by Henry and Froeschner (1988). The latter publication

provides references to the descriptions of the various species and associated literature.

References to keys for identification are included in Henry and Froeschner (1988) and

Maw et al. (2000). Where data label uses common name of host, the scientific name has

been added in parentheses.

Museum abbreviations used in the text are as follows (curators in parentheses):

APM: Alberta Provincial Museum, Edmonton, AB (A.T. Finnamore).

CNC: Canadian Collection of Insects, Agriculture and Agri-Food, Ottawa, ON (R.G.

Foottit).

LEMQ: Lyman Entomological Museum, McGill University, Macdonald College

Campus, St.-Ann-de-Bellevue, QC (T.A. Wheeler).

RBCM: Royal British Columbia Museum, Victoria, BC (R.A. Cannings).

ROM: Royal Ontario Museum, Toronto, ON (D. Currie).

UBC: Spencer Entomological Museum, Department of Zoology, University of British

Columbia, Vancouver, BC (K.M. Needham).

UG: Department of Environmental Biology, University of Guelph, Guelph, ON (S.A.

Marshall).

SPECIES NEW TO CANADA

Infraorder NEPOMORPHA

Family CORIXIDAE

Ramphocorixa acuminata (Uhler)

ON: 2¢ 12, Essex Co., South Gosfield Tp., Wigle Creek Sta. 2, 42°03.3'N 82°46.5'W,

MNR Lot No. 8282-028, 2.x1.1982 (J. Blackburn; H. Tardif) [ROM].

32 J. ENTOMOL. SOC. BRIT. COLUMBIA 97, DECEMBER 2000

Trichocorixa kanza Sailer

ON: 12, Greeley, gravel pit, 2.xi.1983 (D.J. Larson) [CNC].

Trichocorixa louisianae Jaczewski

NS: 12, Sable Is., 8.vi.1966 (W.R.M. Mason) [CNC]; 3¢, Sable Is., West end,

].vii.1967 (H.F. Howden) [CNC]; 473 402, Sable Is., 11-15.ix.1967 (J.E.H.

Martin) [CNC].

Trichocorixa verticalis verticalis (Fieber)

PE: 1c 12, Queens Co., Mount Steward, 21.vili.1971 [CNC].

Infraorder GERROMORPHA

Family HEBRIDAE

Hebrus buenoi Drake & Harris

ON: 1c, Arkell, 20.iv.1976 (W.J. Moolenbeck) [UG]; 2¢, Damascus, 2.vi.1976 (S.A.

Marshall) [UG]; 1¢, Guelph, 23.v.1977 (Kevin Barber) [UG]; 12, Guelph,

18.vii.1977 (W.A. Attwater) [UG]; 16, id., 21.vii.1977 [UG].

Merragata brunnea Drake

ON: 12, Arkell, pond #2 (B.S. Heming) [UG]; 12, Guelph, 22.ix.1983 (Nancy R.

Ennis) [UG]; 1¢, Guelph, 22.1x.1983 (Edward Hippert) [UG]; 1¢, Walpole I.,

11.vii.1977 (E.A. Inns) [UG].

Family VELIIDAE

Microvelia hinei Drake

NS: 26,72, Falmouth, pond #1, pH 6.1, 20.v.1984 (G.G.E. Scudder) [CNC, UBC].

ON: 12, Walpole I., 27.vi.1985 (G.G.E. Scudder) [CNC].

This Ontario record was included in a tabulation in Scudder (1987), but without a

locality record.

Steinovelia stagnalis (Burmeister)

ON: 12, Rondeau Pr. Pk., Marsh Trail, treading 7ypha in marsh, 2.v1.1985 (A. Davies,

J.M. Campbell) [CNC].

Infraorder LEPTOPODOMORPHA

Family SALDIDAE

Chiloxanthus arcticus (Sahlberg)

NT: Io 192, Richards Is., Kidluit Bay, 30.vili.1948 (W.J. Brown) [CNC].

Saldula balli Drake

BC: 1, Vernon, vii.1920 (N.L. Cutler) [UBC].

Infraorder CIMICOMORPHA

Family TINGIDAE

Corythucha bellula Gibson

MB: 7c 72, Aweme, Ribes, 19.v.1919 (N. Criddle) [CNC]; 12, Aweme, Corylus,

30.v.1919 (N. Criddle) [CNC]; 40 12, Aweme, Alnus incarna, 13.vili.1930 (R.M.

White) [CNC]; 12, Darlington, swept from saskatoons (Amelanchier sp.),

24.v.1930 (R.M. White) [CNC]; 23, Lake Audy, alder (A/nus sp.), 1.vi.1941 (R.D.

Bird) [CNC].

ON: 42, Bothwell, 29.v.1929 (G.S. Walley) [CNC]; 7c 122, Haggersville, on

Crataegus, 9.vii.1962 (Kelton & Thorpe) [CNC].

J. ENTOMOL. SOC. BRIT. COLUMBIA 97, DECEMBER 2000 53

Corythucha obliqua Osborn & Drake

BC: 364, Summerland, Ceanothus, 7.vii.1975 (L.A. Kelton) [CNC].

Gargaphia angulata Heidemann

ON: 12, Belle River, 7.vi.1961 (Kelton & Brumpton) [CNC]; 12¢ 152, Grand Bend,

7.1x.1961 (L.A. Kelton) [CNC]; 4c 59, Grand Bend, 15.vi.1962 (Kelton &

Thorpe) [CNC]; 1¢ 12, Pinery Pt. Pk., 8.ix.1961 (J. Brumpton) [CNC].

Hesperotingis antennata Parshley

BC: 26, Cranbrook, Ponderosa pine (Pinus ponderosa), 23.vii.1959 (L.A. Kelton)

[CNC]. |

MB: 12, Aweme, 25.viii.1922 (R.M. White) [CNC].

SK: 12, Cypress Hills, S. Maple Cr. (117), 20.vii.1956 (Lindroth) [CNC]; 2<¢,

Saskatoon, 4.viii.1925 (Kenneth M. King) [CNC].

Hesperotingis fuscata Parshley

AB: 1¢ 12, Brocket, 18 km NNW, 49°41'N 113°54'W, 1350m., pantrap, 18-22.vii.1998

(K. White) [CNC]; 40 19, id., pitfall trap [CNC; Scudder Coll; White Coll.]; 2¢

12, id., 6-10.vili.1998 [CNC]; 23 12, id. 10-14.vili.1998 [CNC]; 1d, id. 14-

18.viii.1998 [CNC]; 1¢ 12, CFB Suffield, NWA, 50°37.678'N 110°18.371'W, pan

trap, 16-28.vi1.1994 (A.T. Finnamore) [APM]; 1¢ 12, id., 16-29.vii.1994 [APM];

1d 12, id., 16-28.vii.1994 [APM]; 1a 22, id., 50°23.466'N 110°36.768'W, 16-

28.vii.1994 [APM].

BC: 16, Fairview, White L., BGxhl, SWm, pan trap P-2, 20.vi-27.vi.1995 (J. Jarrett)

[UBC]; 23, id., pan trap P-9, 27.vi-4.vii.1995 [UBC]; 12, Kilpoola L., PPxhl,

pitfall trap KL3-4, 15.vii-6.vii.1996 (J. Jarrett & G.G.E. Scudder) [UBC]; 12, id.,

pitfall trap KL3-5 [UBC]; 16, id., pitfall trap KL1-5, 23.vi-28.vii.1997 (J. Jarrett)

[UBC].

Hesperotingis occidentalis Drake

AB: 292, Kananaskis, 20.vii.1974 (L.A. Kelton) [CNC]; 1¢, Lundbreck, 7.vii.1970 (L.A.

Kelton) [CNC].

BC: 298, Fernie, 1.vii.1934 (Hugh B. Leech) [CNC].

Leptopharsa heidemanni (Osborne & Drake)

ON: 5¢ 122, Ojibway, 6-7.vi1.1961 (Kelton & Brumpton) [CNC].

Family REDUVIIDAE

Emesaya brevipennis brevipennis (Say)

ON: 1(?), Chatham, 24.1x.1927 (C.W. Smith) [CNC]; 1¢, Essex Co., 10.ix.1955 (W.

Kendrick) [UG]; 12, Freelton, Malaise, 28.viii.1984 (M.T. Kasserra) [UG]; 1¢,

Harrow, 10.ix.1950 (A.H. Kichi) [UG]; 1¢,London, 1958 [CNC]; 12 2 immature,

Pt. Pelee, 28.viil.1920 (N.K. Bigelow) [CNC]; 12, Pt. Pelee, 16.ix.1932 (G.M.

Stirrett) [CNC]; 2d, Pt. Pelee, 8.ix.1954 (R. Lambert) [CNC]; 12, Pt. Pelee,

9.1x.1954 (C.D. Miller) [CNC]; 22, Pt. Pelee, 13.ix.1961 (G. Brumpton) [CNC];

(7), Samia; 5.x,1907 (W.A, Dent) [CNC]; 12, id., 14.xi,1913 [CNC]; le 1°, id,

x.1914 [CNC]; 2d, S. School, 1937 (E.A. Watts, M. Fletcher) [UG]; 1¢ 12, St.

Thomas, x.1920 (H.G. Crawford) [CNC]; 14, Windsor, Malaise, 24.vii.1984 (M.T.

Kasserra) [UG]; 12,Yarmouth, 1946 [CNC].

Empicoris culiciformis (DeGeer)

BC: 19, Chilliwack, 1.vi.1924 [RBCM]; 6c 32, Victoria, 24.v.1921 (W. Downes)

[CNC].

54 J. ENTOMOL. SOC. BRIT. COLUMBIA 97, DECEMBER 2000

ON: 12, Chatham, 29.vi.1928 (G.J. Spencer) [CNC]; 1o¢ 12, Don Mills, in home with

pinned insects infested with carpet beetles, x.1959 (G.B. Wiggins) [ROM]; 192,

Toronto, 12.viii.1964 (B. Pulley) [ROM].

Empicoris parshleyi (Bergroth)

QC: 40 52, Brome, under bark on maple (Acer sp.), 9.vi.1936 (G.S. Walley) [CNC];

12, Covey Hill, 8.vii.1937 (GS. Walley) [CNC]; Ilo, Rigaud, 25.vi.1906

(Beaulieu) [CNC].

Infraorder PENTATOMOMORPHA

Family ARADIDAE

Aradus basalis Parshley

‘NB: 12, Bathurst, vi.1930 (J.N. Knull) [CNC].

QC: 12, Mt. Jacques Cartier, 5.vii.1954 (W.J. Brown) [CNC].

Family COREIDAE

Acanthocephala terminalis (Dallas)

ON: 12, Kent Co., 28.viii.1955 (G. McDonald) [UG]; 1¢, Kent Co., Forest at Hwy.

401, 12.x.1997 (S.A. Marshall) [UG]; 1¢, Lambton Co., North Lambton,

30.vil.1996 (J. Skevington) [UG]; 12, Lambton Co., Pinery Prov. Pk., 21.vii.1992

(J. Carmichel) [UG]; 1¢ 52, Leamington, 29.v.1937 (G.S. Walley) [CNC]; 3 2, id.,

31.v.1937 [CNC]; 3°, id., 1.vi.1937 [CNC]; lo, id., 9: vil93 TENG ae o..id.

11.vi.1937 [CNC]; 12%, Simcoe, 6.vi.1939 (G.E. Shewell)*[EN@i|at2* Simcoe:

24.vi.1939 (T.N. Freeman) [CNC].

Catorhintha mendica Stal

MB: 12, Aweme, 11.vi.1922 (R.M. White) [CNC].

ON: 16, St. Catharines, goldenrod (Solidago sp.), 13.1x.1964 [ROM].

Family RHOPALIDAE

Arhyssus crassus Harris

BC: 12, Osoyoos, 26.vill.1986 (G.G.E. Scudder) [UBC]; 1a 22, Osoyoos, Mt. Kobau,

1525m., 26.vili.1986 (G.G.E. Scudder) [UBC]; 1a, Osoyoos, Mt. Kobau Rd., km

6.7, on flowers of Holodiscus discolor, 10.vi1.1994 (G.G.E. Scudder) [UBC]; 1°,

Osoyoos, Mt. Kobau Rd., km 1.8, PPxhl, WAw, pitfall trap, 23.vi-28.vii.1997 (J.

Jarrett) [UBC].

Aufeius impressicollis Stal

BC: 16, Osoyoos IR, nr. Mud L., 49°13'N 119°31'W, Purshia assoc., BGxhl, AN,

pitfall trap, 4.x.1994-10.iv.1995 (G.G.E. Scudder) [UBC].

Family ARTHENIDAE

Chilacis typhae (Perris)

BC: 22%, Osoyoos, 7.7 km N, jct. Hwy. 97/Rd. 22, 49°05'23"N 119°32'49"W, on Typha

latifolia, 8.x.1999 (G.G.E. Scudder) [UBC]; 73 102, id., 16.x.1999 [UBC]; 40¢

382, Osoyoos, 8 km N, 49°05'32"N 119°32'18"W, on Typha latifolia, 16.x.1999

(G.G.E. Scudder) [CNC, RBCM, UBC].

ON: 16, Nepean, Piney Forest, Lafontaine House, ex MV lite, 18.vii.1991 (M.D.

Schwartz) [CNC]; 2d, id, 27-viilgo lh | ONe|:

J. ENTOMOL. SOC. BRIT. COLUMBIA 97, DECEMBER 2000 55

Family LYGAEIDAE

Kleidocerys modestus Barber

BC: 4¢ 1, Vernon, BX Range, 610m [2000’], 4.vii.1979 (R.A. Cannings) [UBC].

Family CY DNIDAE

Macroporus repetitus Uhler

BC: 2¢ 12, Vaseux Cr., ‘CWS Bench’, 49°16'N 119°30'W, Purshia assoc., BGxhl,

AN, pitfall trap, 9.v-3.vi.1994 (G.G.E. Scudder [UBC]; 2¢ 32, id., 3.vi-8.vii.1994

[UBC]; 22, id., 8.vi-5.vi1.1995 [UBC].

Melanaethus robustus Uhler

ON: 19, Pelee I., Fish Point, 28.vi.1985 (G.G.E. Scudder) [CNC].

Microporus obliquus Uhler

AB: 36, CFB Suffield, NWA, 50°37.678'N 110°18.371'W, 16.vi.1995 (A.T. Finnamore)

[APM]; 1a 22, id., 16-28.vii.1994 [APM]; 1c 22, id., 16-29.vi.1994 [APM]; 132,

id., 28.vil-16.vill.1994 [APM]; 23 12, id., 16.vill-7.1x.1994 [APM].

BC: 282, Oliver, IRI, ‘Watertower’, 49°10'N 119°31'W, Purshia assoc., BGxhl, AN,

pitfall trap, 3.v-7.v1.1995 (G.G.E. Scudder) [UBC]; 12, Osoyoos, Haynes Ecol.

Res., BGxhl, AN recovery after fire, pitfall trap, 9.vii-7.viil.1994 (G.G.E. Scudder)

[UBC]; 1¢, id., 9.vili-3.1x.1995 [UBC]; 1o 22, id., 14.v-9.vi.1996 [UBC]; 12, id.,

9.vi-9.vil.1996 [UBC]; 2c 12, id, 9.vilt-20.1x.1996 [UBC]; 1°, id. 19.vi-

18.vii.1997 [UBC]; 22, id., 12.v-28.vi.1998 [UBC].

Family PENTATOMIDAE

Murgantia histrionica (Hahn)

ON: 420 52, Southampton, on Brassica, 15.1x.1956 (P.N. Vroom) [CNC].

Family SCUTELLERIDAE

Euptychodera corrugata (Van Duzee)

AB: 12, Manyberries, 19 km S, jct. 501-502, 13.viii.1982 (S.G. & R.A. Cannings)

[UBC].

Family THY REOCORIDAE

Corimelaena alpina (McAtee & Malloch)

NB: 192, Fredericton, French Lake, 12.vi.1931 (C.W. Maxwell) [LEMQ].

OC 12, Laval, 7.viul.1938 (LEMQ].

Corimelaena obscura McPherson & Sailer

ON: 24, Kingsville, 8.vii.1977 (A.A. Konecny) [UG].

ACKNOWLEDGEMENTS

Research for this paper was supported by grants from the Natural Sciences and

Engineering Research Council of Canada. I thank the curators of museums mentioned in

the text for loan of material and/or permission to examine the collections in their

institutions. I am indebted to Drs. R.C. Froeschner (National Museum of Natural History,

Smithsonian Institution, Washington, DC) and A. Jansson (University of Helsinki) for help

and confirmation of some of the determinations.

56 J. ENTOMOL. SOC. BRIT. COLUMBIA 97, DECEMBER 2000

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.

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

Canada and Alaska. NRC Research Press, Ottawa.

Schwartz, M.D. and G.G.E. Scudder. 2000. Miridae (Heteroptera) new to Canada, with some taxonomic

changes. Journal of the New York Entomological Society (in press).

Scudder, G.G.E. 1987. Aquatic and semiaquatic Hemiptera of peatlands and marshes in Canada. Memoirs

of the Entomological Society of Canada 140: 65-98.

J. ENTOMOL. SOC. BRIT. COLUMBIA 97, DECEMBER 2000 ST

Pheromone interruption of pine engraver, [ps pint,

by pheromones of mountain pine beetle,

Dendroctonus ponderosae (Coleoptera: Scolytidae)

DANIEL R. MILLER ‘| and JOHN H. BORDEN

CENTRE FOR ENVIRONMENTAL BIOLOGY, DEPARTMENT OF BIOLOGICAL

SCIENCES, SIMON FRASER UNIVERSITY, BURNABY, BRITISH COLUMBIA VSA 1S6,

CANADA

ABSTRACT

The effect of pheromones of Dendroctonus ponderosae Hopkins on the attraction of

Ips pini (Say) to its pheromone, ipsdienol, was investigated in stands of lodgepole pine.

The mixture of cis- and trans-verbenol significantly reduced catches of /. pini in traps

baited with ipsdienol at three locations in British Columbia. exo-Brevicomin had no

effect on catches of / pini, irrespective of the enantiomeric composition of exo-

brevicomin. Ipsdienol did not significantly reduce the attraction of D. ponderosae to

traps baited with cis- and trans-verbenol, and (+)-exo-brevicomin.

Key Words: Coleoptera, Scolytidae, /ps pini, Dendroctonus ponderosae, pheromone

interruption, synomone, exo-brevicomin, cis-verbenol, trans-verbenol, ipsdienol

INTRODUCTION

The pine engraver, /ps pini (Say), and the mountain pine beetle, Dendroctonus

ponderosae Hopkins (Coleoptera: Scolytidae), are common bark beetle species in stands of

lodgepole pine, Pinus contorta var. latifolia Engelmann (Pinaceae), in western North

America (Furniss and Carolin 1980). /ps pini breeds in the phloem tissue of dead, dying or

downed lodgepole pines (Furniss and Carolin 1980). Dendroctonus ponderosae breeds in

the healthy phloem tissue of live, standing pine trees (Unger 1993). During the past 80

years, D. ponderosae has killed more than 500 million pine trees in British Columbia alone

(Unger 1993). Densities of D. ponderosae galleries on infested material range from 10 —

261/m? with optimal brood production densities of 75 — 85/m* (Safranyik and Linton

1998). Population levels of 7. pini can build up during drought conditions, or following

catastrophic events such as logging, fire, windthrow, or epidemics of D. ponderosae, with

attack densities reaching 200-300/m? (Safranyik et al. 1996). At times, populations of /

pini may be sufficiently large that they initiate attacks on live, standing trees. Two years

after the 1988 fire in the greater Yellowstone Park area, 44 % of the lodgepole pines were

infested by J. pini (Amman and Ryan 1991).

In spite of their abundance and similarity in phloem resource requirements, these two

Species maintain ecological and reproductive isolation by assembling on host material in

large non-overlapping, single-species aggregations. Dendroctonus ponderosae generally

infests the lower bole of standing trees whereas /. pini attacks mid- and upper-bole regions,

or the entire tree bole in the absence of D. ponderosae (Furniss and Carolin 1980).

Separation of aggregations seems to be facilitated by semiochemicals. /ps pini uses

ipsdienol (2-methyl-6-methylene-2,7-octadien-4-ol) as an aggregation pheromone (Birch

et al. 1980; Lanier et al. 1980) with both sexes preferring a racemic blend throughout most

' Current address: USDA Forest Service, Southern Research Station, 320 Green Street,

Athens, GA 30602

58 J. ENTOMOL. SOC. BRIT. COLUMBIA 97, DECEMBER 2000

of British Columbia (Miller et al. 1996). Dendroctonus ponderosae produces various

semiochemicals and responds optimally to the combination of exo-brevicomin (exo-7-

ethyl-5-methyl-6,8-dioxabicyclo[3.2.1]octane) and cis- and trans-verbenol (cis- and trans-

4,6,6-trimethylbicyclo[3.1.1]hept-3-en-2-ol) (Borden ef al. 1987; Miller and Lafontaine

1991).

Mutual interruption of pheromone attraction can enhance specificity in bark beetle

aggregations (Byers 1989). Ipsdienol, produced by male /. pini, interrupts the attraction of

D. ponderosae to the semiochemical blend of myrcene, (+)-exo-brevicomin, and cis- and

trans-verbenol (Hunt and Borden 1988). The pheromone blend of (+)-exo-brevicomin and

cis- and trans-verbenol, produced by D. ponderosae, interrupts the attraction of J. pini to

its pheromone ipsdienol (Hunt and Borden 1988). The effects of individual components

are not known. Therefore, our study assessed the effects of (+)-, (-)-, and (+)-exo-

brevicomin, and the mix of cis- and ¢rans-verbenol, separately and in combination, on the

attraction of /. pini to ipsdienol. Specifically, we expected that all these compounds would

reduce trap catches of male and female /. pini to ipsdienol-baited multiple-funnel traps.

MATERIALS AND METHODS

Chemicals and Release Devices. Phero Tech Inc. (Delta, British Columbia) supplied

polyethylene bubble-cap lures containing a 13:87 mixture of trans- and cis-verbenol [both

chemical purities 98%; both enantiomeric compositions 83:17 (-):(+)]. The verbenols were

released at a combined rate of ca. 1.74 mg/d at 24 °C (determined by weight loss). In 1986,

Phero Tech Inc. supplied laminar (+)-exo-brevicomin lures (chemical purity 98%). In

1987, each exo-brevicomin lure consisted of an open polyethylene microcentrifuge tube

(400 mL) (Evergreen Scientific, Los Angeles, California) containing one 3-cm-long glass

capillary tube (i.d. 13 mm; o.d. 15 mm) filled with exo-brevicomin. Phero Tech Inc.

supplied (+)-exo-brevicomin (chemical purity 98%) and B.D. Johnston (Department of

Chemistry, Simon Fraser University, Burnaby, British Columbia) supplied (+)-exo-

brevicomin [chemical purity >99%; enantiomeric composition 97:3 (+):(-)] and (-)-exo-

brevicomin [chemical purity >99%; enantiomeric composition 2:98 (+):(-)]. The release

rates of exo-brevicomin were approximately 0.12 mg/d at 25 °C in 1986 (determined by

collection of volatiles on Porapak-Q) and approximately 0.15 mg/d at 20 °C in 1987

(determined by weight loss).

(+)-Ipsdienol (chemical purity >98%) was obtained from Bedoukian Research Inc.

(Danbury, Connecticut). In 1986, each ipsdienol lure consisted of eight Microcap”

disposable pipettes (2uL) (Drummond Scientific Co., Broomall, Pennsylvania), each

pipette sealed at one end, filled with (+)-ipsdienol and placed in an open polyethylene,

microcentrifuge tube (1.8 mL) (Evergreen Scientific). In 1987, each ipsdienol lure

consisted of a 10-cm length of C-flex® tubing (i.d. 1.6 mm; o.d. 3.2 mm) (Concept Inc.,

Clearwater, Florida), filled with an ethanol solution of (+)-ipsdienol, and heat-pressure

sealed at both ends. The release rates of ipsdienol were approximately 0.08 mg/day at 24

°C in 1986 (determined by weight loss) and approximately 0.6 mg/day at 24 °C in 1987

(determined by collection of volatiles on Porapak-Q). Ethanol, used in the formulation to

reduce the risk of polymerization of ipsdienol, is not attractive to /. pini (Miller 1990).

Experiments. Three experiments were conducted in 1986-1987. In all experiments,

replicates of 8-unit Lindgren multiple-funnel traps (Phero Tech Inc.) were set in mature

stands of lodgepole pine. Replicates were spaced at least 100 m apart, and traps were

spaced 10-15 m apart within each replicate. Each trap was suspended by rope between

trees such that the top of each trap was 1.3-1.5 m above ground level. No trap was within 2

m of any tree.

J. ENTOMOL. SOC. BRIT. COLUMBIA 97, DECEMBER 2000 59

Experiment | tested the effect of ipsdienol, (+)-exo-brevicomin and verbenols on the

attraction of J. pini and D. ponderosae. Ten replicates of five traps/replicate were set on 4

August, 1986, in regular pentagon formations near Princeton, British Columbia. The

following treatments were randomly assigned within each replicate: (1) ipsdienol alone;

(2) ipsdienol and (+)-exo-brevicomin; (3) ipsdienol and verbenols; (4) ipsdienol, (+)-exo-

brevicomin and verbenols; and (5) (+)-exo-brevicomin and verbenols. The experiment was

terminated on 3 September, 1986.

In 1987, experiment 2 tested the effect of (+)-exo-brevicomin and verbenols on the

attraction of /. pini to ipsdienol at three sites in British Columbia: Princeton, Williams

Lake and Radium. At each site, five replicates of four traps/replicate were set in grids of 2

X 2 on 16 July, 7 September, and 9 September, respectively. The following treatments

were randomly assigned within each replicate: (1) tpsdienol alone; (2) ipsdienol and (+)-

exo-brevicomin; (3) ipsdienol and verbenols; and (4) ipsdienol, (+)-exo-brevicomin and

verbenols. Trapping was terminated at the three sites on 29 September, 3 October, and |

October, 1987, respectively.

In 1987, experiment 3 tested the effect of enantiomeric composition of exo-brevicomin

on the attraction of /. pini to ipsdienol. Five replicates of five traps/replicate were set on 20

August, each in a regular pentagon formation near Princeton, British Columbia. The

following treatments were randomly assigned within each replicate: (1) ipsdienol alone;

(2) ipsdienol and (-)-exo-brevicomin; (3) ipsdienol and (+)-exo-brevicomin; (4) ipsdienol

and (+)-exo-brevicomin; and (5) ipsdienol and double (+)-exo-brevicomin. The separate

release rates of (-)- and (+)-exo-brevicomin in treatments 2, 3 and 5 were identical whereas

the combined release rate of both enantiomers in treatments 2, 3 and 4 were identical. The

total release rate of exo-brevicomin in treatment 5 was twice that of exo-brevicomin in

treatment 4. The experiment was terminated on 29 September, 1987.

Sexes of /. pini were determined using declivital characters (Wood 1982) whereas

those of D. ponderosae were determined by dissection and examination of genitalia.

Voucher specimens were deposited at the Entomology Museum, Simon Fraser University.

Statistical Analyses. The data were analyzed with the SYSTAT statistical package

(version 8.0) (SPSS 1998). Trap catch data from all experiments were transformed by

In(Y+1) whereas sex ratio data (for catches > 5) were transformed by arcsineV(Y). All data

were analyzed by one-way ANOVA, followed by Fisher’s least-significant-difference

(LSD) multiple comparison test when P < 0.05. In addition, data from experiment 2 were

analyzed by full-factorial three-way ANOVA using location, verbenol mix and exo-

brevicomin as the model factors.

RESULTS

The treatments in experiment | had a significant effect on catches of /. pini (F 444 =

15.85, P < 0.001) and D. ponderosae (F 42.5 = 4.48, P = 0.006). Three replicates were

excluded in the analyses for D. ponderosae because no beetles were captured in these

replicates. |The combination of (+)-exo-brevicomin and cis- and trans-verbenol

significantly interrupted the attraction of /. pini to its pheromone ipsdienol, reducing mean

catches of /. pini to levels similar to those in traps baited only with (+)-exo-brevicomin and

cis- and trans-verbenol (Fig. 1). Mean catches in traps baited with either ipsdienol and (+)-

exo-brevicomin or ipsdienol and the verbenol mixture were not significantly different from

mean catches in traps baited only with ipsdienol. The response of D. ponderosae was the

converse of /. pini with the highest catches in all traps baited with the verbenol mixture

(Fig. 1). There was no significant effect of treatment on sex ratios for either /. pini (F 72, =

0.36, P = 0.705) or D. ponderosae (F 36 = 2.47, P = 0.099). The mean percentages (+SE)

60 J. ENTOMOL. SOC. BRIT. COLUMBIA 97, DECEMBER 2000

of male /. pini and D. ponderosae in trap catches were 33 (+ 3) % and 47 (4 3) %,

respectively.

2 Ips pini

eB+V ae

eB+Vtlid

V+id

eB + Id

D. ponderosae

0 20 40 60 0 40 80 120

Mean (+SE) number of beetles

Figure 1. Effect of ipsdienol (Id), (+)-exo-brevicomin (eB), and cis- and trans-verbenol

mixture (V) on the attraction of Ips pini and Dendroctonus ponderosae to multiple-funnel

traps in experiment | in 1986 (N = 10). Mean trap catches, within the same figure,

followed by the same letter are not significantly different at P = 0.05 (Fisher’s LSD test).

In experiment 2, the verbenol mixture had a significant effect on catches of /. pini

(Table 1). The effect was consistent for all three regions since no interaction term was

significant. Catches of /. pini to ipsdienol-baited traps were significantly reduced by the

verbenol mixture, with or without (+)-exo-brevicomin (Fig. 2). There was no significant

effect of (+)-exo-brevicomin on trap catches (Table 1). In all three regions, catches of /.

pini in traps baited with ipsdienol and (+)-exo-brevicomin were not significantly different

from those in traps baited with ipsdienol alone (Fig. 2). There was no effect of treatment

on sex ratios of /. pini in trap catches (Table 1). The mean (4SE) percentage of males in

trap catches was 25 (+ 1) %.

Table 1

Analysis of variance on effects of location (Princeton, Williams Lake, and Radium, BC),

verbenol mixture, and (+)-exo-brevicomin on number and sex ratio of /ps pini responding

to ipsdienol-baited multiple-funnel traps in 1987 (Experiment 2).

Trap catch * Proportion of males E

Source df F P df F P

Location (A) 2 24.26 <0.001 2 0.63 0.539

Verbenol mix (B) | 27-73" <0001 l 0.06 0.809

(+)-exo-Brevicomin (C) 1 0.04 0.833 l 0:19" 01669

AB 3 0.15 0:862 2 0:57" 0567

AC 2 0.92 0.406 2 0.31 0.733

Bac l 0.26 0.610 l 1.08 0.304

A BAC 2 0335" > 09703 2 126°" 50i293

Error 48 47

“ Data transformed by In(Y + 1).

” Data transformed by arcsineV(Y).

J. ENTOMOL. SOC. BRIT. COLUMBIA 97, DECEMBER 2000 6]

Radium Williams Lake Princeton

0 100 200 0O 300 600 0 300 600

Mean (+SE) number of Ips pini

Figure 2. Effect of (4)-exo-brevicomin (eB) and cis- and trans-verbenol mixture (V) on

the attraction of Jps pini to ipsdienol (Id) - baited multiple-funnel traps in experiment 2 in

1987 (N = 5). Mean trap catches, within the same figure, followed by the same letter are

not significantly different at P = 0.05 (Fisher’s LSD test).

In experiment 3, the enantiomeric composition of exo-brevicomin had no significant

effect on trap catches of /. pini (F 420 = 0.54, P= 0.707) or sex ratio of captured /. pini

(F 420 = 1.65, P = 0.202) (Fig. 3). The mean (SE) percentage of males in trap catches was

27 (£2) %.

2(+/-)eB + Id

(+)eB + Id

(+/-)eB + Id

(-)eB + Id

0 250 500 0 20 40

Mean (+SE) number of /ps pini Percentage (+SE) male

Figure 3. Effect of enantiomeric composition of exo-brevicomin (eB) on the attraction of

Ips pini to ipsdienol (Id) - baited multiple-funnel traps in experiment 3 in 1987 (N = 5).

Mean trap catches, within the same figure, followed by the same letter are not significantly

different at P = 0.05 (Fisher’s LSD test).

62 J. ENTOMOL. SOC. BRIT. COLUMBIA 97, DECEMBER 2000

DISCUSSION

Bark beetles use semiochemicals to ensure ecological and reproductive isolation (Byers

1989). Host partitioning within the southern pine bark beetle guild of five species occurs

through pheromone specificity and mutual interruption of pheromone attraction (Smith er

al. 1993). The principal pheromones and synomones are ipsenol, ipsdienol, frontalin,

verbenone, brevicomins and verbenols (Smith ef a/. 1993). Similarly in Europe, separation

among six species of ps DeGeer is maintained by pheromone blends of ipsenol, ipsdienol,

amitinol, myrtenol, and verbenols (Kohnle ef a/. 1988, 1993).

The same phenomenon is apparent among western species of bark beetles in North

America as well. Mutual interruption of pheromone response occurs between /. pini and D.

ponderosae in stands of lodgepole pine in British Columbia. Our results substantiate prior

.work by Hunt and Borden (1988) demonstrating that the attraction of / pini to its

pheromone is interrupted by pheromones of D. ponderosae. Specifically, we found that

attraction of /. pini to (+)-ipsdienol was clearly interrupted by the combination of cis- and

trans-verbenol (Figs. 1-2).

Additional work is required to separate the effects of cis-verbenol and trans-verbenol,

and their enantiomeric compositions, on the interruption of pheromone attraction by /. pini.

Our work employed a 13:87 mix of cis- and trans-verbenol with an overall enantiomeric

composition of 83:17 (-):(+), due to the lack of availability of pure compounds. Both cis-

and trans-verbenol are used by D. ponderosae as aggregation pheromones (Miller and

Lafontaine 1991) and likely have similar effects on the response of /. pini to its

pheromone.

Exo-brevicomin had no effect on the attraction of /. pini to ipsdienol (Figs. 1-2).

Pureswaran et al. (2000) demonstrated that exo-brevicomin significantly decreased catches

of male /. pini to (£)-ipsdienol-baited multiple-funnel traps near Princeton, BC. There was

no significant effect on catches of female /. pini (Pureswaran ef a/. 2000). It is possible that

our results with exo-brevicomin were due to an inappropriate dose range. We used devices,

which released exo-brevicomin at rates of 0.12-0.15 mg/d at 20-25 °C whereas Pureswaran

et al. (2000) used devices, which released exo-brevicomin at a rate of ca. 3.1 mg/d at

25.5€,

The enantiomeric composition of exo-brevicomin had no effect on trap catches of /.

pini (Fig. 3). Pureswaran et al. (2000) demonstrated antennal responses of male and female

I. pini to (+)-exo-brevicomin and (+)-endo-brevicomin. The antipodes, (-)-exo-brevicomin

and (-)-endo-brevicomin, elicited no response from /. pini. Further trials with exo-

brevicomin should be conducted with higher release rates of the (+)-enantiomer since the

lack of antennal activity with the (-)-enantiomer should correlate with a lack of field

activity.

In contrast to Hunt and Borden (1988), ipsdienol had no effect on the attraction of D.

ponderosae to the female-produced pheromones, cis- and trans-verbenol (Fig. 1). All our

experiments were conducted in late summer (August and September) whereas Hunt and

Borden (1988) demonstrated significant interruption in pheromone response in

experiments conducted in July. Their experiments conducted in early August failed to

demonstrate interruption of attraction of D. ponderosae to the blend of myrcene, exo-

brevicomin, and cis- and trans-verbenol by ipsdienol. It is possible that discrimination by

D. ponderosae differs during the season, possibly due to differential costs and benefits

related to the onset of colder temperatures (Reid 1962). Since the egg and early larval

stages are susceptible to high mortality from cold temperatures, beetles need to ensure that

eggs hatch and develop to the cold-tolerant 3-rd and 4-th larval stages prior to the arrival

of winter temperatures (Safranyik and Linton 1998). Additional work should be conducted

on the effect of another /. pini pheromone, lanierone (2-hydroxy-4,4,6-trimethyl-2,5-

cyclohexadienl-one), as an interruptant for D. ponderosae. Lanierone produced by male

J. ENTOMOL. SOC. BRIT. COLUMBIA 97, DECEMBER 2000 63

I, pini (Teale et al. 1991) significantly increases catches of /. pini to ipsdienol-baited traps

in British Columbia (Miller et a/. 1997).

Semiochemical specificity and mutual interruption of pheromones in western pine

forests is not limited to 1 pini and D. ponderosae. More than 50 species of bark beetles

have been reported on lodgepole pine, many of which are phloeophagous and maintain

ecological and reproductive separation (Wood 1982). Attraction of /. /atidens (LeConte) to

its pheromone, ipsenol (2-methyl-6-methylene-7-octen-4-ol), is interrupted by (+)-

ipsdienol (Miller and Borden 1992) whereas attraction of /. pini is interrupted by the

pheromone of /. /atidens, ipsenol (Borden et a/. 1992). The attraction of / integer

(Eichhoff) to lanierone is interrupted by ipsdienol whereas the attraction of / pini to

ipsdienol is enhanced by lanierone (Miller e¢ a/. 1997). Mutual interruption of pheromone

attraction also occurs between /. pini and I. paraconfusus Lanier (Birch and Wood 1975;

Birch et al. 1980) and between /. paraconfusus and D. brevicomis LeConte (Byers and

Wood 1980, 1981).

Semiochemical interruptants will play an important role in future integrated pest

management programs for bark beetles (Borden et al. 1992). For example, interruptants

can be used to minimize the likelihood that populations of / pini build up in slash

generated by thinning operations to such levels that they successfully attack and kill

standing trees (Borden ef al. 1992). Verbenone (4,6,6-trimethylbicyclo[3.1.1]hept-3-en-2-

one), an antiaggregation pheromone produced by D. ponderosae (Borden et al. 1987), and

ipsenol, a pheromone produced by J. /atidens (Miller et a/. 1991), interrupt the attraction of

I, pini to its pheromone (Borden ef al. 1992). The combination of verbenone and ipsenol

resulted in a 67% reduction in the number of downed lodgepole pines attacked by /. pini

and a 99% reduction in attack density (Borden ef al. 1992).

A complete understanding of the role and impact of the various pheromones and

kairomones is required to develop effective management programs. For example, the

combination of the interruptants for D. frontalis Zimmermann, verbenone and endo- and

exo-brevicomin, reduced the landings of D. frontalis on live loblolly pine by 84% with a

84% reduction in eggs laid (Payne and Richerson 1979). However, the treatment failed to

prevent tree mortality due to an increase in attacks by another bark beetle, / avulsus

(Eichhoff).

The risks and consequences of interruptants should be carefully considered in

management programs that facilitate interspecific competition to reduce the reproductive

potential of a pest species. Significant reductions in survivorship of D. frontalis can occur

by the practice of simply falling infested trees and abandoning them (Billings 1980). This

fall-and-leave practice apparently increases levels of competition by secondary bark

beetles, predation and parasitism (Billings 1980). Other researchers have used

semiochemicals to induce similar levels of competition with bark beetles in western North

America. Rankin and Borden (1991) used ipsdienol to induce attacks by /. pini on logs

previously infested with D. ponderosae, resulting in a 73% reduction in progeny of D.

ponderosae. Safranyik et al. (1998) obtained a 49% reduction in progeny of D. ponderosae

by baiting standing lodgepole pine with /. pini pheromones, ipsdienol and lanierone, when

baiting was conducted in September. In both studies, the attack densities of D. ponderosae

between control and treated trees were not significantly different. However, Safranyik ef

al. (1998) found that baiting standing trees with /, pini pheromones in August resulted in a

53% reduction in attack density of D. ponderosae with no difference in mean progeny

production between treated and control trees. Attractants used to initiate competition

against a pest species such as D. ponderosae should be applied with due consideration to

timing of application and appropriate combinations of semiochemicals.

64 J. ENTOMOL. SOC. BRIT. COLUMBIA 97, DECEMBER 2000

ACKNOWLEDGEMENTS

We thank K.O. Britton, G.L. DeBarr, D.S. Pureswaran and B.T. Sullivan for editorial

comments. Field and laboratory assistance was provided by L.J. Chong, C. Matteau and L.

Wheeler. Voucher specimens were deposited with the Entomology Museum at Simon

Fraser University. This research was supported in part by an H.R. MacMillan Family Fund

Scholarship and a Simon Fraser University Graduate Research Fellowship to DRM, the

Natural Sciences and Engineering Research Council of Canada, and the Science Council of

British Columbia.

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variation in response of pine engraver, /ps pini, and associated species to pheromone, lanierone.

Journal of Chemical Ecology 23: 2013-2031.

Payne, T.L. and J.V. Richerson. 1979. Management implications of inhibitors for Dendroctonus frontalis

(Col. Scolytidae). Bulletin de la Société Entomologique Suisse 52: 323-331.

Pureswaran, D.S., R. Gries, J.H. Borden and H.D. Pierce, Jr. 2000. Dynamics of pheromone production

and communication in the mountain pine beetle, Dendroctonus ponderosae Hopkins and the pine

engraver, /ps pini (Say). Chemoecology (in press).

Rankin, L.J. and J.H. Borden, J. H. 1991. Competitive interactions between the mountain pine beetle and

the pine engraver in lodgepole pine. Canadian Journal of Forest Research 21: 1029-1036.

Reid, R.W. 1962. Biology of the mountain pine beetle, Dendroctonus monticolae Hopkins, in the East

Kootenay region of British Columbia. I. Life cycle, brood development and flight periods. The

Canadian Entomologist 94: 531-538.

Safranyik, L. and D.A. Linton. 1998. Mortality of mountain pine beetle larvae, Dendroctonus ponderosae

(Coleoptera: Scolytidae) in logs of lodgepole pine (Pinus contorta var. latifolia) at constant low

temperatures. Journal of Entomological Society of British Columbia 95: 81-87.

Safranyik, L., T.L. Shore and D.A. Linton. 1996. Ipsdienol and lanierone increase /ps pini Say

(Coleoptera: Scolytidae) attack and brood density in lodgepole pine infested by mountain pine beetle.

The Canadian Entomologist 128: 199-207.

Safranyik, L., T.L. Shore and D.A. Linton. 1998. Effects of baiting lodgepole pines naturally attacked by

the mountain pine beetle with /. pini (Coleoptera: Scolytidae) pheromone on mountain pine beetle

brood production. Journal of the Entomological Society of British Columbia 95: 95-97.

Smith, M.T., S.M. Salom and T.L. Payne. 1993. The southern pine bark beetle guild: an historical review

of the research on the semiochemical-based communication system of the five principal species.

Virginia Agriculture Experiment Station Bulletin 93-4.

SPSS Inc. 1998. SYSTAT 8.0 Statistics. Chicago, IL. 1086 pp.

Teale, S.A., F.X. Webster, A. Zhang and G.N. Lanier. 1991. Lanierone: a new pheromone component from

Ips pini (Coleoptera: Scolytidae). Journal of Chemical Ecology 17: 1145-1158.

Unger, L. 1993. Mountain pine beetle. Forestry Canada Pest Leaflet 76.

Wood, S.L. 1982. The bark and ambrosia beetles of North and Central America (Coleoptera: Scolytidae), a

taxonomic monograph. Great Basin Naturalist Memoirs 6: 1-1359.

—

J. ENTOMOL. SOC. BRIT. COLUMBIA 97, DECEMBER 2000

Gyrinus cavatus and G. minutus (Coleoptera: Gyrinidae) in

British Columbia with comments on their nearctic

distributions

REX D. KENNER

c/o SPENCER ENTOMOLOGICAL MUSEUM,

UNIVERSITY OF BRITISH COLUMBIA, VANCOUVER, BC V6T 1Z4

ABSTRACT

The distributions of Gyrinus cavatus Atton and G. minutus Fabricius in British

Columbia (BC) were determined by examining adult specimens from a number of

museum collections. Gyrinus cavatus appears to be restricted to the eastern half of the

province except in the far north; G. minutus is more widespread. Outside BC, G.

cavatus is widely distributed with a range extending from Newfoundland to Alaska and

south as far as Kansas, with an apparently isolated population in southern Utah.

Gyrinus minutus, although generally more northern in distribution, extends south along

the Rocky Mountains as far as Colorado.

Key words: Gyrinidae, Gyrinulus, Gyrinus, cavatus, minutus, rockinghamensis,

British Columbia, Yukon, Alaska, nearctic, distribution

INTRODUCTION

The Gyrinidae are a small family of distinctive beetles that are superbly adapted to their

aquatic lifestyle (Ferkinhoff and Gundersen 1983, Hilsenhoff 1990). North American

Gyrinidae are classified in four genera, two of which, Dineutus MacLeay and Gyrinus

Miller, occur in British Columbia (BC) (Roughley 1991). Dineutus is represented by a

single species, Dineutus assimilis (Kirby), whose status in BC is questionable (Hatch

1953). The genus Gyrinus is represented in British Columbia by 17 species (Roughley

1991; Oygur and Wolfe 1991). One subgenus, Gyrinulus Zaitzev, includes three species

North America: Gyrinus minutus Fabricius, G. cavatus Atton and G. rockinghamensis

LeConte. The former two species occur in BC and their distribution forms the subject

this study. Gyrinus rockinghamensis is known from the east coast of North America (Fall

1922; Oygur and Wolfe 1991).

Atton (1990) showed that what was called G. minutus in North America was in fact

two species: G. minutus, with a relatively northern holarctic distribution, and G. cavatus,

with a more southerly distribution. Atton gave only Canadian localities for the two

species and he listed only two localities in BC for G. cavatus: Windermere and Mile 744

on the Alaska Highway. Oygur and Wolfe (1991) did not recognize G. cavatus, thus

their distribution map for G. minutus includes possible G. cavatus records. To better

understand the status of G. cavatus in BC, | have collected locality information by

examining specimens from a number of museums.

MATERIALS AND METHODS

During the course of this investigation, several thousand Gyrinus spp. specimens

from 20 museums and a private collection were examined, including 799 G. minutus and

G. cavatus, and several hundred G. rockinghamensis. In addition, G.G.E. Scudder

(University of British Columbia) supplied a list of records based on the specimens in the

Canadian National Collection in Ottawa (CNC), which I did not examine. No particular

68 J. ENTOMOL. SOC. BRIT. COLUMBIA 97, DECEMBER 2000

effort was made to be comprehensive in coverage of eastern North America. The

following museums and collectors kindly loaned material for this investigation:

American Museum of Natural History, L. Herman

W. F. Barr Entomological Museum, University of Idaho, F. W. Merickel

California Academy of Sciences, D. H. Kavanaugh

Carnegie Museum of Natural History, R. L. Davidson

Entomology Section, Oregon Department of Agriculture, R. L. Westcott

Essig Museum of Entomology, University of California, Berkeley, C. Barr

W. T. James Entomological Collection, Washington State University, R. S. Zack

Lyman Entomological Museum, McGill University, C. C. Hsiung

Museum of Zoology, University of Michigan, M. F. O’Brien

National Museum of Natural History, P. J. Spangler

Natural History Museum of Los Angeles County, B. V. Brown

Oregon State Arthropod Collection, Oregon State University, D. Judd

A. B. Richards, Lakewood, Colorado

Royal British Columbia Museum, R. A. Cannings

Royal Ontario Museum, D. Currie

Snow Entomological Museum, University of Kansas, R. W. Brooks

Spencer Entomological Museum, University of British Columbia, K. Needham

Strickland Museum, University of Alberta, D. Shpeley

University of Nebraska State Museum, G. Hall

University of Wyoming Insect Museum, S. R. Shaw

J. B. Wallis Museum, University of Manitoba, R. E. Roughley.

Gyrinus cavatus, G. minutus and G. rockinghamensis were separated using the

characters given in Fall (1922), Atton (1990) and Oygur and Wolfe (1991). Gyrinus

cavatus has dark abdominal sterna and a pale mesosternum with a medial sulcus and

deep right-triangular anterolateral depressions. Gyrinus minutus is completely dark

ventrally with a sulcate mesosternum having shallow oblique-triangular anterolateral

depressions. Gyrinus rockinghamensis is completely pale ventrally and has a

mesosternum with only a very shallow or no medial sulcus and very shallow or no

anterolateral depressions. Body length is also useful in separating G. cavatus and G.

minutus, it was measured as specified by Atton (1990), using an eyepiece graticule on a

stereomicroscope and is reported as mean +SE with n being the number of specimens. A

detailed list of the specimens examined can be obtained from the author and should

eventually be available on the Internet.

RESULTS AND DISCUSSION

Based on the specimens examined during this study, along with the records from the

CNC and Atton (1990), the distribution of G. cavatus extends from Newfoundland to

Alaska. The following jurisdictions should be added to the distribution given in the

“Checklist of Beetles of Canada and Alaska” (Roughley 1991): AK, ON, PQ and NF

(SK, the type locality, was also omitted). In the contiguous United States, it occurs from

Maine to New Jersey on the east coast, then west from Indiana and Michigan through

Illinois, Iowa, Nebraska and Kansas, north to the Canadian border (Wisconsin,

Minnesota, South and North Dakota and Wyoming). I have no records for Montana or

Idaho but I have seen one specimen labelled simply “Wash” and eight specimens

labelled “Was. T.”or “W. T.” (Washington Territory, an older designation for the Pacific

Northwest, P.J. Spangler, National Museum of Natural History, personal

communication) which may imply its occurrence in Washington. In addition, there is an

apparently disjunct population in Utah.

J. ENTOMOL. SOC. BRIT. COLUMBIA 97, DECEMBER 2000

In spite of its broad range, G. cavatus is relatively uniform morphologically with no

obvious regional variation other than the exception discussed below. Some individual

variability in ventral color was apparent but some of that may be due to differences in

preservation and treatment. Many specimens of G. cavatus have a slight convexity in the

sides of the median sulcus of the mesosternum that results in a slightly oval widening of

the sulcus near its mid-point. In a few specimens, that convexity is exaggerated to the

point where a flat-bottomed oval depression is formed in the medial sulcus. This occurs

in a majority of the specimens from the Aquarius Plateau in south-central Utah but also

occurs in a few specimens from other areas. The specimens from Utah are larger than

elsewhere (Table 1). This larger size is not simply a matter of increasing size with

decreasing latitude as the specimens from Kansas, at about the same latitude as Utah, are

similar in size to those from northern Canada.

Table 1

Body length (BL) of three Gyrinulus species by location: range, mean + SE and sample

size (Nn).

Locality Species BL (3) (mm) BL (?) (mm)

all areas G. cavatus 3.00 — 4.00 3.50 — 4.35

except Utah 3.67 + 0.012 (151) 3.99 = 0012 (140)

Utah G. cavatus 3.85 — 4.10 4.2

3.96 + 0.038 (9) (1)

Canada and G. minutus 3.75 — 4.45 4.00 — 4.85

Alaska 4.11 + 0.013 (144) 4.48 + 0.015 (127)

Wyoming G. minutus 4.00 — 4.45 4.35 — 4.95

4.28 + 0.035 (15) 4 32 0:035 (17)

Colorado G. minutus 4.3 -

(1)

all areas G. rockinghamensis 3.45 — 4.05 3.55 — 4.30

3.69 + 0.015 (70) 3.98 + 0.023 (47)

The distribution of G. minutus also stretches across Canada and Alaska although I

have very few records from the eastern provinces: one record from Newfoundland, three

from Quebec and one labelled “H. B.” (Hudson Bay, R.L. Davidson, Carnegie Museum

of Natural History, personal communication). It is restricted to the more northerly parts

of Manitoba and Saskatchewan and is widespread in the Northwest and Yukon

Territories and Alaska. In Alberta, it is found as far south as Edmonton, except in the

Rocky Mountains, where it is found at least as far south as Jasper. Its distribution

appears to extend south along the Rocky Mountains with records from Wyoming and

Colorado. It seems likely that it will be found in the southern Canadian Rockies and

Montana.

Gyrinus minutus is also relatively uniform in appearance over its large range in North

America; Atton (1990) found no difference between nearctic and palearctic specimens.

The male specimens from Wyoming and Colorado are slightly larger than those from

Canada although the females are little different (Table 1).

The distributions of G. cavatus and G. minutus in BC, Yukon Territory and Alaska,

based on this work and the sources cited above, are shown in Figure 1. In BC, G. cavatus

appears to be restricted to the eastern half of the province except in the far north; G.

minutus iS more widespread but is not found west of the Coast Mountains. The few

records for these species in BC suggest that either they are relatively uncommon or that

limited collecting was done in the appropriate habitat or season. The lack of data

precludes any possibility of determining habitat preferences which dictate the

distributions. I have two sites in BC at which both species were collected, Barkerville

and Summit Lake at mile 392 on the Alaska Highway. In addition, there are four such

70 J. ENTOMOL. SOC. BRIT. COLUMBIA 97, DECEMBER 2000

Figure 1. The distribution of Gyrinus cavatus (A) and G. minutus (B) in British

Columbia, Yukon Territory and Alaska based on the data on the labels of the specimens I

examined, the data on the labels of the specimens in the Canadian National Collection

(Ottawa) and localities given in Atton (1990). Alaska is plotted on a smaller scale than

British Columbia and Yukon Territory.

J. ENTOMOL. SOC. BRIT. COLUMBIA 97, DECEMBER 2000

sites in Alaska, three in the Yukon, four in the Northwest Territories and three in

Alberta.

I have records for both Gyrinus cavatus and G. rockinghamensis in New Jersey but

the overlap in their ranges is probably more extensive. The two species are superficially

very similar, being almost identical in size (Table 1) and both have pale mesosterna. |

have seen several examples of G. cavatus specimens which were misidentified as G.

rockinghamensis (see, for example, Figure 5 in Oygur and Wolfe (1991) which is clearly

a male G. cavatus). The characters discussed above should be adequate to reliably

separate the two species.

Studies such as the one reported here can only give a general sense of the overall

distribution for the species concerned. The locality labels on most specimens, especially

the older ones, give only the general location and carry no information on the habitat in

which the collection was made. Even so, one can look for correlations between the

ranges and geographic features or ecological zones. Although this study accomplished

what it set out to do, it generates several new questions. For example, what is the

southern limit for G. minutus across the prairie provinces and in eastern North America?

Roughley (1991) indicates that G. minutus occurs in every province and territory in

Canada except Prince Edward Island, but Roughley (1991) is based in part on data that

predate the description of G. cavatus. Do some of these records actually refer to G.

cavatus? Another question concerns the apparent gap in the distribution of G. cavatus

between New Jersey and Indiana. Does G. cavatus not occur in the intervening states?

Another study similar to this one but with a more eastern geographical focus would

probably answer such questions.

ACKNOWLEDGEMENTS

I thank the curators and staff of the various museums that loaned material for this

study, A. B. Richards for promptly informing me of the Colorado record for G. minutus,

R. E. Roughley for advice and copies of keys, G. G. E. Scudder for transcribing the

records from the Canadian National Collection in Ottawa, L. Lucas for producing the

maps in Figute 1 and K. Needham for allowing me unlimited access to the collections of

the Spencer Entomological Museum and space to work.

REFERENCES

Atton, F. M. 1990. Gyrinus (Gyrinulus) cavatus sp.nov. from North America described and compared

with Gyrinus (Gyrinulus) minutus Fabricius (Coleoptera: Gyrinidae). Canadian Entomologist 122:

651-657.

Fall, H. C. 1922. The North American species of Gyrinus (Coleoptera). Transactions of the American

Entomological Society 48: 269-306.

Ferkinhoff, W. D. and R. W. Gundersen. 1983. A key to the whirligig beetles of Minnesota and adjacent

states and Canadian provinces (Coleoptera: Gyrinidae). Scientific Publications of the Science

Museum of Minnesota 5: No. 3. pp. 53.

Hatch, M. H. 1953. The Beetles of the Pacific Northwest, Part 1: Introduction and Adephaga.

University of Washington Press, Seattle. pp. 340.

Hilsenhoff, W. L. 1990. Gyrinidae of Wisconsin, with a key to adults of both sexes and notes on

distribution and habitat. The Great Lakes Entomologist 23: 77-91.

Oygur, S. and G. W. Wolfe. 1991. Classification, distribution and phylogeny of North American (north

of Mexico) species of Gyrinus Miiller (Coleoptera: Gyrinidae). Bulletin of the American Museum

of Natural History 207. pp. 97.

Roughley, R. E. 1991. Gyrinidae. In “Checklist of Beetles of Canada and Alaska”, Bousquet, Y. (ed.),

Res. Branch; Agriculture Canada Pub. 1861/E. pp. 430.

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J. ENTOMOL. SOC. BRIT. COLUMBIA 97, DECEMBER 2000 73

Occurrence and inheritance of a colour pattern dimorphism

in adults of Hyalophora euryalus (Lepidoptera: Saturniidae)

W.D. MOREWOOD!

DEPARTMENT OF BIOLOGY, UNIVERSITY OF VICTORIA

P.O. BOX 3020 STATION CSC, VICTORIA, BC V8W 3N5

ABSTRACT

A white prothoracic collar and white abdominal rings are among the characters used to

distinguish adults in the genus Hyalophora Duncan (Lepidoptera: Saturniidae) from

those in the related genera Callosamia Packard and Eupackardia Cockerell. However,

some adults of H. euryalus (Boisduval) on southern Vancouver Island, British

Columbia, were found to lack these white body markings. Controlled rearing indicated

that the “brown” phenotype is produced by a recessive allele at a single autosomal

locus, and examination of museum specimens showed that it is fairly common on

southern Vancouver Island and has been present there for at least half a century.

INTRODUCTION

Adults in the genus Hyalophora Duncan (Lepidoptera: Saturniidae) are large reddish

brown moths with white crescent-shaped discal spots, a white prothoracic collar, and white

segmental rings on the abdomen; these white markings are the key characters used to

separate this genus from the related genera Callosamia Packard and Eupackardia

Cockerell (Ferguson 1972; Lemaire 1978). In the course of other studies (Morewood

1991la, 1991b) an atypical adult phenotype, characterized by the absence of the white

prothoracic collar and abdominal rings (Fig. 1), was discovered in Hyalophora euryalus

(Boisduval) on southern Vancouver Island, British Columbia. This phenotype appears to

be unique in the genus Hyalophora and has not been reported previously; for example,

Tuskes et al. (1996) made no mention of the white prothoracic collar or abdominal rings in

their discussion of adult variation for any of the taxa within the genus. The objectives of

the current study were to document this adult colour pattern dimorphism, determine its

pattern of inheritance, and provide preliminary estimates of its prevalence and distribution.

MATERIALS AND METHODS

A small colony of H. euryalus was established from eggs laid by wild adult females

captured in Saanich, BC, one female on 16 May 1991 and one female on 6 May 1992 (Fig.

2). Larvae were reared indoors on cuttings of Douglas-fir, Pseudotsuga menziesii (Mirb.)

Franco ssp. menziesii, in large plastic buckets with screened lids. Pupae were overwintered

in small cages outdoors, and in the spring, adults were mated using two different methods.

In most cases mating cages constructed from coffee cans, as described by Miller and

Cooper (1976), were used to mate reared females to wild males to reduce inbreeding. In

some cases matings were obtained between reared adults by keeping them together in a

larger cage outdoors overnight. Each spring the phenotypes of all reared adults, as well as

wild males attracted by the caged females, were recorded as either “normal” (having a

white prothoracic collar and abdominal rings) or, for simplicity, “brown” (lacking the

“normal” white body markings such that the body appeared brown overall). Twenty-two

' Current Address: Department of Biological Sciences, Simon Fraser University, 8888

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J. ENTOMOL. SOC. BRIT. COLUMBIA 97, DECEMBER 2000

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J. ENTOMOL. SOC. BRIT. COLUMBIA 97, DECEMBER 2000 75

different broods were reared between 1991 and 1999, and the study was terminated when

in the summer of 1999 all larvae were lost to disease. Due to logistic constraints, only

small numbers of larvae were reared from most broods, making analysis of the distribution

of adult phenotypes within individual broods unlikely to yield meaningful results.

Therefore, broods representing crosses of apparently equivalent genotypes were grouped

together and the overall distribution of phenotypes for each type of cross was used to

evaluate the pattern of inheritance.

To provide a broader estimate of the prevalence and distribution of the brown

phenotype, all of the adult specimens of H. eurya/lus were examined in three major insect

collections in southwestern BC, namely those in the Spencer Entomological Museum at

the University of British Columbia in Vancouver and in the Pacific Forestry Centre and the

Royal British Columbia Museum in Victoria.

RESULTS

Adults were easily classified as having either the normal or the brown phenotype, with

no intermediate forms, and both phenotypes were common in both sexes (Figs. 1, 2). The

normal pattern of white body markings, in particular the location of the white prothoracic

collar, was often detectable as a faint greyish “smudge” in adults with the brown

phenotype; however, in no case was there any difficulty in assigning an individual to one

phenotype or the other. Crosses in which both parents were brown produced only brown

progeny (Brood numbers 3, 5, 9, 16, and 18 in Fig. 2 and Table 1) but crosses in which

both parents were normal had the potential to produce progeny of both phenotypes (Brood

number 4 in Fig. 2 and Table 1). Crosses in which one parent was brown and the other was

heterozygous normal (Brood numbers 2, 6, 7, 8, 10, 11, 12, 13, 14, and 20) produced

progeny in almost exactly the expected 1:1 ratio overall (Table 1), including both

phenotypes in progeny of each sex (Fig. 2).

1991

1992 O- :

i993 CHEO 24

1994 Ores C+

1995 Pog

1996 --OO

1997 C) i |

i908 Slo] OAD Ov ah ZEEE

1999 O See

Figure 2. Pedigree for the adults of Hyalophora euryalus reared during the course of this

study. Circles represent females, squares represent males; open symbols indicate the

“normal” phenotype, filled symbols indicate the “brown” phenotype. Numbers above

matings, and above the two founding females, are the “Brood numbers” in Table 1.

76 J. ENTOMOL. SOC. BRIT. COLUMBIA 97, DECEMBER 2000

Table 1

Observed and expected adult phenotypes of Hyalophora euryalus reared in Saanich,

British Columbia, between 1991 and 1999.

Type of cross Brood numbers’ Observed phenotypes — Expected phenotypes

(parental genotypes) (see Fig. 2) Normal Brown Normal Brown

Normal x Brown 2 0 a

(Bb x bb) 6 4 2,

fi D 4

8 1] 4

10 2 5

1] S 3

12 l 0

I3 5 7

14 6 4

20 6 8

Total: 40 44 42 42

Brown x Brown 3 0 22

(bb x bb) 5 0 5

9 0 is

16 0 31

18 0 4

Total: 0 qi) 0 77

Normal x Normal 4 4 2

(Bb x Bb) Total: 4 2 3 ]

| = ee en Ge i a ae en nr i i at ois. Se

Excluding broods 1, 15, 17, 19, 21, and 22, for which both parental genotypes could not

be conclusively determined.

The phenotypes of 28 wild males attracted by reared females caged in Saanich were

recorded during the course of this study; of these, seven (25%) had the brown phenotype.

A total of 40 adult specimens of H. euryalus were examined at the Spencer Entomological

Museum, approximately one-third of these being from southern Vancouver Island and the

adjacent Gulf Islands and the remainder from various localities in the southern interior of

BC as far north as Riske Creek and as far east as Cranbrook. Only three of these specimens

(7.5% of the total) had the brown phenotype, one from Nanaimo (1951), one from

Langford (no collection date), and one from Salmon Arm (1961). At the Pacific Forestry

Centre 18 adult specimens of H. euryalus were examined, over half of these being from

southern Vancouver Island and the remainder from the northern Okanagan region of the

southern interior, and no brown phenotypes were found. A total of 52 adult specimens of

H. euryalus were examined at the Royal British Columbia Museum, almost half being

from southern Vancouver Island and the Gulf Islands and the remainder of those with

locality data being from the southern interior of BC as far north as Williams Lake and as

far east as Cranbrook. Six of these specimens (11.5% of the total) had the brown

phenotype, two from Saanich (1959 and 1990), one from Nanoose (1995), one from

Galiano Island (1989), and two with no locality data (and no collection dates). Overall, of

50 museum specimens from southern Vancouver Island and the Gulf Islands, six (12%)

had the brown phenotype and of 40 museum specimens from the mainland (all but one

from the interior) of BC, only one (2.5%) had the brown phenotype (the remaining 20

museum specimens had no locality data).

J. ENTOMOL. SOC. BRIT. COLUMBIA 97, DECEMBER 2000 77

DISCUSSION

The overall distribution of phenotypes was consistent with a simple Mendelian pattern

of inheritance. Specifically, the presence or absence of the white prothoracic collar and

abdominal rings appears to be controlled by a single autosomal gene with two alleles, a

dominant allele (“B”) producing the normal phenotype and a recessive allele (“‘b’’)

producing the brown phenotype (Table 1). The distribution of phenotypes among the

progeny of crosses in which the parental female was normal and the parental male was

brown indicates that the brown phenotype is not sex-linked. Considering that the female is

the heterogametic sex in Lepidoptera, such crosses (Brood numbers 2, 10, and 12) should

produce only normal males and brown females if the trait is sex-linked, but that was not

the case (Fig. 2).

Other atypical phenotypes with similar patterns of inheritance have been reported

previously for various Lepidoptera in the context of biochemistry or evolution or both.

Waldbauer and Sternburg (1972) reported that a “blue” larval phenotype arose in their

laboratory colony of Hyalophora cecropia (L.) and was inherited as a simple autosomal

recessive. Based on the work of Clark (1971), they suggested that it was produced by a

mutation affecting the biochemical pathway that produces cuticular pigments from dietary

carotenoids. Stimson and Meyers (1984) reported that a “white” adult phenotype known

since 1965 in the Monarch butterfly, Danaus plexippus (L.) (Nymphalidae), in Hawaii is

inherited as a simple autosomal recessive. They presented evidence that it might be

increasing in frequency due to a lack of natural predators in Hawaii compared to North

America where predators would selectively remove adults lacking the normal aposematic

colour pattern. Adult females of the Eastern Tiger Swallowtail, Papilio glaucus L.

(Papilionidae), have either the black and yellow phenotype typical of the species or a

melanic phenotype that is thought to mimic the distasteful Pipevine Swallowtail, Battus

philenor (L.) (Papilionidae) (Brower and Brower 1962). Clarke and Sheppard (1962)

presented evidence that the melanic phenotype is completely sex-linked and Koch et al.

(1998) proposed a biochemical mechanism by which it could be controlled through a

single locus on the Y chromosome.

The museum specimens and wild individuals of H. euryalus observed during the

current study indicate that the brown phenotype has existed for at least half a century and

is fairly common on southern Vancouver Island, which suggests that the trait arose in this

region and that it offers some selective advantage or at least is not deleterious. The single

brown specimen from Salmon Arm indicates that this phenotype may also occur more

rarely in the interior of BC in Hyalophora “kasloensis” - a population of hybrid origin and

uncertain taxonomic standing (Tuskes et al. 1996; Collins 1997) which formerly was

considered to be a subspecies of H. euryalus (Morewood 1991a) - and suggests that there

might be gene flow between populations of Hyalophora in the interior and on the coast.

Among the colour patterns on the wings of various Lepidoptera, including many

Saturniidae, are well-developed “eyespots” that are thought to provide some protection

from vertebrate predators. The discal spots of Hyalophora can hardly be considered to

resemble typical eyes, however. The discal spots of H. euryalus are white, more-or-less

crescent-shaped, and much larger and more elongate on the hindwings than on the

forewings (Fig. 1). In the normal posture of living moths (as opposed to spread museum

specimens), the discal spots on the forewings and hindwings, respectively, of H. euryalus

might be imagined to resemble the narrowed eyes and bared canine teeth of a small feline

predator. Such a resemblance might provide some protection against predation by birds

and perhaps this resemblance is disrupted by the presence of the white body markings

typical of adults of Hyalophora.

78 J. ENTOMOL. SOC. BRIT. COLUMBIA 97, DECEMBER 2000

ACKNOWLEDGEMENTS

A number of people, most importantly Harry Morewood, assisted with rearing when other

commitments required my absence for long periods. Thanks to Karen Needham, Bob

Duncan, and Dave Blades for facilitating access to the specimens in the Spencer

Entomological Museum, the Pacific Forestry Centre, and the Royal British Columbia

Museum, respectively. Special thanks to John Cody and Daria C. Nutsch for the

suggestion that the elongated hindwing discal spots of H. euryalus resemble “fangs”.

REFERENCES

Brower, L.P. and J. VZ. Brower. 1962. The relative abundance of model and mimic butterflies in natural

populations of the Battus philenor mimicry complex. Ecology 43: 154-158.

Clark, R.M. 1971. Pigmentation of Hyalophora cecropia \arvae fed artificial diets containing carotenoid

additives. Journal of Insect Physiology 17: 1593-1598.

Clarke, C.A. and P.M. Sheppard. 1962. The genetics of the mimetic butterfly Papilio glaucus. Ecology 43:

159-161.

Collins, M.M. 1997. Hybridization and speciation in Hyalophora (Insecta: Lepidoptera: Saturniidae): A

reappraisal of W. R. Sweadneris classic study of a hybrid zone. Annals of Carnegie Museum 66: 411-

455.

Ferguson, D.C. in Dominick, R.B. et al. 1972. The Moths of America North of Mexico, fasc. 20.2B,

Bombycoidea (in part).

Koch, P.B., D.N. Keys, T. Rocheleau, K. Aronstein, M. Blackburn, S.B. Carroll and R.H. ffrench-

Constant. 1998. Regulation of dopa decarboxylase expression during colour pattern formation in wild-

type and melanic tiger swallowtail butterflies. Development 125: 2303-2313.

Lemaire, C. 1978. Les Attacidae Americains: The Attacidae of America (=Saturniidae). Attacinae. Edition

C. Lemaire, 42 boulevard Victor Hugo, Neuilly-sur-Seine, France.

Miller, T.A. and W.J. Cooper. 1976. Portable outdoor cages for mating female giant silkworm moths

(Saturniidae). Journal of the Lepidopterists’ Society 30: 95-104.

Morewood, W.D. 1991a. Larvae of Hyalophora euryalus kasloensis (Lepidoptera: Saturniidae). Journal of

the Entomological Society of British Columbia 88: 31-33.

Morewood, W.D. 1991b. Cold hardiness of Hyalophora euryalus kasloensis (Saturniidae) from the

Okanagan Valley, British Columbia. Journal of the Lepidopterists’ Society 45: 236-238.

Stimson, J. and L. Meyers. 1984. Inheritance and frequency of a color polymorphism in Danaus plexippus

(Lepidoptera: Danaidae) on Ohahu, Hawaii. Journal of Research on the Lepidoptera 23: 153-160.

Tuskes, P.M., J.P. Tuttle and M.M. Collins. 1996. The Wild Silk Moths of North America. Cornell

University Press, Ithaca, New York.

Waldbauer, G.P. and J.G. Sternburg. 1972. A mutant blue cecropia larva. Entomologia Experimentalis et

Applicata 15: 250-251.

J. ENTOMOL. SOC. BRIT. COLUMBIA 97, DECEMBER 2000 79

Aphid (Homoptera: Aphididae) accumulation and

distribution near fences designed for cabbage fly

(Diptera: Anthomyiidae) exclusion

M. K. BOMFORD

DIVISION OF PLANT AND SOIL SCIENCES, P.O. BOX 6108

WEST VIRGINIA UNIVERSITY, MORGANTOWN, WV USA 26506-6108

R. S. VERNON

PACIFIC AGRI-FOOD RESEARCH CENTRE, P.O. BOX 1000

AGASSIZ, BC, CANADA VOM 1A0

PEETER PATS

NORDIC ACADEMY FOR ADVANCED STUDY NEDRE VOLLGATE 8,

OSLO, NORWAY N-0158

ABSTRACT

Aphids accumulate near exclusion fences designed to intercept Delia radicum (L.)

movement into fields. Aphid accumulations increase with increasing fence height, but

are not affected by fence overhang length. Overall aphid levels are higher in small (4.3

m square) enclosed plots than in unenclosed plots. Enclosing large (38 m square) plots

does not alter overall aphid catches, but does alter aphid distribution within enclosures.

In large enclosures aphid accumulations are higher at enclosure perimeters than

interiors, with the highest accumulations near enclosure corners. This concentric

distribution is not observed in unfenced areas, and ts not altered by the addition of a

trap crop outside an enclosure.

Key words: Myzus persicae, Delia radicum, physical control

INTRODUCTION

The brassica pest Delia radicum (L.) (Diptera:Anthomyiidae) tends to fly close to the

ground (Vernon and MacKenzie 1998), where it can be intercepted by mesh exclusion

fences erected around brassica plantings to reduce crop damage (Vernon and MacKenzie

1998; Pats and Vernon 1999; Bomford et a/. 2000). In contrast, aphids (Homoptera:

Aphididae) commonly migrate at altitudes between 10 and 2,000 m (Isard et a/. 1990).

Exclusion fences are unlikely to intercept aphid movement due to aphids’ tendancy to

move close to the ground only when making short, local flights or preparing to alight.

Aphids are known to alight in areas where wind speeds are low, perhaps due to passive

deposition (Lewis and Dibley 1970), or active behaviour (Kennedy and Thomas 1973).

Since an exclusion fence may act as a partial windbreak, aphid accumulations may occur

inside the fence, which could be a concern to growers wishing to adopt this pest

management tool. This paper reports observations of aphid distribution inside fenced

enclosures during several experiments initially conducted to test D. radicum exclusion by

mesh fences.

80 J. ENTOMOL. SOC. BRIT. COLUMBIA 97, DECEMBER 2000

MATERIALS AND METHODS

Fence height study. Exclusion fences consisted of wooden frameworks covered with 1-

mm-mesh nylon window screening (Stollco Industries Ltd., Port Coquitlam, BC),

enclosing 5 m square plots. At the top of each fence, the vertical screen was bent over a

horizontal wooden sill (5 cm wide) along the top of the fence posts, to form a 22 cm

outward overhang, angled downward at 30-35° along triangular pieces of plywood secured

to the fence posts. There was no screen overhang projecting into the inside of the

enclosures. Fence heights (from ground level to the bottom of the outward overhang) of

30, 60, and 90 cm were tested. The open check plots had the same structure as the 30-cm

fence, but without the vertically oriented nylon screen. The trial was arranged as a four by

four Latin square, with adjacent blocks 4 m apart.

Fences were installed by 26 April 1991 in a SO by 55 m field located at Abbotsford,

BC. The field had been planted in raspberries for the 3 years previous, and had been kept

virtually weed free during the previous growing season. To prevent weed growth, soil on

the inside of the enclosures was covered with landscape fabric (Lumite 994, Division of

Synthetic Ind., Norcross, Georgia), and soil in a | m strip centred along the fence

perimeter was covered with black plastic. On 29 April, 1991, twenty 2-week-old

rutabagas, Brassica campestris var. napobrassica (L.) ‘Laurentian,’ were transplanted into

the plots along each of five parallel rows cut into the landscape fabric. Exposed soil around

rutabagas was weeded weekly.

On 4 July 1991, counts of aphids, aphids parasitized by aphidiid wasps (Hymenoptera:

Aphidiidae), and syrphid fly larvae (Diptera: Syrphidae) were recorded for 15-23 rutabaga

leaves from each of the five crop rows of each plot. The mean number of insects per leaf

was calculated for each insect in each treatment. Data were analyzed by ANOVA, and

treatment means were separated using Tukey’s test.

Standard fence design. A modified version of the fence used in the previous study

was used in all remaining studies. Aluminum framed window screens of | mm black nylon

mesh (210 cm long by 120 cm high) (Stollco Industries, Port Coquitlam, BC) were

supported between wooden fence posts (7.5 cm by 9 cm wide by 120 cm high) to form

vertical panels. At the top of each panel a wooden fence top (2 cm high by 8 cm wide by

210 cm long) rested on the top edge of the aluminum frame. From this wooden top,

separate strips of 1-mm-mesh nylon screen were attached to form collection overhangs of

specified lengths angled downward at 45° on both sides of the fence, and held in place by

plywood triangles attached to the tops of the fence posts. All exposed fence components

were black.

Sticky trap design. Sticky traps were used to monitor winged aphid populations in all

remaining studies. Traps were made from sheets of white cardboard (4-ply Railroad Board;

Domtar Fine Papers, Toronto, ON) painted on both sides with yellow, semigloss enamel

paint (Yellow 776, Cloverdale Paint and Chemicals, Surrey, BC), cut into 10 by 14 cm

rectangles and dipped in a commercial insect adhesive (Stiky Stuff, Olson Products,

Medina, OH). Traps were attached to wooden stakes, with the bottom edge (14 cm long)

15 cm above the ground, and were oriented to face north-south.

Overhang length studies. The experimental site was a regularly mowed field of mixed

grass near Abbotsford, BC. The trials were arranged in a randomized complete block

design with four replicates, 30 m apart. Each replicate contained three 7 x 7 m square

treatment plots, 10 m apart, covered with black woven landscape fabric to prevent the

growth of weeds. The three treatment plots in each block were as follows: (1) an unfenced

control plot, (2) a plot enclosed by a fence with a 25-cm-long collection overhang, and (3)

a plot enclosed by a fence without an overhang (trial 1: 13 July — 10 August, 1994), a

fence with a 12.5 cm collection overhang (trial 2: 12-30 August, 1994) or a fence with a 50

cm collection overhang (trial 3: 23 August — 14 September, 1995). The positions of plots

J. ENTOMOL. SOC. BRIT. COLUMBIA 97, DECEMBER 2000 8]

in each block were randomized at the start of each trial. Fences enclosed a 4.3 x 4.3 m

square in the centre of each plot.

At the beginning of each trial, black plastic flats of 50 6- to 15-d-old radish, Raphanus

sativus (L.) ‘Cavalrondo,’ seedlings were evenly spaced throughout a 3.5 x 3.5 m square in

the centre of each plot. Radishes were watered daily for the duration of each trial.

Winged aphid catches on sticky traps placed 1.5 m north-east and south-west of the

center of each plot were recorded every 2-6 d throughout each trial, following trap

replacement. Data were transformed (square root (x + 0.5)) to correct for heterogeneity of

variance. For each trial, mean aphids per trap were calculated for each 2-6 d trapping

session for each treatment, and the effect of treatment and block on mean aphids per trap

for each 2-6 d trapping session was tested by ANOVA and means separated using Tukey’s

test (Zar 1984). Data from all 2-6 d trapping sessions in each trial were then pooled, the

effect of treatment and block on mean aphids per trap tested by ANOVA, and means

separated using Tukey’s test.

Concentric enclosure study. A 41 x 41 m square in a regularly mowed field of mixed

grass near Abbotsford, BC was covered with black landscape fabric to prevent weed

invasion, and to provide a uniform environment throughout the experimental area. Four

concentric enclosures were constructed in the centre of the experimental area using

standard exclusion fences. The innermost enclosure was a 4.5 x 4.5 m square; the next a

13.5 x 13.5 m square; the next a 22.5 x 22.5 m square; and the outermost a 31.5 x 31.5 m

square.

On 23 June 1994, 324 flats of 50 7-d-old radish seedlings were arranged in a 1 m grid

(18 rows and columns) throughout the experimental area. Eighty-one sticky traps were

arranged throughout the experimental area in a 9 row and 9 column grid, with 4.5 m

between consecutive traps. All traps were replaced at 3-7 (mean 5) d intervals, until 17

August 1994 — a total of 10 trapping sessions. Winged aphid catches on each trap were

recorded for each trapping session.

Traps were grouped into one of five levels, according to their location (Table 1). Mean

aphid catches for each level were calculated and ranked for each trapping session.

Trapping sessions were treated as replicates in time. Friedman’s test (Zar 1984) was used

to test the null hypothesis that mean aphid catches were equivalent for each level; rankings

were separated using a variation of Tukey’s test for multiple comparisons of

nonparametric data (Zar 1984). Cumulative aphid distribution throughout the experimental

area was mapped using 3-dimensional graphing software (MSGraph 8.0, Microsoft 1997).

Table 1

Trap locations by level in concentric enclosure study.

Distance from outer Trap location in relation to fenced

Level fence (m) Traps _enclosure(s)

] +250 52 Outside 31.5 m enclosure

2 -2.50 24 Between 22.5 and 31.5 m enclosures

3) -6.75 16 Between 13.5 and 22.5 m enclosures

4 -11.25 8 Between 4.5 and 13.5 m enclosures

5 -15.75 l Inside 4.5 m enclosure

Large enclosure studies. Three 38 x 38 m squares in a regularly mowed field of mixed

grass near Abbotsford, BC were covered with black landscape fabric. Treatment areas

were arranged in a line oriented roughly perpendicular to the main southwest wind

direction, with adjacent plots ~20 m apart. Sticky traps and flats of 10-d-old radishes were

evenly spaced throughout each experimental area, according to the design of the previous

experiment.

82 J. ENTOMOL. SOC. BRIT. COLUMBIA 97, DECEMBER 2000

Experimental areas were randomly assigned to one of three treatments: (1) no fence

(Control); (2) a 38 by 38 m square standard fence, enclosing all of the radish plants

(Fence); and (3) a 30 by 30 m square standard fence, with radish plants also encircling the

fence to act as a trap crop (Fence + Trap crop). Due to the large size of the treatment plots,

replication was conducted over time in 1995. The treatments were initially established on

29 May 1995, and were subsequently re-randomized on three additional occasions (11

July, 15 August, 12 September) to allow four replicated blocks. Traps were changed every

3-7 d (mean, 4 d) for a period of 21-28 d (mean, 24 d). Cumulative aphid distributions for

each treatment were mapped using 3-dimensional graphing software (MSGraph 8.0,

Microsoft 1997). ANOVA was used to test for effects of treatment and block (time) on

overall aphid catches. The average aphid catch for each enclosure treatment and block

(time) on sticky traps immediately inside the enclosure was compared to that on the

remaining traps using the Wilkoxon paired-sample test (Zar 1984).

RESULTS

All studies. Aphids caught in all studies were predominantly Myzus persicae (Sulzer),

but the proportion was not quantified. Total aphid catches varied considerably from one

trapping session to another, following no apparent trends. Differences between treatments

tended to be most pronounced for trapping sessions with relatively high aphid catches.

Fence height study. A total of 3,637 aphids were found on the 1,244 leaves sampled.

More aphids were found in plots enclosed by 90-cm-high fences than in plots without

fences or plots with 30-cm-high fences (P=0.008) (Fig. 1A). Aphid accumulations

increased with increasing fence height over the range of fence heights tested, but were not

significantly higher inside plots surrounded by 30- and 60-cm-high fences than in

unfenced control plots (Fig. 1A). No block effect was detected.

A total of 127 aphids were parasitized by aphidiid wasps and 35 syrphid fly larvae

were found on the leaves sampled. Both of these aphid biocontrols were more numerous

inside plots enclosed by 90-cm-high fences than in other experimental plots (P=0.007,

aphidius; P=0.002, syrphid) (Fig. 1A,B). No block effect was detected.

Overhang length studies. A total of 18,526 winged aphids were caught on sticky traps

over the course of the three overhang length trials. Significant (P<0.05) treatment, block,

and trapping date effects were detected in all trials. A significant interaction between

treatment and trapping date was attributed to a positive correlation between total aphid

catch and strength of the treatment effect. More winged aphids were caught on sticky traps

inside the fenced enclosures than in unfenced check plots in each of the trials (Table 2).

The presence of overhangs, and overhang length had no effect on aphid catches (Table 2).

Concentric enclosure study. A total of 37,894 winged aphids were caught on sticky

traps over the course of this study, averaging 468 aphids per trap and 46.8 aphids per trap

for each trapping session. Aphid catches varied tremendously between trapping sessions,

ranging from 0.6 aphids per trap on 5 July, to 237.0 aphids per trap on 11 August.

Trap location had a significant (P<0.001) effect on aphid catches, with traps within the

outer two enclosures (levels 2 and 3) catching the most aphids (Table 3; Fig. 2). Traps in

level 2 caught more aphids than those in levels 1, 4, or 5; traps in level 3 caught more than

those in level 4 (Table 3). Aphid catches peaked near the inner corners of the largest

enclosure (Fig. 2). Catches were below average around the outer perimeters and towards

the center of the study area (Fig. 2).

Large enclosure study. A total of 25,419 aphids were caught throughout this study,

averaging 26 aphids per trap for each block (time). The mean aphid catches (+ SE), were

2006 + 1098, 2085 + 802, and 2264 + 1148 in the Check, Fence, and Fence + Trap Crop

treatments, respectively. No significant difference in mean aphid catches were detected

J. ENTOMOL. SOC. BRIT. COLUMBIA 97, DECEMBER 2000

between treatments. The block effect was highly significant (P<0.0001), indicating

variation in aphid catches over time.

8.0

A: Aphid

6.0

4.0

2:0

0.0

0.4

0.3

0.2

Insects per leaf (x + S.E.)

C: Syrphid

0 30

Enclosure height (cm)

Figure 1. Counts of living aphids (A), aphids parasitized by aphidiid wasps (B), and

syrphid larvae (C) on leaves of rutabagas growing inside exclusion fences ranging from 0-

90 cm in height (n=4). Bars with the same letter are not significantly different, Tukey’s

test, P<0.05.

Table 2

Average aphid catch on sticky traps in unfenced plots of radish and plots of radish

enclosed by 120-cm-high exclusion fences with varying overhang lengths.

Mean aphid catch by trial, n=4°

Trial | Thiai2 Trial 3

Treatment (13/7/94 (12/8/94 (23/8/95

- 10/8/94) -30/8/94) -14/9/95)

No fence 97.9 a DA ORIG: 154.0 a

Fence without overhang 364.7 b - -

Fence with 12.5 cm overhang - 930.20 -

Fence with 25 cm overhang 494.2 b 923.9) T1395

Fence with 50 cm overhang - - 658.1 5

“Means within a column followed by the same letter are not significantly different,

Tukey’s test, P<0.05.

84 J. ENTOMOL. SOC. BRIT. COLUMBIA 97, DECEMBER 2000

Aphid catches were above average near the outer edges of fenced enclosures

particularly near the enclosure corners (Fig. 3B,C). This pattern was not observed in the

unfenced control plots (Fig. 3A). Aphid catches on traps immediately inside the enclosure

fences were higher (P<0.001) than for the remaining traps inside both the Fence and Fence

+ Trap Crop treatments. This concentric distribution was not observed in the control plots.

Table 3

Winged aphid catches, by level, in five levels of a concentric enclosure study. Rank sum is

the sum of aphid catch ranking for each of 10 trapping sessions, according to Friedman’s

analysis of variance by ranks.

Level Total Aphids per trap per session, Rank sum, n=10°*

traps n=10.(x. £S.E.)

1 (outer traps) 32 36.8 + 18.2 265.45

2 24 60.5 #:30.5 45.0 c

3 16 5004-2417 40.0 bc

4 8 40.3+21.4 20°5°4

5 (center trap) l AS 3 ce LOK 23.0 ab

“Rank sums followed by the same letter are not significantly different, Tukey’s test,

P<0.05.

Figure 2. Contour map showing aphid distribution in an experimental area with concentric

exclusion fences (heavy lines). Sticky traps were placed at grid nodes. Contour lines show

total aphid catch per trap after 10 trapping sessions (x = 468), at intervals of 50, and are

labeled at intervals of 150. Areas with total catches below 450 aphids per trap

(approximate average) are shaded gray. One square = 4.5 by 4.5 m.

J. ENTOMOL. SOC. BRIT. COLUMBIA 97, DECEMBER 2000

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86 J. ENTOMOL. SOC. BRIT. COLUMBIA 97, DECEMBER 2000

DISCUSSION

We conclude that exclusion fences trigger aphid accumulations near enclosure

perimeters. Inside small enclosures, where all enclosed areas are relatively near the

enclosure perimeter, exclusion fences increase aphid numbers overall. In large enclosures

higher aphid catches near enclosure edges are counterbalanced by comparatively low aphid

catches far from enclosure edges, resulting in an altered aphid distribution within

enclosures, but no overall change in aphid catches.

Aphid accumulations inside exclusion fences are affected by fence height, but not

overhang length. In small enclosures 30 - 90-cm-high, aphid accumulations increase with

fence height. Lewis and Dibley (1970) hypothesized that small insects, such as aphids, are

passively carried by wind currents, and deposited in the lee of windbreaks by swirling

eddies, which are larger for taller windbreaks, assuming constant windbreak permeability.

Kennedy and Thomas (1973) agreed that aphids accumulate in areas where windspeeds are

low, but argued that this was an effect of aphid behaviour rather than passive deposition.

Whether due to active behaviour or passive deposition, both authors agree that aphid

accumulations will be highest where windspeed is reduced, as we observed near exclusion

fences.

At our study locations the prevailing daytime wind blew from the southwest. Prevailing

night winds blew from the northeast. The regular reversals in the local prevailing wind

direction made it difficult to establish any relationships between the location of aphid

accumulations within enclosures and wind direction, particularly since we made no

observations of the time of day when winged aphids were caught.

Our observation that overhang length has no effect on aphid accumulations conflicts

with the finding that overhangs reduce cabbage fly movement into fenced enclosures

(Bomford ef al. 2000). This may be because exclusion fences intercept the low-flying

cabbage flies, but not aphids, which maintain a higher altitude before alighting. Overhangs

will only reduce insect movement into enclosures if insects fly into the exclusion fence,

then encounter the overhang as they attempt to move up and over the fence.

More syrphid fly larvae, which feed on aphids, and aphids parasitized by aphidiid

wasps were found inside 60-cm-high fences than in contro] plots. These insects may have

been attracted to the higher concentrations of their aphid hosts within the small enclosures,

or they may accumulate in the same low windspeed areas where aphids tend to alight. The

fact that the exclusion fences did not reduce immigration of these predators and parasites

suggests that this physical control tactic could compliment efforts to use these beneficial

insects for the biological control of aphids.

The highest aphid accumulations in large enclosures occurred near enclosure corners.

Corner traps likely catch aphids moving from two directions, whereas traps near the

middle of an edge likely catch only aphids coming from one direction. Traps placed inside

small enclosures catch aphids coming from all directions, resulting in the marked increase

in aphid catches observed in small enclosures relative to control plots.

Positioning an exclusion fence between a perimeter trap crop and the main crop had no

effect on overall aphid accumulations, as compared to control plots without a fence, or

plots entirely enclosed by a fence. Plot size was held constant in these experiments, such

that allowing room for a trap crop required a reduced enclosure size. The area of reduced

aphid accumulations in the interior of the Fence + Trap Crop plots was correspondingly

smaller than the area of reduced aphid accumulations in the fully enclosed plots.

In our concentric enclosure study all traps were the same distance from a mesh fence,

yet traps towards the outer edge of the study areas caught more aphids than traps towards

the center of the study area. This was the same distribution pattern observed in our large

J. ENTOMOL. SOC. BRIT. COLUMBIA 97, DECEMBER 2000 87

enclosure study, suggesting that local aphid distributions are better predicted by the

distance from the outer edge of an enclosed area than distance from a fence.

Extrapolating from our observations in experimental plots, we would expect aphids to

accumulate near the outer edges of fields enclosed by exclusion fences. By comparison,

fields without exclusion fences should have similar overall aphid levels, but aphids will be

more randomly distributed throughout the field area. The more predictable aphid

distribution within fields surrounded by exclusion fences could allow producers to target

field edges for insecticide applications intended for aphid control, reducing control costs,

insecticide use, and soil compaction, while preserving an area of refuge for biological

control organisms in field interiors.

ACKNOWLEDGEMENTS

Thanks to Amanda Bates and Sherah van Laerhoven for help with field work. This

study was supported by grants from Energy, Mines and Resources (PERD program)

(MKB, RSV), and by the Wenner-Gren Foundations, Sweden (PP). Contribution No. 644

of the Pacific Agri-Food Research Centre, Agassiz, British Columbia, Canada VOM 1A0.

REFERENCES

Bomford, M.K., Vernon, R.S. and Pats. P. 2000. Importance of collection overhangs on the efficacy of

exclusion fences for managing cabbage flies (Diptera: Anthomyiidae). Environmental Entomology 29:

795-799.

Isard, S.A., Irwin, M.E. and Hollinger, S.E. 1990. Vertical distribution of aphids in the planetary boundary

layer. Environmental Entomology 19: 1473-1484.

Kennedy, J.S. and Thomas, A.A.G. 1973. Behaviour of some low-flying aphids in wind. Annals of

Applied Biology 76: 143-159.

Lewis, T. and Dibley, G.C. 1970. Air movement near windbreaks and a hypothesis of the mechanism of

the accumulation of airborne insects. Annals of Applied Biology 66: 477-484.

Microsoft. 1997. Microsoft Graph 8.0. Microsoft Corp. Redmond, WA.

Pats, P. and Vernon, R.S. 1999. Fences excluding cabbage maggot flies and tiger flies (Diptera:

Anthomyiidae) from large plantings of radish. Environmental Entomology 28: 1124-1129.

Vernon, R.S. and MacKenzie, J.R. 1998. The exclusion fence: A novel tool for Anthomyiid fly

management. The Canadian Entomologist 130:153-162.

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

‘

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J. ENTOMOL. SOC. BRIT. COLUMBIA 97, DECEMBER 2000 89

Parasitism of the eggs of Lygus shulli and

Lygus elisus (Heteroptera: Miridae) by Anaphes tole

(Hymenoptera: Mymaridae)

ROBERT R. McGREGOR’, DAVID R. GILLESPIE,

DONALD M.J. QUIRING and DAWN HIGGINSON

PACIFIC AGRI-FOOD RESEARCH CENTRE, AGRICULTURE and AGRI-FOOD

CANADA, P.O. BOX 1000, AGASSIZ, BC, VOM 1A0

ABSTRACT

Females of the egg parasitoid Anaphes iole Girault (Hymenoptera: Myrmaridae)

accepted and oviposited in eggs of both Lygus shulli Knight and L. elisus Van Duzee

(Heteroptera: Miridae) when presented on sections of green bean pod in the laboratory.

Resulting A. iole larvae developed normally on eggs of both host species and emerged

as adults. The wings of 4. iole emerging from L. shulli eggs were significantly larger

than those from L. elisus probably because the eggs of L. shulli were larger. Anaphes

iole females parasitized only approximately 50% of the eggs available of either host

species. This may indicate that 50% of the hosts were suitable and rejected, that 50%

were unsuitable for oviposition, or that the structure of bean pods prevents females

from finding or ovipositing in 50% of hosts. Anaphes iole has potential for biological

control of Lygus spp. on greenhouse vegetables in southwestern British Columbia.

Key words: Anaphes iole, Mymaridae, Lygus shulli, Lygus elisus, Lygus hesperus,

Miridae, egg parasitoids, biological control, greenhouse vegetables

INTRODUCTION

Plant bugs in the genus Lygus (Heteroptera: Miridae) feed on a wide variety of native

plant species and agricultural crops throughout North America (Schwartz and Footit 1992;

Schwartz and Footit 1998). Sequential migration among different seasonally-occurring

host-plant species is typical of the biology of Lygus (Schwartz and Footit 1992). As a

consequence, Lygus species migrate into, and feed upon, acceptable agricultural crops.

Economic damage occurs when Lygus nymphs or adults feed on reproductive tissues or

apical meristems (Schwartz and Footit 1992). Damage has been reported on cotton (Leigh

et al. 1988), alfalfa (Sorenson 1936), canola (Butts and Lamb 1991; Wise and Lamb 1998),

strawberries (Norton and Welter 1996) and conifer seedlings (Schowalter 1987). In

southwestern British Columbia (BC), Lygus bugs are sporadically-occurring pests of

greenhouse vegetable crops like cucumber and sweet pepper (Gillespie ef a/. 1999). Here,

we report on parasitism of the eggs of L. shulli Knight and L. elisus Van Duzee by the egg

parasitoid Anaphes iole Girault (Hymenoptera: Mymaridae), and discuss the potential of A.

iole for biological control of Lygus species in BC vegetable greenhouses.

Lygus hesperus Knight, L. shulli and L. elisus are the three most common species of

Lygus in the Fraser Valley of southwestern BC (Gillespie et al. 1999). Lygus shulli and L.

elisus are sporadically found in vegetable greenhouses throughout the growing season, and

their feeding damages both cucumbers and peppers (Gillespie et a/. 1999). Lygus hesperus

occurs only on late-season pepper crops, and there is no definitive evidence, at present, that

Department of Biology, Douglas College, P.O. Box 2503, New Westminster, BC, V3L

5B2, Canada

90 J. ENTOMOL. SOC. BRIT. COLUMBIA 97, DECEMBER 2000

this species causes economic damage (Gillespie et al. 1999). Invasion of greenhouses

probably occurs when Lygus bugs disperse from surrounding weedy habitat (Gillespie et

al. 1999).

Augmentative releases of the egg parasitoid Anaphes iole have been used to manage

populations of L. hesperus in field plantings of strawberries in California (Norton ef al.

1992; Norton and Welter 1996; Udayagiri and Welter 2000). Although A. iole is available

commercially (Biotactics Inc., Riverside, California) and could potentially be used for

biological control of Lygus spp. in BC, no information is currently available regarding

parasitism of L. shulli and L. elisus by this parasitoid. In this paper, we report on the

acceptance and suitability of L. shulli and L. elisus eggs as hosts for A. iole.

MATERIALS AND METHODS

Host eggs for parasitism bioassays. Adults of L. shulli and L. elisus were collected at

field sites near the Pacific Agri-Food Research Centre in Agassiz, BC in July 1999. Lygus

adults were identified to species using Henry and Froeschner (1988). Adults of each

species were held at ambient conditions in the laboratory in groups of 5-10 in cylindrical

500-ml plastic containers with screened lids. Lygus adults were provided with 5-10

sections of green bean (3 to 5 cm in length) for feeding and oviposition. Bean sections

were inspected daily for Lygus eggs and those with two or more eggs present were

removed from the containers and stored in a refrigerator at 10°C until used in parasitism

bioassays. Bean sections were replaced in oviposition containers when removed (as above)

or when their condition deteriorated (e.g. showing signs of fungal growth).

Parasitism bioassays. Adult Anaphes iole of unknown age were obtained directly

from a commercial supplier (Biotactics Inc., Riverside, California). Female A. io/e were

selected for use in oviposition bioassays by examining abdominal morphology under a

dissecting microscope to determine sex. Individual female A. io/e were introduced using a

moistened fine paintbrush into 62-ml plastic cups (Solo Cup No. P200, Solo Cup

Company, Urbana, Illinois) each containing a single section of green bean with eggs of

either L. shulli (91 cups) or L. elisus (82 cups). The number of eggs available for

parasitism on each bean section ranged from 2 to 36 for L. elisus and from 2 to 68 for L.

shulli. The cups were held in a growth chamber for 24 h to allow A. jole to oviposit (L:D

16:8 h photoperiod and temperatures of 23.0 + 0.2°C (+SD) during photophase and 18.0 +

0.2°C (+SD) during scotophase; relative humidity was not controlled). After 24 h, the

female A. iole were removed with a moistened paintbrush and the cups were returned to

the growth chamber and checked every second day for emergence of Lygus nymphs or A.

iole adults until no further emergence occurred. Emerging Lygus nymphs were removed,

counted and discarded. Emerging A. jo/e adults in each cup were removed and stored

separately, as families, in 70% (w/v) ethanol. The total numbers of Lygus nymphs and 4A.

iole males and females that emerged from each bean section were recorded.

Wing size of A. iole adults. The right wing of A. io/e adults that emerged from bean

sections was measured as an index of body size. From the offspring stored in ethanol, we

selected at random, one male and one female adult from each of 32 randomly-selected

families reared on L. shulli and 31-32 families reared on L. elisus. The length (from the

point of attachment to the tip of the wing) and width (at the widest point) of each right

forewing were measured and recorded.

Size of Lygus eggs. Adults of L. shulli and L. elisus were collected at field sites near

the Pacific Agri-Food Research Centre at Agassiz, BC in July of 2000. The ovaries of 10

adult females of each species were dissected. The length (from tip to tip) and width (at the

widest point) of all mature, fully chorionated eggs for each female were measured and

recorded.

J. ENTOMOL. SOC. BRIT. COLUMBIA 97, DECEMBER 2000 9]

Data analysis. Total number of viable host eggs in parasitism bioassays was calculated

as the totals of Lygus and A. io/e emerging from each bean section. This calculation gives a

conservative estimate of the original number of Lygus eggs per bean section as it assumes

that no Lygus or Anaphes eggs died during the test. The proportion of eggs parasitized per

female was calculated as the number of A. io/e emerging divided by the total number of

host eggs. The proportion of female 4. io/e emerging from each bean section (sex ratio)

was calculated as the number of females emerging divided by the total of both sexes

(F/(F+M)). Proportion of Lygus eggs parasitized, number of A. iole emergents, number of

Lygus emergents, total number of host eggs and sex ratio of A. iole emergents (all per bean

section) were compared between L. shulli and L. elisus using Mann-Whitney tests.

Proportional data were arcsin-square root transformed before analysis. The length and

width of the wings of male and female wasps emerging from L. shulli and L. elisus were

compared using Mann-Whitney tests. The length and width of the eggs of L. shulli and L.

elisus were compared using Mann-Whitney tests. All statistical analyses were conducted

using Systat (Wilkinson ef a/, 1997) and Sigmastat (Fox ef al. 1995).

RESULTS

Parasitism bioassays. Female A. io/e oviposited in eggs of both Lygus species, and

both species were suitable for the development of A. jo/e from larva to adult. There was no

significant difference in the proportion of eggs parasitized per A. io/e female for either L.

elisus or L. shulli (Table 1). The number of A. jole and Lygus emerging and the total

number of host eggs per bean section were significantly higher for L. shulli than for L.

elisus (Table 1). Previous observations indicate that L. shu/li is more likely to oviposit on

green beans than L. e/isus (D.M.J. Quiring, personal observation). There was no significant

difference in the sex ratio of A. iole emerging from L. shu/li compared to L. elisus eggs

(Table 1).

Table 1

Proportion of Lygus eggs parasitized, number of Anaphes and Lygus emerging, number of

Lygus host eggs, and sex ratio (Means + SE) of A. jole emerging per bean section

containing host eggs of L. shulli and L. elisus.

Variable L. shulli L. elisus Mann-Whitney test

U P

Proportion of eggs 0.46+0.04 0.49+0.04 3052 0.50

parasitized (n=9T) (n=82)

Number of Anaphes 10.4+1.1 5.1+0.5 2986 0.01

emerging (n=91) (n=82)

Number of Lygus 10.9+ 1.0 6.0 + 0.6 2549 < 0.001

emerging (n=91) (n=82)

Total number of host ore eS 11.1+40.8 1791 =02001

eggs (i911) (n=82)

Sex ratio 0:55-0:03). °0,52.40.04 199] 0.48

(F/(F+M)) (n=64) (n=67)

Wing size of A. iole adults. The mean length and width of the right forewings of both

male and female A. io/e emerging from the eggs of L. shulli were significantly greater than

those emerging from the eggs of L. elisus (Table 2).

92 J. ENTOMOL. SOC. BRIT. COLUMBIA 97, DECEMBER 2000

Table 2

Length and width (Mean + SE) of right forewings of Anaphes iole males and females

reared on the eggs of Lygus shulli and Lygus elisus.

Variable L. shulli L. elisus Mann-Whitney test

rm in mE MEMM MN

Female wing length 0.75+0.01 O72 0.01 204 < 0.001

(mm) (n=32) (n=32)

Female wing width 0.17+<0.005 0.15+< 0.005 202 < 0.001

(mm) (n=32) (n=32)

Male wing length 0.79+<0.005 0.75+0.01 I33 < 0.001

(mm) (n=32) (n=31)

Male wing width 0.18+<0.005 0.17+<0.005 343 0.033

(mm) (n=32) (n=31)

Size of Lygus eggs. The mean length of L. shulli eggs (0.99 + 0.01 mm) was

significantly greater than that of L. e/isus eggs (0.91 + 0.01 mm; Mann Whitney test: U =

132, P<0.001), as was the mean width of L. shulli eggs (0.26 + 0.00 mm) compared to L.

elisus eggs (0.23 + 0.00 mm; Mann Whitney test: U = 224, P<0.001).

DISCUSSSION

Anaphes iole females readily accepted, and oviposited in, eggs of both L. shulli and L.

elisus. Their offspring developed successfully in both species of host eggs and emerged as

adults. Anaphes iole are available commercially and readily parasitize all three Lygus spp.

(L. shulli, L. elisus, and L. hesperus) found in greenhouses in the Fraser Valley of

southwestern BC. Anaphes iole thus has a strong potential for biological control of Lygus

spp. in BC vegetable greenhouses.

No significant differences were found between the two host species in the proportion of

available eggs parasitized by A. iole females. The proportion of eggs parasitized was

approximately 50% for both species despite the fact that, on average, nearly twice the

number of host eggs were oviposited on bean sections by L. shulli compared with L. elisus.

Anaphes iole females typically carry between 30 and 40 mature eggs in their abdomens,

and an individual female can parasitize approximately 30 Lygus eggs per day (S.

Udayagiri, University of California, personal communication). Assuming that no A. jole

eggs died after oviposition in our experiments, the 4. io/e females in this study oviposited

substantially fewer eggs (on either host species) during 24 h than they had available (5

eggs on average on bean sections with L. e/isus eggs and 10 eggs on average on those with

L. shulli eggs). It is possible that 50% of the eggs of both species were either suitable hosts

that were rejected by females, or unacceptable hosts for oviposition. Alternatively, 50% of

Lygus eggs could have been inaccessible for oviposition, or impossible to locate, by A. iole

on bean sections. Recently, it has been shown that plant structure can influence the

oviposition success of A. iole. A lower proportion of Lygus eggs were parasitized by A.

iole females on strawberry fruits than on petioles, leaflets or calyx tissue (Udayagiri and

Welter 2000). Achenes (one-seeded fruitlets) on strawberry fruits apparently hinder access

by A. iole females to Lygus eggs present on fruits (Udayagiri and Welter 2000). If A. ole is

to be used for biological control of Lygus spp. in BC greenhouses, it will be critical to

determine whether plant structure affects the ability of A. iole females to locate and

parasitize eggs on cucumbers and peppers.

Anaphes iole adults that emerged from L. shulli eggs had larger wings than adults from

L. elisus eggs. Assuming that wing size correlates with body size, L. shulli eggs may be of

higher quality for development of A. iole than L. elisus eggs. This is likely a consequence

J. ENTOMOL. SOC. BRIT. COLUMBIA 97, DECEMBER 2000 93

of the larger size of L. shulli eggs compared to L. elisus eggs. Alternatively, the difference

in wing size we observed may be caused by some influence of the host not related to egg

size. Host-induced variation in antennal morphology unrelated to host size was found in A.

iole by Huber and Rajakulendran (1988). Determining whether wing-size variation

between hosts reflects differences in host-egg size or other effects will require further

research.

ACKNOWLEDGEMENTS

We thank the British Columbia Greenhouse Research Council, the Investment

Agriculture Foundation of British Columbia, and Agriculture and Agriculture and Agri-

Food Canada Matching Investment Initiative for financial assistance. This is contribution #

639 from the Pacific Agri-Food Research Centre (Agassiz).

REFERENCES

Butts, R.A and R.J. Lamb. 1991. Pest status of Lygus bugs (Heteroptera: Miridae) in oilseed Brassica

crops. Journal of Economic Entomology 84: 1591-1596.

Fox, E., Shotton, K. and Ulrich, C. 1995. SigmaStat Statistical Software, Version 2.0. Jandel Corporation,

Chicago, Illinois.

Gillespie, D.R., R.G. Footit, L. Shipp, M.D. Schwartz, K. Wang and D.M.J. Quiring. 1999. Lygus bugs in

protected crops — improving our understanding pp. 1-8. In: R.G. Footit and P.G. Mason (Eds.)

Proceedings of the Lygus working group meeting. Agriculture and Agri-Food Canada, Research

Branch. 54pp.

Henry, T.J. and R.C. Froeschner. 1988. Catalog of the Heteroptera or true bugs of Canada and continental

United States. St. Lucie Press, New York, N.Y.

Huber, J.T. and S.V. Rajakulendran. 1988. Redescription of and host-induced antennal variation in

Anaphes iole Girault (Hymenoptera: Mymaridae), an egg parasite of Miridae (Hemiptera) in North

America. The Canadian Entomologist 120: 893-901.

Leigh, T.F., T.A. Kerby and P.F. Wynholds. 1988. Cotton square damage by the plant bug, Lygus hesperus

(Hemiptera: Heteroptera: Miridae), and abscission rates. Journal of Economic Entomology 81: 1328-

leer

Norton, A.P. and S.C. Welter. 1996. Augmentation of the egg parasitoid Anaphes iole (Hymenoptera:

Mymaridae) for Lygus hesperus (Heteroptera: Miridae) management in strawberries. Environmental

Entomology 25: 1406-1414.

Norton, A.P., S.C. Welter, J.L. Flexner, C.G. Jackson, J.W. Debolt and C. Pickel. 1992. Parasitism of

Lygus hesperus (Miridae) by Anaphes iole (Mymaridae) and Leiophron uniformis (Braconidae) in

California strawberry. Biological Control 2: 131-137.

Schowalter, T.D. 1987. Abundance and distribution of Lygus hesperus (Heteroptera: Miridae) in two

conifer nurseries in Western Oregon. Environmental Entomology 16: 687-690.

Schwartz, M.D., and R.G. Footit. 1992. Lygus bugs on the prairies: biology, systematics and distribution.

Agriculture:and Agri-Food Canada, Technical Bulletin 1992-4E. 44pp.

Schwartz, M.D., and R.G. Footit. 1998. Revision of the nearctic species of the genus Lygus Hahn, with a

review of the palaearctic species (Heteroptera: Miridae). Memoirs on Entomology International,

Volume 10. Associated Publishers.

Sorenson, C.J. 1936. Lygus bugs in relation to occurrence of shrivelled alfalfa. Journal of Economic

Entomology 29: 454-457.

Udayagiri, S. and S.C. Welter. 2000. Escape of Lygus hesperus (Heteroptera: Miridae) eggs from

parasitism by Anaphes iole (Hymenoptera: Mymaridae) in strawberries: plant structure effects.

Biological Control 17: 234-242.

Wilkinson, L., Hill, M., Welna, J.P. and Birkenbeuel, G.K. 1997. SYSTAT, Version 7.0. SYSTAT Inc..

Evanston, IIlinois.

Wise, I.L. and R.J. Lamb. 1998. Economic threshold for plant bugs, Lygus spp. (Heteroptera: Miridae), in

canola. The Canadian Entomologist 130: 825-836.

J. ENTOMOL. SOC. BRIT. COLUMBIA 97, DECEMBER 2000 95

Early records of alien species of Heteroptera

(Hemiptera: Prosorrhyncha) in Canada

D.I. BARNES, H.E.L. MAW

EASTERN CEREAL AND OILSEED RESEARCH CENTRE, AGRICULTURE AND

AGRI-FOOD CANADA, OTTAWA, ON KIA 0C6

G.G.E. SCUDDER

DEPARTMENT OF ZOOLOGY AND CENTRE FOR BIODIVERSITY RESEARCH,

UNIVERSITY OF BRITISH COLUMBIA, VANCOUVER, BC V6T 1Z4

ABSTRACT

Early records for 45 alien species of Heteroptera in the provinces across Canada are

reported. Nine of the records constitute new provincial records, with 69 others

documenting published records of occurrence in detail for the first time. There are also 51

reported occurrences in provinces prior to published literature records.

INTRODUCTION

Many alien Heteroptera have become important agricultural crop pests in Canada (Beirne

1972), and more can be expected. The potential invasion appears greatest for those bug species

that either overwinter as adults, or oviposit in plant tissue (Lattin and Oman 1983).

Research has thus been initiated on the origins, distribution patterns, pathways of spread,

and ecological impact of alien Heteroptera in Canada (Scudder and Foottit 2000). It is

important to understand not only the history of alien species occurrence and impact in the past,

but also to undertake future pest risk assessment and the potential effects of impending climate

change. New evidence suggests that invasive species share traits that will allow them to

capitalize on the various elements of the forthcoming global change (Dukes and Mooney 1999)

Hence, over the past few years, most of the major collections of insects across Canada have

been visited, specimens of alien species of Heteroptera examined and named, the capture

information databased and the locality data georeferenced. These data have been combined

with published records to document the distribution of species in Canada, their probable place

of introduction, and their subsequent dispersal.

In the process, many new provincial records were discovered. Most of these have been

tabulated by Maw ef al. (2000), but the voucher data for these records have not been

published. For 45 of the total 79 alien Heteroptera in Canada, we thus document 9 new

provincial records (indicated by an asterisk) that are not included in Maw et al. (2000), plus 69

others that document published records of occurrence in detail for the first time. We cite only

the earliest records for the provinces.

Our research on collections also shows that some of the alien species have been collected

in provinces earlier than published records currently indicate. Since early records are important

for tracing the spread of these species, we document also 51 reported occurrences prior to the

literature records.

The order of species in the text follows Maw et al. (2000). Provinces (by acronym) are

arranged in alphabetical order.

J. ENTOMOL. SOC. BRIT. COLUMBIA 97, DECEMBER 2000

Collection abbreviations are as follows:

CNC: Canadian National Collection, Agriculture and Agri-Food, Ottawa, ON.

FRNF: Department of Forest Resources and Lands, Corner Brook, NF.

LEMQ: Lyman Entomological Museum, Macdonald College, McGill University, Ste-Anne-

de Bellevue, QC:

MU: Department of Biology, Memorial University, St. John’s, NF.

NSM: Nova Scotia Museum, Halifax, NS.

ROM: _ Royal Ontario Museum, Toronto, ON.

UBC: — Spencer Entomological Museum, University of British Columbia, Vancouver, BC.

UG: Department of Environmental Biology, University of Guelph, Guelph, ON.

USNM: National Museum of Natural History, Smithsonian Institution, Washington, DC.

Infraorder CIMICOMORPHA

Family ANTHOCORIDAE

Anthocoris confusus Reuter

NS: Kentville, 15-17.vii.1966 (L.A. Kelton) [CNC]. Reported from Nova Scotia

without date by Kelton (1978).

Orius minutus (Linnaeus)

BC: Abbotsford, 7.1x.1950 [CNC]. Previously reported from British Columbia in

1951 by Tonks (1953) and Downes (1957).

Family MIRIDAE

Tribe MIRINI

Adelphocoris lineolatus (Goeze)

AB: Elkwater Park, 28.vi1.1952 (L.A. Konotopetz) [CNC]. BC: no data, 1964 (J.R.

Hill) [LEMQ]. ON: Vineland Station, 1.1x.1942 ( W.L.Putman) [CNC]. PE: Wood

Island, 9.vili.1929 (R.P. Gorham) [CNC]. OC: Mont Royal, 26.viii.1944 [LEMQ].

Recorded from Alberta in 1968 (Craig 1971), Ontario in 1949 (Phillips 1951) and

Quebec (Moore 1950; Larochelle 1984) without dates.

Camptozygum aequale (Villers)

QC: Saint-Antoine-Abbé, 3.vi.1983 (Larochelle & Lariviére) [LEMQ].

Capsus ater (Linnaeus)

NB: Nashwaak Bridge, 22.vi.1934 (C.E. Atwood) [ROM]. PE: Woods Island,

11.vii.1966 (L.A. Kelton) [CNC]. Kelton collected this species at 9 localities

throughout New Brunswick in late June to early July of 1966.

Closterotomus norwegicus (Gmelin)

BC: Royal Oak, 24.vii.1917 (W. Downes) [CNC]. NB: Fredericton, 22.viii.1940

(R.P. Gorham) [CNC]. PE: Brackley Beach, 24.vii.1940 (J. McDunnough) [CNC].

Recorded from British Columbia (Kelton 1959) and Prince Edward Island (Kelton

1982a) without dates.

Phytocoris populi (Linnaeus)

* MB: Bald Head Hills (20.9 km (13 mi) n of Glenboro), 8.viii.1958 (J.G. Chillcott)

[CNC]. NS: Halifax, 7-8.viii.1966 (L.A. Kelton) [CNC]. *ON: Gull L., 6.vili.1918

(H.S. Parish) [CNC].

J. ENTOMOL. SOC. BRIT. COLUMBIA 97, DECEMBER 2000 97

Phytocoris tiliae (Fabricius)

NS: Halifax, 7-8.viii.1966 (L.A. Kelton) [CNC].

Stenotus binotatus (Fabricius)

MB: Falcon Lake, 25.vi.1972 (L.A. Kelton) [CNC]. MS: Kings Co., 15.vii.1948

(Ralph J. Day) [NSM]. ON: Trenton, 26.vi.1911 (Evans) [CNC]. PE: Cavendish

National Park, 9.vii.1966 (L.A. Kelton) [CNC]. OC: Aylmer, 27.vi.1924 (C.H.

Curran) [CNC]. Recorded from Ontario by Gibson (1912) without year, and

Manitoba (Kelton 1980), Ontario and Quebec (Carvalho 1959) without dates.

Tribe STENODEMINI

Leptopterna dolabrata (Linnaeus)

*AB: Highwood River Picnic Area, Hwy. 541, 24.vii.1994 (M.D. Schwartz) [CNC].

MB: Moose Lake, 25.vi.1972 (L.A. Kelton) [CNC]. MS: Halifax, 1899 [CNC].

Osborn (1918) and Parshley (1923) recorded specimens from Nova Scotia collected

in 1915; recorded from Manitoba (Kelton 1980) without dates.

Megaloceroea recticornis (Geoffroy)

*AB: Waterton Park, 3.vili.1994 (M.D. Schwartz) [CNC]. NB: Fredericton,

23.vi.1966 (L.A. Kelton) [CNC]. MS: Bible Hill, 12.vii.1966 (L.A. Kelton) [CNC].

PE: Cavendish National Park, 9.vii.1966 (L.A. Kelton) [CNC]. QC: North Hatley,

25.vil.1929 (G.S. Walley) [CNC]. Recorded from Quebec by Larochelle (1984)

without date, and Wheeler and Henry (1992) recorded collections from Nova Scotia

in 1971.

Pithanus maerkeli ((Herrich-Schaeffer)

AB: Elkwater, 16.vii.1952 (A.R. Brooks) [CNC]. NB: Fundy National Park,

6.vi1.1966 (L.A. Kelton) [CNC]. ON: Rondeau Park, 13.vi.1929 (G.S. Walley)

[CNC]. PE: Cavendish National Park, 9.vii.1966 (L.A. Kelton) [CNC]. QC: Laniel,

10-11.vit.1963 (L.A. Kelton) [CNC]. Previously recorded from Alberta, New

Brunswick, Ontario, Prince Edward Island and Quebec, but without dates (Kelton

1966).

Subfamily ORTHOTYLINAE

Tribe HALTICINI

Halticus apterus (Linnaeus)

NB: Fredericton, 192[9?] [CNC]. NF: Pasadena, 24.viii-3.ix.1985 (B. 4820) [CNC;

FRNF]. PE: Rustico, 4.vili.1966 (L.A. Kelton) [CNC].

Orthocephalus coriaceus (Fabricius)

BC: Galiano Island (north end), 1.vii.1976 (G.G.E. Scudder) [UBC]. NB:

Woodstock, 22.vi.1966 (L.A. Kelton) [CNC]. VF: Baie D’Espoir, low vegetation in

field, 11.vii.1985 (L.H. Hollett) [FRNF]. VS: Lockeport, 2.viii.1958 (J.R. Vockeroth)

[CNC]. ON: Niagara Falls, 7.vii.1955 (L.A. Kelton) [CNC]. PE: Cavendish National

Park, 9.vii.1966 (L.A. Kelton) [CNC]. OC: Georgeville, 27.vi.1936 (G.S. Walley)

[CNC]. Reported in 1973 from Ontario (Reid ef al. 1976). Kelton collected this

species from 9 localities throughout New Brunswick in late June 1966.

J. ENTOMOL. SOC. BRIT. COLUMBIA 97, DECEMBER 2000

Orthocephalus saltator (Hahn)

NF: Carbonear, 2.viii. 1925 (J.M. Swaine) [CNC]. OC: L'Anse-Pleureuse, 29.vii.1984

(Larochelle & Lariviére) [LEMQ].

Tribe ORTHOTYLINI

Blepharidopterus angulatus (Fallén)

NB: Saint John (Rockwood Park), 4.viii.1954 (J.F. Brimley) [CNC]. NF: Pasadena,

18.vili.1984 (D. Langor) [FRNF]. PE: Cavendish National Park, 3-5.vii.1966 (L.A.

Kelton) [CNC]. Reported from Prince Edward Island without date (Kelton 1982a).

Heterotoma planicornis (Pallas)

BC: Victoria, 6.vili.1923 (W. Downes) [CNC]. NS: Halifax, 10.viii.1980 (L.A.

Kelton) [CNC]. Recorded from British Columbia in 1933 (Downes 1957), and Nova

Scotia without date (Kelton 1982a).

Melanotrichus concolor (Kirschbaum)

NS: Shelburne, 2.vili.1958 (J.R. Vockeroth) [CNC].

Melanotrichus flavosparsus (Sahlberg)

AB: Manyberries (Experimental Range), 4.vili.1951 (D.F. Hardwick) [CNC]. BC:

Vernon, 22.1x.1923 (D.G. Gillespie) [CNC]. MB: Aweme, 19.vi.1930 (R.M. White)

[CNC]. NB: Fredericton (Experimental Station), 18.vi.1925 [CNC]. N7: Fort

Simpson, 19.v1.1950 (D.P. Whillans) [CNC]. ON: Trenton, 16.1x.1903 (Evans)

[CNC]. PE: Cavendish National Park, 9.vi1.1966 (L.A. Kelton) [CNC]. SK: Estevan,

27.vill.1929 (P.C. Brown) [CNC]. Previously reported from Ontario in 1914 (Gibson

1917), and Alberta, Manitoba and Saskatchewan (Kelton 1980) without dates.

Melanotrichus virescens (Douglas & Scott)

NS: Shelburne, 2.vii.1958 (J.R. Vockeroth) [CNC].

Orthotylus nassatus (Fabricius)

ON: Niagara, 25.vii.1963 (L.A. Kelton) [CNC].

Subfamily PHYLINAE

Tribe PHYLINI

Amblytylus nasutus (Kirshbaum)

BC: Saanich District, 6.vii.1950 (B.P. Beirne) [CNC]. ON: Marmora, 13.vi.1952

(J.C. Mitchell) [CNC].

Asciodema obsoletum (Fieber)

BC: Vancouver (UBC), 6.vii.1959 (G.G.E. Scudder) [UBC]. Recorded from British

Columbia in 1963 (Waloff 1966).

Atractotomus magnicornis (Fallén)

NF: St. John’s, 2.x.1980 (D. Northcott) [MU]. NS: Halifax, 22.vii.1976 (L.A. Kelton)

[CNC]. ON: Aldershot, 6.vii.1955 (L.A. Kelton) [CNC].

J. ENTOMOL. SOC. BRIT. COLUMBIA 97, DECEMBER 2000 99

Atractotomus mali (Meyer-Diir)

BC: Oliver, 19.vii.1970 (L.A. Kelton) [CNC]. PE: Charlottetown, 7-10.viii.1976

(L.A. Kelton) [CNC]. Reported from Prince Edward Island without date by Kelton

(1982a).

Campylomma verbasci (Meyer-Diir)

NS: Kentville, 19.vii.1923 (R.P. Gorham) [CNC]. ON: Jordan, 27.vili.1915 (W.A.

Ross) [UG]. PE: Dalvay House, 13.viii.1940 (G.S. Walley) [CNC]. Previously

reported from Ontario in 1950 (Phillips 1951) and Nova Scotia in 1931 by Gilliatt

(1935).

Compsidolon salicellum ((Herrich-Schaeffer)

ON: Presque Isle Provincial Park, 14.vili.1991 (M.D. Schwartz) [CNC]. PE:

Burlington, 4.vili.1966 (L.A. Kelton) [CNC]. Previously recorded from Prince

Edward Island in 1976 by Kelton (1982b).

Lepidargyrus ancorifer (Fieber)

BC: Langley, 17.vii.1959 (L.A. Kelton) [CNC]. ON: Rainy River District, 30.vi.1924

(J.F. Brimley) [CNC].

Lopus decolor (Fallén)

BC: Saltspring Island, 30.vii.1981 (G.G.E. Scudder) [UBC]. NB: Fredericton,

192[9?] [CNC]. NF: Pasadena, 3.vii1.1984 (D. Langor) [FRNF]. ON: Norway Point

(Lake of Bays), 15.vii.1922 (J. McDunnough) [CNC]. PE: Brackley Beach,

29.vii.1940 (G.S. Walley) [CNC]. Recorded from Ontario (Carvelho 1958) without

date.

Megalocoleus molliculus (Fallén)

NB: Grey Mills [?=Grey Hills], 13.vii.1921 (R.P. Glorham]) [CNC]. VF: Silverdale,

31.vii.1980 (L.A. Kelton) [CNC]. MS: Parrsboro (Ottawa House), 9.vili.1943 (J.

McDunnough) [CNC]. ON: Strathroy, 12.vii.1936 [CNC]. PE: Wood Island, ix.1929

[CNC]. OC: Forestville, 8.viii.1950 (R. de Ruette) [CNC]. Previously recorded from

Ontario in 1972 by Reid et al. (1976), Nova Scotia in 1967 by Wheeler and Henry

(1992) and Quebec (Larochelle 1984) without date.

Parapsallus vitellinus (Scholtz)

*NS: Kentville, 3-6.vi1.1976 (L.A. Kelton) [CNC]. ON: Saint Catharines, 22.v1.1961

(Kelton & Brumpton) [CNC]. Previously recorded from Ontario in 1978 (Henry and

Wheeler 1979).

Plagiognathus arbustorum (Fabricius)

BC: Lulu Is., 23.viii.1954 (W. Downes) [UBC]. NF: St. John’s, Pippy Park,

14.1x.1982 [FRNF]. Previously recorded from British Columbia in 1959 (Kelton

1982c) and 1963 (Waloff 1966).

J. ENTOMOL. SOC. BRIT. COLUMBIA 97, DECEMBER 2000

Plagiognathus chrysanthemi (Wolff)

*AB: Waterton Park, 3.viil.1994 (M.D. Schwartz) [CNC]. BC: Creston, 7.viii.1948

(D.B. Waddell) [CNC]. NB: Fredericton, 1.viii.1919 (R.P. Gorham) [CNC]. NF:

Carbonear, 2.vili.1925 (J.M. Swaine) [CNC]. *SK: Melville 5 km E, Rt. 16,

18.vi1.1990 (M.D. Schwartz) [CNC]. Previously reported from British Columbia in

1951 (Tonks 1953), Newfoundland in 1949 (Lindberg 1958), and New Brunswick

(Kelton 1982a) without date.

Sthenarus rotermundi (Scholtz)

*OC: St-Jean-Baptiste[-de-]Rouville, 22.vi.1992 (J. F. Roch) [CNC].

Tribe PILOPHORINI

Pilophorus perplexus Douglas & Scott

BC: Victoria, 3.vili.1927 (W. Downes) [UBC]. ON: Guelph, 15.vii.1916 [CNC; UG].

PE: Cavendish, 13-19.viii.1976 (L.A. Kelton) [CNC]. Reported from Ontario by

Knight (1941) and from Ontario and Prince Edward Island by Schuh and Schwartz

(1988) without dates.

Family TINGIDAE

Dictyla echii (Schrank)

OC: Mont Royal, ex. Echium vulgare L., 14.vii.1997 (J.F.R.) [CNC; USNM].

Dictynota fuliginosa Costa

BC: Victoria, 29.vil.1957 (N.H. Anderson) [CNC]. Previously reported from British

Columbia in 1959 (Scudder 1960).

Stephanitis rhododendri Horvath

NB: Saint John (Rockwood Park), 12.viii.1953 (J.F. Brimley) [CNC]. VF: Miguels

Lake, 28.i1x.1983 (L.H. Hollett) [CNC]. ON: Vineland Station, 16.vii.1963 (W.L.

Putnam) [CNC]. OC: Mount Lyall, 12.vii.1933 (W.J. Brown) [CNC].

Family REDUVIIDAE

Empicoris pilosus (Fieber)

NS: Halifax, 22.vii.1976 (L.A. Kelton) [CNC]. ON: DeGrassi Point, 14.vili.1915

(E.M. Walker) [ROM]. QC: Brome, 9.v1.1948 (G.S. Walley) [CNC].

Empicoris vagabundus (Linnaeus)

NF: Crooked Lake, 19.viti. 1984 (L.H. Hollett) [CNC]. MS: Halifax, 8.viii.1966 (L.A.

Kelton) [CNC]. ON: Prince Edward County, |.viti.1940 (J.F. Brimley) [CNC]. PE:

Rustico, 4.viil.1966 (L.A. Kelton) [CNC]. OC: Québec, 24. viii. 1959 (A.E. Strasby)

[CNC].

Family BERYTIDAE

Berytinus minor ((Herrich-Schaeffer)

OC: Ste-Anne-de-Bellevue, 24.v.1947 (G.A. Moore) [LEMQ]. Previously reported in

Quebec without date (Moore 1950).

Family RHYPAROCHROMIDAE

Megalonotus sabulicola (Thomson)

ON: Prince Edward County, 9.v.1965 (J.F. Brimley) [CNC].

J. ENTOMOL. SOC. BRIT. COLUMBIA 97, DECEMBER 2000 101

Stygnocoris rusticus (Fallén)

AB: Millet, 11.viii.1993 (A.S. McClay) [CNC]. *NVB: Saint Andrews , 30.1x.1984

(G.G.E. Scudder) [CNC]. NF: St. John’s, Long Pond, 12.vili.1977 (D. Larson)

[CNC]. QC: Mont Royal, 27.viii.1907 [CNC]. Previously reported from Quebec in

1916 (Criddle 1922).

Stygnocoris sabulosus (Schilling)

AB: Brocket, 19 km N, 49°43'N 113°45'W, 1410 m, pan trap, 10-14.vili.1998 (K.

White) [CNC]. NB: Fredericton, 30.vii.1949 (D.G. Cameron) [CNC]. ON: Guelph,

10.1x.1971 (C.D. Rollo) [UG]. PE: Charlottetown, 7-10.viil.1976 (L.A. Kelton)

[CNC].

Family PENTATOMIDAE

Picromerus bidens (Linnaeus)

NB: Kouchibouguac National Park, 21.vii.1977 (S.J. Miller) [CNC]. NS: Amherst

Shore, 11.viii.1980 (J. Gilhen) [NSM]. ON: York Co., Whitchurch, 8.vili.1974

(Antonucci) [ROM]. PE: Charlottetown, 7-10.vili.1976 (L.A. Kelton) [CNC].

Reported from New Brunswick, Nova Scotia, Ontario and Prince Edward Island

(Lariviére and Larochelle 1989) without dates, but recorded from Quebec in 1968 by

Kelton (1972).

ACKNOWLEDGEMENTS

Research was supported by grants to G.G.E. Scudder from the Natural Sciences and

Engineering Research Council of Canada. We are indebted to D.C. Currie, R.G. Foottit, R.C.

Froeschner, J. Heron, L.H. Hollett, D.J. Larson, S.A. Marshall, M.D. Schwartz and K. White

for support, loan of material or help with records.

REFERENCES

Beirne, B.P. 1972. Pest insects of annual crop plants in Canada, IV, Hemiptera-Homoptera; V, Orthoptera; V1,

Other groups. Memoirs of the Entomological Society of Canada 85:1-73.

Carvalho, J.C.M. 1958. A catalogue of the Miridae of the world. Part II. Subfamily Phylinae. Arquiros do Museu

Nacional, Rio de Janeiro 45(2):1-216.

Carvalho, J.C.M. 1959. A catalogue of the Miridae of the World. Part IV. Subfamily Mirinae. Arquiros do

Museu Nacional, Rio de Janeiro 48(4): 1-384.

Craig, C.H. 1971. Distribution of Adelphocoris lineolatus (Heteroptera: Miridae) in western Canada. The

Canadian Entomologist 103:280-281.

Criddle, N. 1922. The entomological record, 1921. Annual Report of the Entomological Society of Ontario

52(1921):57-70.

Downes, W. 1957. Notes on some Hemiptera which have been introduced into British Columbia. Proceedings of

the Entomological Society of British Columbia 54:11-13.

Dukes, J.S. and H.A. Mooney. 1999. Does global change increase the success of biological invaders? Trends in

Ecology and Evolution 14:135-139.

Gibson, A. 1912. The entomological record, 1911. Annual Report of the Entomological Society of Ontario

42(1911):89-112.

Gibson, A. 1917. The entomological record, 1916. Annual Report of the Entomological Society of Ontario

47(1916):137-171.

Gilliatt, F.C. 1935. Some predators of the European red mite, Paratetranychus pilosus C.&F., in Nova Scotia.

Canadian Journal of Research 13(D)(2):19-38.

Henry, T.J. and A.G. Wheeler, Jr. 1979. Palearctic Miridae in North America: records of newly discovered and

little-known species (Hemiptera: Heteroptera) Proceedings of the Entomological Society of Washington

81:257-268.

102 J. ENTOMOL. SOC. BRIT. COLUMBIA 97, DECEMBER 2000

Kelton, L.A. 1959. Male genitalia as taxonomic characters in the Miridae (Hemiptera). The Canadian

Entomologist Supplement 11:1-72.

Kelton, L.A. 1966. Pithanus maerkeli (Herrich-Schaffer) and Actitocoris signatus Reuter in North America

(Hemiptera: Miridae). The Canadian Entomologist 98:1305-1307.

Kelton, L.A. 1972. Picromerus bidens in Canada (Heteroptera: Pentatomidae). The Canadian Entomologist

104:1743-1744.

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

Canada Department of Agriculture Publication 1639:101 pp.

Kelton, L.A. 1980. The insects and arachnids of Canada. Part 8. The plant bugs of the Prairie Provinces of

Canada (Heteroptera: Miridae). Agriculture Canada Research Publication 1703:408 pp.

Kelton, L.A. 1982a. Plant bugs on fruit crops in Canada. Heteroptera: Miridae. Research Branch, Agriculture

Canada Monograph 24:1-201.

Kelton, L.A. 1982b. Description of a new species of Plagionathus Fieber and additional records of European

Psallus salicellus in the Nearctic Region (Heteroptera: Miridae). The Canadian Entomologist 14:169-172.

Kelton, L.A. 1982c. New and additional records of Palearctic Phy/us Hahn and Plagiognathus Fieber in North

America (Heteroptera: Miridae) The Canadian Entomologists 114:1127-1128.

Knight, H.H. 1941. The plant bugs, or Miridae, of Illinois. Illinois Natural History Survey Bulletin 22:1-234.

Lariviere, M.-C. and A. Larochelle. 1989. Picromerus bidens (Heteroptera: Pentatomidae) in North America,

with a world review of distribution and bionomics. Entomological News 100:133-146.

Larochelle, A. 1984. Les punaises terrestres (Heteroptera: Geocorises) du Quebec. Fabreries, Supplement 3:5 13

Pp.

Lattin, J.D. and P. Oman. 1983. Where are the exotic insect threats? pp. 93-137. In: Wilson, C.L. and C.L.

Graham (Eds.) Exotic Plant Pests and North American Agriculture. Academic Press, New York.

Lindberg, H. 1958. Hemiptera Heteroptera from Newfoundland, collected by the Swedish-Finnish Expedition of

1949 and 1951. Acta Zoologica Fennica 96:1-25.

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

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

Moore, G.A. 1950. Catalogue des Hémipteéres de la Province de Québec. Le Naturaliste Canadien 77:233-271.

Osborn, H. 1918. The meadow plant bug, Miris dolabratus. Journal of Agricultural Research 15:175-200.

Parshley, H.M. 1923. Records of Nova Scotian Hemiptera-Heteroptera. Proceedings of the Acadian

Entomological Society 1922:102-108.

Phillips, J.H.H. 1951. An annotated list of Hemiptera inhabiting sour cherry orchards in the Niagara Peninsula,

Ontario. The Canadian Entomologist 83:194-205.

Reid, D.G., C.C. Loan and R. Harmsen. 1976. The mirid (Hemiptera) fauna of Solidago canadenisis

(Asteraceae) in south-eastern Ontario. The Canadian Entomologist 108:561-567.

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

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

Scudder, G.G.E. 1960. Dictyonota fuliginosa Costa (Hemiptera: Tingidae) in the Nearctic. Proceedings of the

Entomological Society of British Columbia 57:22.

Scudder, G.G.E. and R.G. Foottit. 2000. Alien bugs (Insecta: Hemiptera) in Canada: Distribution patterns,

pathways of spread and ecological impact (in) Claudi, R. (ed.) Ecosystem Globalization: Threat to Canadian

Biodiversity. Natural Resources Canada, Ottawa (in press).

Tonks, N.V. 1953. Annotated list of insects and mites collected on brambles in the lower Fraser Valley, British

Columbia, 1951. Proceedings of the Entomological Society of British Columbia 49:27-29.

Waloff, N. 1966. Scotch broom (Sarothamnus scoparius (L.) Wimmer) and its insect fauna introduced into the

Pacific Northwest of America. Journal of Applied Ecology 3:293-311.

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

biology, and origin, with emphasis on North America. The Thomas Say Foundation 15:282 pp.

J. ENTOMOL. SOC. BRIT. COLUMBIA 97, DECEMBER 2000 103

The ambrosia beetle, Gnathotrichus retusus (Coleoptera:

Scolytidae) breeding in red alder, Alnus rubra (Betulaceae)

SUSANNE KUHNHOLZ', JOHN H. BORDEN

and RORY L. McINTOSH?

CENTRE FOR ENVIRONMENTAL BIOLOGY,

DEPARTMENT OF BIOLOGICAL SCIENCES, SIMON FRASER UNIVERSITY,

8888 UNIVERSITY DRIVE, BURNABY, BC, V5A 1S6

ABSTRACT

Brood adult ambrosia beetles recovered from well established galleries in a wind-

thrown red alder, A/nus rubra Bongard, on Burnaby Mountain, Burnaby, British

Columbia, were identified as Gnathotrichus retusus LeConte. The tree was attacked to

a height of 25.8 m. Galleries penetrated up to 17.5 cm into the wood. The mean density

of gallery entrance holes (t SE) was 120 + 31.9 per m’ of the bark surface. The mean

production of brood in five completely dissected galleries was 13.2 + 5.5. These results

show conclusively that G. retusus in British Columbia can breed successfully in an

angiosperm host.

Key Words: Guathotrichus retusus, Alnus rubra, ambrosia beetle

INTRODUCTION

The ambrosia beetle, Gnathotrichus retusus LeConte, is widely distributed throughout

western North America, from Alaska to Baja California (Bright 1976; Wood 1982). This

monogamous, univoltine species commonly attacks dying, standing, or recently cut or

fallen coniferous trees in the genera Picea, Pinus, Pseudotsuga, Tsuga and Abies, but is

also recorded as attacking angiosperm trees, specifically alders, A/nus spp. and Populus

trichocarpa Torrey (Nijholt 1981; Wood 1982). Wood (1982) synonymized G. alni

Blackman (Blackman 1931) and G. retusus, because he failed to see distinguishing

characteristics. The reproductive biology and the breeding success of G. retusus in

angiosperm trees are not well documented, although Nijholt (1981) reported live striped

ambrosia beetles, Trypodenron lineatum Olivier, and G. retusus and their progeny in red

alder, A/nus rubra Bongard, that had been killed with 2,4-D. We describe and verify the

successful attack and brood production by Guathotrichus retusus in red alder.

OBSERVATIONS

Two large red alder trees of similar size were found fallen over a trail on the south

slope of Burnaby Mountain Park in Burnaby, British Columbia, Canada, at the beginning

of August 1998. The trees were partially rooted and were most likely wind-thrown in a

severe storm in January of 1997, and attacked in the spring of 1998. No evidence of root

rot or other disease was apparent, but reddish-brown frass on the bark was observed around

the gallery entrances of the alder bark beetle, A/niphagus aspericollis LeConte, a common

bark beetle in this locale (Borden 1969). Creamy-white frass occurred at the entrance to

' Author to whom correspondence should be addressed

* Current address: Forest Ecosystems Branch, Saskatchewan Environment & Resource

Management, Box 3003, Prince Albert, Saskatchewan, S6V 6G1, Canada

104 J. ENTOMOL. SOC. BRIT. COLUMBIA 97, DECEMBER 2000

ambrosia beetle galleries, later identified, using beetles recovered from the galleries, to be

those of G. retusus.

One of the trees was measured in detail. The tree was 30 m high, 43.9 cm in diameter at

1 m height from the root collar, and had major branches at 17, 19.3, 21.8, and 22 m. Attack

by G. retusus occurred from the root collar to a height of 25.8 m (8.7 cm diam.). On 16

August 1998, three disks-ca. 20 cm long were cut from the tree with a chain saw at 2, 8.5,

and 15 m up the bole. The densities of G. retusus gallery entrances on these disks ranged

from 36 tol99 per m’, representing a mean attack density (tSE) of 120 + 32 per m? of bark

surface. With the first cut into the tree at 3 m height the trunk shattered lengthwise and

revealed extensive gallery systems, from which numerous living callow and mature adult

beetles emerged. These galleries penetrated the wood to a depth of 17.5 cm, and were

stained dark brown, very similar to G. retusus galleries in conifers. They contained adults,

callow adults, larvae, and eggs, samples of each life stage were collected in 70% ethanol.

Voucher specimens of mature adults have been deposited in the collection of the Pacific

Forestry Centre, Canadian Forest Service, Victoria, BC.

From the cut discs, 11 galleries were dissected, revealing all life stages. One gallery

dissected in February 1999, from a disk that had been left outside in the shade, contained

egg niches, larvae, pupae, and both callow and mature adults, indicating that as in conifers

(Prebble and Graham 1957; Chamberlin 1958) G. retusus in red alder can overwinter in

any life stage. Galleries were typically forked two to four times, and curved in all

directions, not only on one plane as in coniferous hosts (Wood 1982), possibly due to the

lack of dense annual rings as found in conifers. Pupal and adult cradles (tSE) were 4.6

+0.4 mm long, 2 mm wide, and 1 mm apart (m=25), staggered in an alternate pattern in one

plane on both sides of a main gallery. The mean number of summed niches and cradles in

five completely dissected galleries was 13.2 + 5.5, a similar level of brood production as

occurs in conifers (Liu and McLean 1993).

Our observations indicate that attack and brood production by G. retusus in red alder is

consistent with descriptions of the beetles’ biology in coniferous host species (Liu and

McLean 1993). These results show conclusively that G. retusus in British Columbia can

breed successfully in an angiosperm host.

ACKNOWLEDGEMENTS

This research was supported by the Natural Sciences and Engineering Research Council of

Canada, Forest Renewal B.C., the Science Council of B.C. and 19 forest industry

companies.

REFERENCES

Blackman, M.W. 1931. Revision of Gnathotrichus. Journal of the Washington Academy of Science.

21:264-276.

Borden, J.H. 1969. Observations on the life history and habitats of A/niphagus aspericollis (Coleoptera:

Scolytidae) in southwestern British Columbia. The Canadian Entomologist 101:870-878.

Bright, D.E. 1976. The Insects and Arachnids of Canada, Part 2: The bark beetles of Canada and Alaska.

Canada Department of Agriculture Publication Number 1576, 241 pp.

Chamberlin, W.J. 1958. The Scolytidae of the Northwest. Oregon State College Press, Corvallis, Oregon,

205 pp.

Liu, Y.-B. and J.A. McLean. 1993. Observations on the biology of the ambrosia beetle Gnathotrichus

retusus (LeConte) (Coleoptera: Scolytidae). The Canadian Entomologist 125:73-83.

Nijholt, W.W. 1981. Ambrosia beetles in alder. Canadian Forestry Service Research Notes 1(2):12.

Prebble, M.L. and K. Graham.1957. Studies of attack by ambrosia beetles in softwood logs on Vancouver

Island, British Columbia. Forest Science 3:90-112.

Wood, S.L. 1982. The bark and ambrosia beetles of North and Central America (Coleoptera: Scolytidae), a

taxonomic monograph. Great Basin Naturalist Number 6, 1163 pp.

J. ENTOMOL. SOC. BRIT. COLUMBIA 97, DECEMBER 2000 105

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Figure 10. Features referred to in the Key to the Major Workers of the Species. a) Clypeus

bearing a notch in the ventral border; b) clypeus without a ventral notch; c) profile of a

rounded epinotum; d) profile of an angulate epinotum.

a | | b

Cet

Figure 11. Features referred to in the Key to the Major Workers of the Species. a) Concave

occipital border; b) non-concave occipital border; c) profile of pronotum with basal and

declivitous faces meeting at an angle; d) profile of non-angulate pronotum.

Erratum

The Editor regrets that Figs. 10 and 11, pg. 55, Volume 96, December 1999 were inadvertently

reversed and should have been printed as shown above.

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Directors of the Entomological Society of British Columbia 2000-2001 .....esss+ssesceeeeeveee2

Allison, J.D., R.L. McIntosh, J.H. Borden and L.M. Humble. A new parasitoid (Diptera:

Tachinidae) of Acanthocinus princeps (Coleoptera: Cerambycidae) in North America.....3

Progar, R.A., M.T. AliNiazee and J.L. Olsen. The economic and environmental impact of an

IPM program on hazelnuts in Oregon... «255... oo) scnudcsscne 0 «oc sstse dete annie ne nee

Li, S.Y. and I.S. Otvos. Enhancement of the activity of a nuclear nolphegee virus by an

optical brightener in the eastern hemlock looper, Lambdina fiscellaria fiscellaria

(Lepidoptera: Geometridae)

Mayer, D.F., E.R. Miliczky, B.F. epee and C.A. Johansen. The bee fauna Bieta

Apa) of southeastern Washington... 2. sccs.ccie pcs chu a

Dodds, K.J., D.W. Ross and G.E. Daterman. A comparison of traps and trap trees ee

capturing Douglas-fir beetle, Dendroctonus pseudotsugae (Coleoptera: Scolytidae)...

Garland, J.A. The ee of Canada eee recent acquisitions ees in British

Columbia and Yukon.. ene senna diinySSeient ene ee awe

Miller, D.R. and B.S. Lada, Couey of a-pinene and myrcene on attraction of

mountain pine beetle, Dendroctonus ponderosae (Coleoptera: Scolytidae) to pheromones

in stands of western white pine

Kenner, R.D. Somatochlora kennedyi (Odonata: Corduliidae): a new species for British

Columbia, with notes on geographic variation in size and wing venation

Scudder, G.G.E. Heteroptera (Hemiptera: Prosorrhyncha) new to Canada. Part |

Miller, D.R. and J.H. Borden. Pheromone interruption of pine engraver, Ips pini, by

pheromones of mountain pine beetle, Dendroctonus ponderosae (Coleoptera:

Scolytidae)

Kenner, R.D. Gyrinus cavatus and G. minutus (Coleoptera: Gyrinidae) in British Columbia

with comments on their nearctic distributions

Morewood, W.D. Occurrence and inheritance of a colour pattern dimorphism in adults of

Hyalophora euryalus (Lepidoptera: Saturniidae)

Bomford, M.K., R.S. Vernon and P. Pats. Aphid (Homoptera: Aphididae) accumulation and

distribution near fences designed for cabbage fly (Diptera: Anthomytidae) exclusion .....79

McGregor, R.R., D.R. Gillespie, D.M.J. Quiring and D. Higginson. Parasitism of the eggs of

Lygus shulli and Lygus elisus (Heteroptera: Miridae) by Anaphes iole (Hymenoptera:

Mymaridae)

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

(Hemiptera: Prosorrhyncha) in Canada

Kiihnholz, S., JH. Borden and R.L. McIntosh. The ambrosia beetle, Gnathotrichus retusus

(Coleoptera: Scolytidae) breeding in red alder, A/nus rubra (Betulaceae)

NOTICE TO CONTRIBUTORS. ooo ooo sci eves wsnsetasenies ust cubeatiesen taaauadtenpeneuen «dele ane

Journal

of the

Entomological Society

of British Columbia

Volume 98 ISSN #0071-0733

Issued December 2001

\ \

Lohans 2 Entomological Society

"2001 of British Columbia

a) iy sr)

i £UU¢

COVER: ‘Beetles’ is the art of Richard Hunt, Victoria BC.

Richard kindly granted permission to use the design on the cover of Volume 98 of the

Journal, marking the 100" anniversary of the Entomological Society of British Columbia.

‘Beetles’ was the emblem for the International Congress of Entomology, Vancouver, BC,

Canada, 3-9 July 1988.

The Journal of the Entomological Society of British Columbia is published annually in

December by the Society.

Designed and typeset by David Raworth and David Holden.

Printed by Reprographics, Simon Fraser University, Burnaby, BC, Canada.

Printed on Recycled Paper.

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER, 2001 l

Preface

The Entomological Society of British Columbia marks its 100" anniversary in 2001. To

commemorate the occasion, the Executive of the ESBC decided to publish 14 invited, six-

page papers that sample the numerous entomological subjects that have been explored in

the province of British Columbia during the last 50 years. The response to this idea has

been overwhelming. In some cases the authors thought that it was important to review work

during the last 100 years, and many authors extended the length of their articles in order to

do the topic some justice. Given the finite resources of the ESBC, several topics have been

partially covered. Other topics have not been specifically examined, for example,

biological control and amateur entomology. However, the reader will catch a glimpse of

the enormous breadth and depth of entomological studies that have been conducted in the

Province. The scientific contribution of entomologists, on all scales from local to

international, is clear. Deep thanks are due to the authors and reviewers for taking the time

to contribute to this volume; to several authors who found additional funding to defray the

publication costs of their papers; and to Richard Hunt for permission to use his art on the

cover of this Volume.

David A. Raworth Editor

g) J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

Journal

of the

Entomological Society

of British Columbia

Volume 98 Issued December 2001 ISSN #0071-0733

Directors of the Entomological Society of British Columbia 2001-2002.........................4

HISTORICAL CONTRIBUTIONS

Belton, P., J.C. Arrand and H.R. MacCarthy. The second 50 years of entomology in British

Columbia — a brief perspective ..2......2.:c0nsiecasteereeenssndes Tniaetenecaneh Acluaieracden eae eee eege >

The G.J. Spencer Memorial Lecture Series at the University of British Columbia.................. 13

The H.R. MacCarthy Pest Management Lecture Series (Simon Fraser University and the

University of British Columbia) ..cs:.c:..:cssescne.coasercechentesaravseensendoasveousnuenyen eae eee IS)

Cannings, R.A., S.G. Cannings and G.G.E. Scudder. Insect collections, surveys and

conservation in British Columbia in the 20" CONGULY...2:.<csniedanaseesetoredgue eee tee See ee AF

Cannings, R.A. and G.G.E. Scudder. An overview of systematics studies concerning the insect

fauna of British Columbia. :...2:cscccccactses1eseyes-cuonssaenoneseanenganendatedaeboueteton neee eee eee eM 33

Scudder, G.G.E., K.M. Needham, R.D. Kenner, R.A. Cannings and S.G. Cannings. Aquatic

insects in British Columbia: 100 years of Studly......:...ccs.cc¢-ctseceedessenneersce tee een ems 61

Bennett, R.G. Spiders (Araneae) and araneology in British Columbia...............cccccccssereee ees 83

Gillespie, D.R. Arthropod introductions into British Columbia — the past 50 years...............91

Ring, R.A. Research in adaptations of arthropods in British Columbia................:ccscceeee es 99

Myers, J.H. and D.A. Raworth. Insect population ecology in British Columbia............... OF

Roitberg, B. and G. Gries. Behavioral and chemical ecology in British Columbia............113

Belton, P., A. Borkent and B. Costello. Arthropods that attack man and domestic animals in

British Columbia (195 1 = 2001). .c.csssssseceseessncasstsaceceenoesadceecsesDeseteesceeecct ye ee eee eee hs

Anderson, G.S. Forensic entomology in British Columbia: A brief history...........0.....66 AZT

van Westendorp, P. and D.M. McCutcheon. Bees and pollination in British Columbia........137

Vernon, R.S. Fifty years of entomological research in orchard and vegetable crops in British

CN 01 i saica zeae ode swseainencn sagas so cavneee leyaeunapee tees sescenae opp tenecnesteas eg Sete scene eee eee 143

Hall, P.M., J.M. Kinghorn, B.S. Lindgren, J.A. McLean and L. Safranyik. History of forest

insect investigations in British Columbia. I. Forest entomology education, research, and

INSECE MANAG CIIENE, sce../.savesesciecueaveseotnacaponee ducgedlveotasnadheaanceteel cee agin ae eat ee tee ee eee SB)

Van Sickle, A., R.L. Fiddick and C.S. Wood. History of forest insect investigations in British

Columbia. II. The Forest Insect and Disease Survey in the Pacific Region.................. 169

Rajala, R.A. History of forest insect investigations in British Columbia. III. The Vernon

Laboratory and federal entomology in British Columbia.................5-.25...-.cs1-asesoeostese ane 7)

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 3

SCIENTIFIC CONTRIBUTIONS - 2001 “a

VanLaerhoven, S.L. and G.S. Anderson. Implications of using development rates of blow fly

(Diptera: Calliphoridae) eggs to determine postmortem interval..................c:eeeeeee es 189

Allison, J.D. and J.H. Borden. Observations on the behavior of Monochamus scutellatus

(Coleoptera: Cerambycidae) in northern British Columbia... ee eeeeeeeee eee eats 195

Carcamo, H.A., L. Dosdall, M. Dolinski, O. Olfert and J.R. Byers. The cabbage seedpod

weevil, Ceutorhynchus obstrictus (Coleoptera: Curculionidae) —a review ..............5. 201

Fitzpatrick, S.M., J.A. Newhouse, J.T. Troubridge and K.A. Weitemeyer. Rearing the

cranberry girdler Chrysoteuchia topiaria (Lepidoptera: ee on reed canary grass

Phalaris arundinacea (Festucoideae: Panicoideae)............cc..0 cee iecaeees eel

Knight, A.L. Monitoring the seasonal population densify of poe pyrusana (uesidnoters

Tortricidae) within a diverse fruit crop production area in the Yakima Valley, WA.......217

Kucera, J.R., G.E. Haas and M.K. MacDonald. Fleas (Siphonaptera) from sciurid ant anna

rodents on the eastern slope of the Cascade Range, Kittitas County,

IW ASHI EN MOI ci a ahs asccesce etter neem wee tee eae rae cee eeeeat cat. vant dase anna eel eeae wer tesmne ns teen 227),

Horton, D.R. and P.J. Landolt. Use of Japanese-beetle traps to monitor flight of the Pacific

coast wireworm, Limonius canus (Coleoptera: Elateridae), and effects of trap height and

CO) OB rekeae ecencten eee kecsinctrse cy ncetgataageru asic saetnn cn cet cars mak ceseresioe ou tane tA eNea Se nce ueateeta Mewes ores 235

Poirier, L.M. and J.H. Borden. Qualitative analyses of larval oral exudate from eastern and

western spruce budworms (Lepidoptera: Tortricidae)...........cccccsccseeesecesseeseesseeessee eens 243

Holsten, E.H., R.E. Burnside and S.J. Seybold. Verbenone interrupts the response to

aggregation pheromone in the northern spruce engraver, /ps perturbatus (Coleoptera:

Scolytidae), in south-central and interior Alaska... cceeeseceseeeseeeseeessessssesseeeseeesen ees aot

Vernon, B., E. Lagasa and H. Phillip. Geographic and temporal distribution of Agriotes

obscurus and A. lineatus (Coleoptera: Elateridae) in British Columbia and Washington as

determined by pPRETOMOMNE trap SULVEYS, 2i5:--c:7e- peeessseresecobseyasencwecereesocaeseve-ctutee a alces <i 25d

NOTICE POICON ERIBUDORS 52s oacs pause dettenste esee, stcerevtety napieced tire ageneeee LON,

4 J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER, 2001

DIRECTORS OF THE ENTOMOLOGICAL SOCIETY OF

BRITISH COLUMBIA FOR 2001-2002

President

Lorraine Maclauchlan

BC Ministry of Forests, Kamloops

President-Elect

Gail Anderson

Simon Fraser University, Burnaby

Past-President

Rob Cannings

Royal British Columbia Museum, Victoria

Secretary / Treasurer

Robb Bennett

BC Ministry of Forests, 7380 Puckle Rd., Saanichton BC V8M 1W4

Editorial Committee (Journal)

Ward Strong (Editor) Dave Raworth Peter Belton

Ken Naumann Lorraine Maclauchlan

H.R. MacCarthy (Editor Emeritus)

Editor (Boreus)

Cris Guppy

Directors

Cris Guppy (1st) Ian Wilson (1st)

Rene Alfaro (2nd) Keith Deglow (2nd) Tracey Hueppelsheuser (2nd)

Honorary Auditor

Rob Cannings

Regional Director of National Society

Terry Shore

Canadian Forest Service, Victoria

Web page, (Bill Riel): http://www.harbour.com/commorgs/ESBC/

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 5

The second 50 years of entomology in British Columbia

—a brief perspective

P. BELTON

DEPARTMENT OF BIOLOGICAL SCIENCES, SIMON FRASER UNIVERSITY,

BURNABY, BC, CANADA VSA 1S6

J.C. ARRAND

1501 1035 BELMONT AVE., VICTORIA, BC, CANADA V8S 3T5

H.R. MACCARTHY

101 6001 YEW ST, VANCOUVER, BC, CANADA V6M 3Y7

Fifty years ago the history of entomology in the Province was reviewed in seven

articles dealing with different regions or subjects in our journal, then called the

Proceedings (Vol. 48, 1952).

It was the heyday of chlorinated hydrocarbon pesticides and new government

laboratories were being set up to study not only pest control with these new compounds

but also biology, biological control and the identification of difficult groups of insects like

root maggots and wireworms.

Entomological training in the Province was in the capable hands of Prof. G.J. Spencer

of the University of British Columbia, the Society’s President in 1951. Many of his

students became well-known in their own right, such as D.A. Chant, K. Graham, J.D.

Gregson, G.P. Holland, H.R. MacCarthy, and A.L. Turnbull. Between 1934 and 1937,

Spencer was also in charge of summer operations at the Dominion Insect Unit in

Kamloops following the untimely death of Eric Hearle, the Province’s founding medical

entomologist. Spencer was much more than just a capable academic. He retired in 1953

and the role of guiding entomologist at UBC was taken over in 1958 by Prof. G.G.E.

Scudder for the remainder of the century. Other entomologists expanded the topics covered

at that University when C.S. (Buzz) Holling was appointed to the new Institute of Animal

Resource Ecology, set up to study the interactions between fisheries and forestry. Holling

was its first full-time director and appointed another forest entomologist, W.G. Wellington

to it in 1970. The latter took over as director from 1973 to 1979, retiring in 1986. J.H.

Myers joined the institute in 1972. The Faculty of Forestry also employed entomologists

including Spencer’s former student, K. Graham, Professor from 1948-1977 and later, J.A.

McLean from 1977 to the present. The Faculty of Agriculture, Plant Science also

employed an entomologist, J. Sandess in the late 1960s, succeeded by B. Philogéne, R.H.

Elliott and in 1983 by the present incumbent, M.B. Isman.

R.A. Ring was hired to teach entomology for a year at UBC in 1964, and after a spell at

the Biosystematics Research Institute, Ottawa, returned and was appointed by the

University of Victoria in 1966, where he continues to teach.

There was a major influx of entomologists to the newly founded Simon Fraser

University in 1967. J.H. Borden had been appointed a year earlier but was soon joined by

B.P. Beirne and seven of his staff from the Federal Research Institute for Biological

Control in Belleville ON. Beirne set up the Pestology Centre, which flourished (with a

change of name to Centre for Pest Management) for about 30 years as an integral part of

the Department of Biological Sciences. The Masters of Pest Management Degree

6 J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

Programme started in 1973 and is still attracting students from all over the world. Over

250 MPM degrees have been awarded since its inception.

The Province’s newest University of Northern BC in Prince George opened in

September 1994 and one of the first faculty appointed was a former J.H. Borden student,

the forest entomologist and inventor, B.S. Lindgren.

By 1951, the predominantly British Victorian naturalists, who had collected and listed

the insects in many different groups during the first 50 years, were being replaced, as

Marshall (1952) put it, by Canadian ‘spray blokes’.

The Dominion Department of Agriculture was then well represented in the Province.

The oldest laboratory was the original Experimental Farm Station set up in Agassiz in

1886 by T. Sharpe with the aim of serving the farming community and developing our

embryonic agricultural industry. Many of the ornamental trees and shrubs he planted, after

clearing the forest at the turn of the century, still stand impressively around the new

research building that opened there in 2001. R. Glendenning was in charge of the

entomology laboratory at Agassiz for 37 years until his retirement in 1953 and in 1956 a

substation for soft fruit was set up in Abbotsford.

The Plant Protection Laboratory in Vancouver had also been running for many years.

Vancouver was the site of a Stored Product Insect Laboratory and a Biological Control

Investigations Laboratory with J.H. McLeod from the then Dominion Parasite Laboratory

in Belleville in charge. Turnbull and Chant were both employed there after they graduated

from UBC in the 1950s. A fourth laboratory, for Plant Pathology, was already on the UBC

campus and a Dominion Field Crop Insect Laboratory was opened there in 1955.

The Dominion was represented in the Interior by Livestock Insect and Field Crop

Insect Laboratories in Kamloops and a Fruit Insect Laboratory in Summerland. There was

another Field Crop Insect Laboratory in Victoria. The Kamloops Research Station was

directed by R.H. Handford from 1962 to 1970. He was the western expert on the biology

and control of grasshoppers since the death of N. Criddle and had a wealth of experience in

their management (Riegert 1980).

In 1960 a new Research Station was opened on the UBC campus. One of its main

functions was to act as a National Plant Virus Research Laboratory, but the outlying Plant

Pathology, Stored Product and Field Crop Insect Laboratories together with a substation in

Chilliwack were brought together in a seven-man Entomology Section under H.R.

MacCarthy. The station had modern rearing facilities, greenhouses, library and an

administrative section. After some 30 years the Department of Agriculture closed the

Vancouver Research Station in 1996 and the entomologists moved to Agassiz and

Summerland.

Entomology in the Province was directed in 1951 from the Vernon courthouse by the

Provincial Entomologist. C.L. Neilson, a talented field man and administrator, was in

Vernon at that time and by 1955 had replaced E.R. Buckell’s associate I.J. Ward as

Provincial Entomologist after Ward’s untimely death. Later the headquarters moved to,

and remains in, Victoria.

J.C. Arrand was appointed Assistant Provincial Entomologist in 1957 and maintained

entomological continuity in the Ministry, retiring as Director of the Crop Protection

Branch in 1987. During his tenure, the Province was involved in research into alternatives

to organochloride insecticides. Several crop pests had developed resistance to DDT by that

time and secondary pests were emerging as the populations of beneficial insects were

reduced by the ‘pesticide treadmill’. Several mishaps in the 1960s persuaded the Social

Credit government of the time to order the Department of Agriculture to draft regulations

for the Pharmacy Act in 1965 and to establish a new Analytical Laboratory for pesticides.

In 1963, the Provincial Entomology and Plant Pathology Branches and the Field Crops

Branch moved to larger buildings in Cloverdale and there were similar but smaller

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 7

laboratories in Kamloops, Sidney, Summerland and Victoria. In 1995 the Provincial

Laboratories in Cloverdale were closed and most entomologists and their programs moved

to modern facilities in the Abbotsford Regional Office.

Forest insect pests were managed by the Canada Department of Agriculture (Science

Service) in 1951, and H.A. Richmond, who had carried out forest insect surveys on

horseback for R. Hopping in Vernon in the 1920s, was in charge of the Victoria laboratory

until 1955. In the interior, the laboratory in Vernon, headed by R. Hopping for two decades

solved many of our urgent forest pest problems. D.A. Ross became its head in 1955, but

moved to Victoria in 1970 when the Vernon laboratory closed. L.H. McMullen moved to

Victoria from Vernon in 1955 and became head of entomological research there in 1965S.

Many other well known entomologists have worked in these laboratories and more details

are given elsewhere in this Volume.

In the 50" ESBC Anniversary volume, Spencer (1952) summarised the ‘status of our

knowledge of the insects of British Columbia’. He pointed out that federal officers of the

Division of Entomology sent their specimens to the Canadian National Collection in

Ottawa and that complete lists of Provincial records would be almost impossible to obtain.

Many of the specimens of the more colourful Lepidoptera, Hemiptera and Coleoptera were

divided between the National and Provincial collections with most of Spencer’s own

collection at ‘The University’. R. Hopping’s “huge” beetle collection from Vernon was left

to the California Academy of Science where H.B. Leech went, taking his own collection of

water beetles with him. At the Provincial Museum in Victoria, G. Hardy, the

Botanist/Entomologist, collected Lepidoptera and Coleoptera until his retirement in 1959

but then the collections were almost completely neglected until 1970 when a severe insect

infestation was discovered. B. Ainscough volunteered to look after the collection until a

full-time curatorial division was established. R.H. Carcasson took over as Senior Curator

in 1973 and the collection continued to recover as he and later R.A. Cannings, were in

charge of it. Two museum handbooks were published dealing with insects of the Province,

‘The Dragonflies’ (#35) by R.A. Cannings and K.M. Stuart (1977) and ‘The Mosquitoes’

(#41) By P. and E.M. Belton (1983).

Our Journal, apart from its name, has not changed greatly. The 50" ESBC Anniversary

volume had 17 research papers (as well as the seven retrospectives) in 104 pages. It had 5

full-page advertisements and eight smaller ones, some of which were congratulations. It

was typeset, printed on good quality paper, stitched in ‘signatures’ and ‘perfect bound’ in

‘sugar-bag blue’ card by Chapman and Warwick in Vancouver. The latest issue, Vol. 97,

2000, had 16 research papers in 105 pages. It was laser printed, not typeset, and then

trimmed to size (‘camera ready’ apart from a few figures) by the Editor and printed on

glossy coated paper using photo-offset by the Reprographics Unit at Simon Fraser

University. It is also ‘perfect bound’ in blue, but is without advertising. Stages in this

evolution included green and even a white, stapled cover. R.A. Ring, introduced computer

printing in 1992, when the design and printing was done by the Graphics Group at the

University of Victoria. The scientific quality and reputation of the Journal has remained

one of the highest in Canada.

Our Society also continues to attract enthusiastic and dedicated entomologists (Riegert

1991) and a list of its presidents during the second half-century is appended. On June 14th

1951, 53 members were photographed at the 50" annual general meeting at UBC and a

similar number attended our 100" in September 2001 at the Summerland Research Station

(Figs. 1,2). A handful of the 1951 veterans maintain an interest in entomology but none

attended the 100" meeting. The proportion of student members attending has increased

greatly over the years, D.A. Chant and L.E. Wade may have been the only graduate

students there in 1951 but today, with four Universities and many Regional Colleges, we

DECEMBER 2001

J. ENTOMOL. SOC. BRIT. COLUMBIA 98

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10 J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

can expect them to make up almost half of those registering. The last issue of Boreus, the

Society’s newsletter, first issued in a formal binding by R.A. Cannings in April 1981 was

mailed in June 2001 to 160 ‘more or less paid up’ members according to our Secretary-

Treasurer.

REFERENCES

Belton, P. and E.M. Belton. 1983. The Mosquitoes of British Columbia, Handbook No.41, British

Columbia Provincial Museum, Victoria. 189pp.

Cannings, R.A. and K.M. Stewart. 1977. The Dragonflies of British Columbia, Handbook No.35, British

Columbia Provincial Museum, Victoria. 254pp.

Marshall, J. 1952. Applied entomology in the orchards of British Columbia, 1900-1951. Proceedings of the

Entomological Society of British Columbia 48: 25-31.

Riegert, P.W. 1980. From Arsenic to DDT, A History of Entomology in Western Canada, University of

Toronto Press. 357pp.

Riegert, P.W. 1991. Entomologists of British Columbia, Friesen Printers, Altona MB. 90pp.

Spencer G.J. 1952. The 1951 status of our knowledge of the insects of British Columbia. Proceedings of

the Entomological Society of British Columbia 48: 36-41.

APPENDIX

Presidents of the Entomological Society of British Columbia 1951 - 2001

YEAR PRESIDENT = AFFILIATION

1951 GJ. Svencer Zoology, University of British Columbia, Vancouver

1952 H.Olds Agriculture Canada, Vancouver

1953. J. Marshall Agriculture Canada, Summerland

1954 J.H. McLeod Agriculture Canada, Vancouver

1955 M.H. Hatch University of Washington, Seattle

1956 R.H.Handford Agriculture Canada, Kamloops

1957 _-H. Andison Agriculture Canada, Victoria

1958 M.D. Proverbs Agriculture Canada, Summerland

1959 P 22K Agriculture Canada, Vancouver

1960 __—D.A. Ross Canada Dept. of Forestry,

Forest Entomology Laboratory, Vernon

1961 H.R. MacCarthy Agriculture Canada, Vancouver

1962 CL. Neilson BC Department of Agriculture, Victoria

1963 D.P. Pielou Agriculture Canada, Summerland

1964 ~R.R. Lejeune Canada Dept. of Forestry, Forest Biology Laboratory, Victoria

1965 M.G. Thompson Canada Dept. of Forestry, Forest Biology Laboratory, Victoria

1966 ‘J.C. Arrand BC Department of Agriculture, Victoria

1967 GG.E. Scudder Zoology, University of British Columbia, Vancouver

1968 _F.L. Banham Agriculture Canada, Summerland

1969 ~H. Madsen Agriculture Canada, Summerland

1970 W.T. Cram Agriculture Canada, Vancouver

1971 D.G. Finlayson Agriculture Canada, Vancouver

1972 R.A. Ring Biology, University of Victoria, Victoria

1973. J. A.Chapman Canada, Dept. Fisheries and Environment,

Pacific Forestry Research Centre, Victoria

1974 R.D.McMullen Agriculture Canada, Summerland

1975 _ T. Finlayson Biology, Simon Fraser University, Burnaby

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

1976

1977

1978

1979

1980

198]

1982

1983

1984

1985

1986

1987

1988

1989

1990

199]

1992

£993

1994

1995

1996

1997

1998

1999

2000

2001

J.R. Carrow

H.S. Gerber

A.L. Turnbull

P. Belton

R.H. Elliot

A.R. Forbes

L. Safranyik

J.A. McLean

R.A. Ring

N.P. Angerilli

R.A. Cannings

B.D. Roitberg

M.B. Isman

C.S. Guppy

D.A. Raworth

J.E. Cossentine

R.S. Vernon

T.L. Shore

S.M. Fitzpatrick

L.A. Gilkeson

G.S. Anderson

D.R. Gillespie

B.S. Lindgren

M.B. Isman

Canada, Department of Environment,

Pacific Forestry Research Centre, Victoria

BC Ministry of Agriculture, Cloverdale

Biology, Simon Fraser University, Burnaby

Plant Science, University of British Columbia, Vancouver

Agriculture Canada, Vancouver

Canada, Department of Environment,

Pacific Forestry Research Centre, Victoria

Forestry, University of British Columbia, Vancouver

Biology, University of Victoria, Victoria

Agriculture Canada, Summerland

BC Provincial Museum, Victoria

Biology, Simon Fraser University, Burnaby

Plant Science, University of British Columbia, Vancouver

Royal BC Museum, Victoria

Agriculture Canada, Vancouver

Agriculture Canada, Summerland

Agriculture Canada, Vancouver

Forestry Canada, Pacific Forestry Centre, Victoria

Agriculture Canada, Vancouver

BC Ministry of Environment, Victoria

Biology, Simon Fraser University, Burnaby

Agriculture and Agri-Food Canada, Agassiz

Biology, University of Northern BC, Prince George

Faculty of Agricultural Science,

University of British Columbia, Vancouver

N.N. Winchester Biology, University of Victoria, Victoria

R.A. Cannings

Royal BC Museum, Victoria

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J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 13

The G.J. Spencer Memorial Lecture Series

at the University of British Columbia

A landmark in the past 35 years has been the G.J. Spencer Memorial Lecture series at

the University of British Columbia. This series of lectures was to commemorate the

achievements of G.J. Spencer, and ran from 1967 to 1999. The 33 lectures by eminent

entomologists exposed faculty, students and guests to some of the major entomological

research accomplishments in the world. The following gives a brief note on Spencer

attached to the annual brochure circular each year, the list of lectures and their titles.

Prof. Emeritus George Johnston Spencer was born of missionary parents in Yercaud,

South India, January 18, 1888 and died at his home in Vancouver, Canada, July 24, 1966.

Prof. Spencer was renowned as a teacher. In 1924 Prof. Spencer was appointed Assistant

Prof. at the University of British Columbia, and in 1945 Prof. at the same Institution.

Retiring in 1953 he was elected Prof. Emeritus, Special Lecturer and Curator of the

Entomological Museum.

In the early years of the University of British Columbia on its Point Grey campus, Prof.

Spencer played a major role in establishing the Department of Zoology and his particular

pride was the fine Entomological Museum that he established and which now bears his

name. When he came to the University there were "less than a handful" of unlabelled

specimens; when he left in 1966 the Museum contained over 300,000 specimens of

perfectly mounted and labelled insects belonging to all orders. As a scientist he directed

much of his energy to assembling a representative collection of the insect fauna of British

Columbia. Prof. Spencer was a prodigious collector, even when ailing in the early months

of 1966. His favourite study area was always the Dry Belt of British Columbia, an area that

he insisted was "God's Own Country".

1967 | Prof. Sir V.B. Cambridge University Jan Swammerdam, preformation and insect

Wigglesworth growth.

1968 | Prof. H.A. Case Western Reserve Control systems in insect development.

Schneiderman University

1969 | Prof. K.D. Roeder Tufts University Sonar and countersonar; the interaction of bats

and moths.

1970 | Prof. G. Hoyle University of Oregon Neural mechanisms underlying the behaviour of

invertebrates.

1971 | Prof. Th. Rockerfeller University Genetics of behaviour in Drosophila.

Dobzhansky

1972 | Prof. M. Locke University of Western Insect cells, and the study of basic problems in

Ontario cell biology.

1973 | Prof. L.P. Brower Amherst College Experimental proof of the palatability spectrum

in natural populations of the monarch butterfly.

1974 | Prof. V.G. Dethier Hunger in the blowfly; a physiological analysis.

1975 | Prof. D. Pimentel Cornell University The Economy of Insect Population.

1976 Prof. C.M. Williams Harvard University Hormones, Genes, and Metamorphosis.

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

1977 | Prof. F.J. Ayala University of California, The Genetics of Speciation: a Study with

Davis Drosophila.

1978 | Prot. T.R.E. Imperial College,

Southwood University of London

1979 | Prof. F. Engelmann University of California, Production of a yolky egg; aspects of hormonal

Los Angeles control.

1980 | Prof. E.B. Edney

Population Biology of Checkered-Spot

Prof. P. Ehrlich

Butterflies: testing a theory in the field.

1982 | Prof. E. Bursell University of Bristol The relationship of the tsetse fly and its host.

1983 | Prof. G. Dover Cambridge University Molecular drive and the origin of insect species.

1984 | Prof. K.G. Davey York University Sex among the arthropods.

1985

—"

Some Patterns of Nature.

University of British

Columbia

Water balance in land Arthropods: some

problems and solutions.

Stanford University

Prof. J.S. Edwards University of Washington | Origin of flight in insects: an exercise in

evolutionary neuroethology.

1986 Prof. C.S. Goodman

Stanford University Embryonic development of the insect nervous

system: the generation of neural specificity.

Prof. H. Dingle

University of California,

Davis

The genetic architecture of insect life histories.

1987

Newly-formed species: recognition and

1988 Prof. H.L. Carson

characteristics.

1989 | Prof. R.G.H. Downer | University of Waterloo

1990 Prof. J.G. Hildebrand

University of Hawaii

From semiochemical to behavior: Mechanisms

underlying pheromonal communication in

moths.

University of Arizona

Prof. J.H. Borden Semiochemicals: the essence of integrated

management of the mountain pine beetle.

Simon Fraser University

Prof. G.M. Hewitt University of East Anglia | Ice Ages, Species Substructure and the

Significance of Hybrid Zones.

1992

The Juvenile Hormone of Insects: Elixir,

1993 | Prof. G.R. Wyatt

Nemesis and Enigma.

1994 | Prof. R.D. Alexander | University of Michigan Species Problems in the Singing Insects.

1995 Prof. I.W.B.

Thornton

Queen's University

La Trobe University The recolonization of Krakatau.

Lepidopteran reproductive strategies and

changing habitat quality.

Prof. Jeremy N. Laval University

McNeil

1997 Dr. A.O. Nicholls

Conservation Evaluation, where to from here?

An Australian Perspective.

CSIRO, Australia

1998 | Prof. E. Bernays University of Arizona Why do insect herbivores specialize on plant

hosts?

Prof.G.G.E.. Scudder Insects in biodiversity conservation: some

perspectives from the South Okanagan.

University of British

Columbia

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 15

The H.R. MacCarthy Pest Management Lecture

Series (Simon Fraser University and the

University of British Columbia)

The purpose of the H.R. MacCarthy Pest Management Lecture is to present an annual

lecture by a distinguished pest management scientist or practitioner. The venue of the

lecture alternates between Simon Fraser University and the University of British

Columbia. The lecture is managed by a committee consisting of representatives from

Simon Fraser University (former Centre for Pest Management), the University of British

Columbia (Faculty of Agricultural Sciences), Agriculture and Agri-Food Canada, the

Professional Pest Management Association of British Columbia, and the Entomological

Society of British Columbia. It is funded by revenues from the H.R. MacCarthy

Endowment Fund. The following biographical note is part of the program for each year’s

lecture.

Dr. MacCarthy began his career in agricultural research in 1948 as a student assistant at

the Field Crop Insect Laboratory at Kamloops. Mac grew up in England, was an

agriculturist in Australia, a cattle rancher at Princeton, BC for 9 years, and spent nearly 6

years in war service with the Canadian Infantry Corps. After returning from war service in

1946, Mac attended the University of British Columbia, receiving his B.A. in Zoology in

1950. He went directly on to graduate study at the University of California at Berkeley

and was awarded his Ph.D. in 1953.

He returned to Kamloops and worked there until 1955 when he was appointed Officer-

in-Charge of the Field Crop Insect Laboratory on the campus of the University of British

Columbia. He was named head of the Entomology Section of the Vancouver Research

Station in 1959. Mac’s research was largely on the transmission of potato leaf roll virus by

aphids. Collaborative work by him and other scientists at the station has led to almost

complete control of potato leaf roll virus in the province.

Mac has been an adjunct professor at Simon Fraser University’s Centre for Pest

Management since 1974. Immediately following his retirement from Agriculture Canada

in 1976, he became a sessional lecturer at Simon Fraser University, and was acting director

of the Centre for Pest Management for more than two years. He also held the title of

Honorary Lecturer at the University of British Columbia for.almost two decades, and was

editor of the Journal of the Entomological Society of British Columbia for over three

decades. His specialty has always been to improve the English of thesis writers and others

who need it — a category for which he has yet to identify an exception. Mac celebrated his

90" birthday on June 22, 2001.

Affiliation Title

ol eee eet Implimentation of IPM programs — the impact

of agroecological and socio-economic

conditions

eee Texas A&M University Intelligent geographic information systems and

integrated pest management

1992 | Dr. S. Finch Horticulture Research Integrated pest management in field vegetable

International, crops — the challenge facing research scientists

Wellesbourne, U.K.

Prof. R.J. Prokopy University of Integration of management practices for insect,

Massacheusetts weed and disease pests can be viewed as a

stepwise process ultimately affected by socio-

political concerns

1994 | Prof. G. Norton University of Queensland Pragmatic economics for pest management

16 J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

Prof. W. Fry

Prof. T.C. Baker Sex pheromones of moths: promise, premise

and practice

1997 | Prof. F. Gould Evolutionary potential of crop pests

University implications for integrated pest management

Canada, Lethbridge

1995 Cornell University Re-emergence of potato late blight,

Phytophthora infestans: sex and the single

fungus

1999 | Prof. J.T. Trumble University of California, Ethics, environment and economics inIPM: a

REISS nheside | cocci

2000 | Prof. J. Rosenheim University of California, Predators that eat predators: implications for

2001 Is Solenopsis invicta, the imported fire ant, as

invincible as its specific name implies?

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 17

Insect collections, surveys and conservation in

British Columbia in the 20" century

ROBERT A. CANNINGS'!

ROYAL BRITISH COLUMBIA MUSEUM,

675 BELLEVILLE STREET, VICTORIA, BC, CANADA V8W 9W2

SYDNEY G. CANNINGS

BRITISH COLUMBIA CONSERVATION DATA CENTRE,

MINISTRY OF SUSTAINABLE RESOURCE MANAGEMENT,

PO BOX 9344 STN PROV GOVT. VICTORIA, BC, CANADA V8W 9M1

GEOFFREY G.E. SCUDDER

DEPARTMENT OF ZOOLOGY, UNIVERSITY OF BRITISH COLUMBIA,

VANCOUVER, BC, CANADA V6T 1Z4

INTRODUCTION

In this brief summary of insect collections, surveys and conservation efforts in British

Columbia during the 20" century, we emphasize the years since 1950, occasionally

referring to activities in the first half of the century for historical perspective. Because of

restricted space, our intent here is to stress accomplishments rather than the historical

aspects of entomologists and their work.

Researchers and managers in many fields of biology have recognized that invaluable

information on biological diversity is contained with specimens in natural history

collections. Such collections are also crucial in education, essential references for

identification of specimens, and critical for studies in environmental biology, ecology,

evolution and other fields. Consequently, there is an effort in many parts of the world to

emphasize the continued and future importance of collections and to make specimen-based

information available on the Internet to researchers and others (Bisby 2000; Edwards et al.

2000). Collections of British Columbia insects are an important source of information

relevant to a number of provincial initiatives in the assessment and conservation of our

biodiversity. Miller (1985, 1993) and the Biological Survey of Canada (1991) emphasize

the importance of biological collections in these and other roles. The Biological Survey of

Canada (Terrestrial Arthropods) promotes, develops and coordinates national initiatives in

systematic and faunistic entomology. It has published several briefs outlining guidelines

for successful arthropod surveys (Biological Survey of Canada 1994, 1996).

Over the years, many biological surveys have been undertaken in the province.

However, there has been little attempt to document those inventories that involve insects

and ascertain where the results from these might be located. Indeed, it is often unclear if

voucher or residue material from these studies are available for further study or

verification of records. Furthermore, we must maintain provincial initiatives in collection

growth at a time when funding is being curtailed. Ironically, with the decline of funding

for taxonomic research, insects collected in association with various fish/forestry

interaction programs may be our best, and perhaps only, source of new records of British

‘Authors are in alphabetical order, not necessarily in order of the importance of their

contribution.

18 J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

Columbia insects in the future. All faunal surveys conducted in the province should be

required to submit voucher specimens to a museum collection (Miller and Nagorsen 1992)

before the project is considered complete. In this way, identifications can be confirmed and

valuable records made part of the developing biodiversity database. Guidelines for the

preparation and deposition of such collections are part of the provincial government’s

Resources Inventory Committee standards (Resources Inventory Committee 1999).

This paper summarizes the current situation with respect to collections of BC insects. It

also outlines some of the major surveys that have been undertaken, and notes a few of the

major results. The last section details some of the ongoing biodiversity conservation

initiatives that need entomological input.

COLLECTIONS

Many collections around the world contain insects collected in BC, but only a few have

large holdings of specimens from the province. Scudder (1996) gives a more

comprehensive list of collections housing provincial material. In general, the larger

collections containing BC specimens, both inside and outside the province, are dominated

by Coleoptera and Lepidoptera material, although some other orders, such as Hemiptera

and Odonata, are well represented in the province’s collections. Of the larger orders,

Diptera and Hymenoptera show the most serious gaps in taxonomic coverage, and the

apterygote taxa are poorly covered.

Provincial collections contain limited type material, especially holotypes. Most type

specimens of BC taxa are housed in larger collections, especially the Canadian National

Collection of Insects, Arachnids and Nematodes in Ottawa.

COLLECTIONS IN BRITISH COLUMBIA

Royal British Columbia Museum (RBCM)

The Royal British Columbia Museum’s entomology collections in Victoria began

accumulating with the Provincial Museum’s establishment in 1886. E.H. Blackmore, a

well-known lepidopterist, volunteered as curator from 1913 to 1928. Through the first half

of the 1900s, holdings grew through gifts of specimens and the energetic collecting of staff

members E.M. Anderson (1903-1916) and G.A. Hardy (1924-1928/ 1941-1953). Although

Hardy was primarily a botanist, his insect collections on southern Vancouver Island,

especially of Lepidoptera and Coleoptera, formed the backbone of the collection. The

collections were never fully organized and properly stored, however, and between Hardy’s

retirement in 1953 and the arrival of the collection’s first full-time curator, R.H.

Carcasson, a lepidopterist (1973-1978), they suffered considerable neglect and damage.

Carcasson was assisted by B.D. Ainscough (1972-1983) and A. Mackie (1974-1975). R.A.

Cannings, the present curator, succeeded Carcasson in 1980. C.S. Guppy (1987-1993) and

D.C.A. Blades (1997-present) have served as collections managers.

The RBCM has approximately 250,000 specimens; about 55,000 of these are

Lepidoptera, 60,000 Coleoptera and 35,000 Diptera. Cannings’ specialty, the Odonata,

number 35,000. Important collections included are those of A.W. Hanham (Lepidoptera

and Coleoptera), G.O. Day (Lepidoptera), T.A. Molliet (Lepidoptera), G.A. Hardy

(Geometridae, Noctuidae and Cerambycidae), G. Straley (Lepidoptera), F.C. Whitehouse

(Odonata) (in part), J. Grant and R. Guppy. The RBCM has a small collection of fossil

insects, mainly from the Eocene shales of the Interior.

Spencer Entomological Museum (SEM)

This teaching and research museum is located in the Department of Zoology at the

University of British Columbia (UBC) in Vancouver. It was founded in 1953 on the

aie

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 19

retirement of G.J. Spencer, and is named after this outstanding teacher, prodigious

collector and investigator of insect biology. Currently the museum contains 600,000

specimens (500,000 pinned, 75,000 alcohol preserved, 25,000 slide mounted) in 110 12-

drawer metal cabinets.

As well as containing specimens from student collections, and voucher material

associated with UBC student theses, the SEM houses some of the most important private

collections made in the province over the years. These include the Buckell, Cannings (pre-

1980) and Whitehouse (in part) collections of Odonata; the Buckell and Spencer

collections of Orthopteroid insects; the Downes and Scudder collections of BC Hemiptera

(the Alice McDouglass collection of aphids is on permanent loan to the CNC, see below);

the Spencer collection of Phthiraptera and Siphonaptera slides; the Blackmore, Kimmich

and Llewellyn Jones collections of Lepidoptera; the R. Guppy (in part) and Stace-Smith

collections of Coleoptera (including many of the Hatch types described from the latter

collection); and the Foxlee collection of Diptera and Hymenoptera. The museum also

holds, on permanent loan, the small insect collection formerly maintained by the

Vancouver City Museum.

G.G.E. Scudder has been the academic curator since 1965. Curators in the museum

have included R.A. Cannings, S.G. Cannings, K.M. Stuart and J. van Reenen; K. Needham

is the current curator. R. Kenner has been a volunteer since 1995. Because of budget

retrenchments, the museum has been closed to the public since 1993.

Pacific Forestry Centre

The collection of the Canadian Forest Service, Pacific and Yukon Region, is housed at

the Pacific Forestry Centre in Victoria. This collection was built mostly through the efforts

of the Forest Insect and Disease Survey (FIDS) during the more than 50 years of its

existence. Some of the specimens came from the federal laboratory in Vernon, which was

amalgamated with the Victoria laboratory in 1969. Recently, the surveys of forest canopy

biodiversity made by the University of Victoria have provided many new accessions. The

Pacific Forestry Centre collection consists primarily of forest species. It is particularly well

represented in Lepidoptera, bark and wood-boring Coleoptera, sawflies and hymenopteran

parasitoids and predators of these groups. The collection holds approximately 100,000

specimens of 7,000 species.

D. Evans curated the collection from 1949 to 1985, assisted for much of this time by D.

Ruppel. More recently, the collection has been maintained by B. Duncan, L. Humble and

J. Seed. Entomologists associated with the Vernon collection before its assimilation

included D. Ross and J. Grant.

British Columbia Department of Agriculture

The BC Department of Agriculture in Kelowna contains a small collection of insects

relevant to agriculture in BC.

Agriculture and Agri-Food Canada

The AAFC centre at Agassiz also houses a small collection of insects relevant to BC

agriculture. Some of these collections were formerly at AAFC stations in Vancouver and

elsewhere in BC.

Other University Collections

Small insect collections are also contained in the Department of Biology at the

University of Victoria, the Department of Biological Sciences at Simon Fraser University

in Burnaby, and in the Department of Natural Resources at the University of Northern

British Columbia in Prince George. These collections have specimens obtained during

20 J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

teaching assignments and voucher material associated with research theses in these

universities.

Private Collections

Several people in the province maintain collections that contain important material for

documenting insect distribution and biology in the province. The most striking of these are

the large Lepidoptera collections of C. Guppy, N. Kondla, J. Shepard and J. Troubridge.

These collections made important contributions to the data analyzed and mapped in the

recently published ‘Butterflies of British Columbia’ (Guppy and Shepard 2001). The

extent and significance of private collections in the province is unknown, and an initiative

to document these resources would be useful.

COLLECTIONS OUTSIDE BRITISH COLUMBIA

Many other collections in Canada, the USA and elsewhere contain specimens of BC

insects (Scudder 1996). Undoubtedly the most important is the Canadian National

Collection of Insects, Arachnids and Nematodes at Agriculture and Agri-Food Canada in

Ottawa. This contains not only the abundant material collected in BC by government

scientists and technicians over the years, but it also houses the Ricker collection of

Plecoptera and the Glendenning and McGillivray collection of aphids, and much of the

Forbes and Chan collection of these insects.

BC specimens are also held by the Lyman Entomological Museum at Macdonald

College of McGill University in Ste.-Anne-de-Bellevue, Québec, and in the Royal Ontario

Museum in Toronto. The Hopping collection of BC Coleoptera and the H.B. Leech

collection of water beetles are in the California Academy of Sciences in San Francisco,

together with the large collection of beetles collected by D. Kavanaugh in the Queen

Charlotte Islands and elsewhere in the province. The entomological collection at Oregon

State University in Corvallis houses the Hatch collection of beetles, rich in BC specimens.

The American Museum of Natural History in New York and the National Museum of

Natural History in Washington, DC also contain much BC material. J. Bergdahl (Spokane,

WA) has a large collection of ground beetles that includes much material from BC,

especially the Kootenays, Vancouver Island and the Gulf Islands. L. Crabo (Bellingham,

WA) owns a significant collection of noctuid moths from the province.

COLLECTION DATABASES

Rapid access to the information contained in collections has become a major concern

and a large portion of the meagre resources available to collections has been directed to the

production of electronic databases. Much of the collection in the RBCM is databased and

some of the insect orders in the SEM have also been computerized. The major private

butterfly collections were databased during the production of ‘Butterflies of British

Columbia’ (Guppy and Shepard 2001).

SURVEYS

Over the years surveys and inventories have increased our understanding of BC insects

and their distribution. The important and wide-reaching FIDS initiative has already been

mentioned. Many of the private collections made in the past, and many of those being

made today, were made by collectors interested in surveying the distribution of species in

their favourite groups. Recently, entomologists and naturalists have joined in informal

insect forays (Cannings 1996), usually to some unstudied area, in an attempt to further

knowledge of insect status and distribution. However, in the last 40 years or so, several

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 5)

projects with a more or less formal survey component have been undertaken in the

province. The distinction between surveys and general collecting is often not definite, and

we have tried to include only the former. Most of these surveys are summarized below,

emphasizing the publications produced. The surveys are arranged according to the general

type of environment sampled (aquatic habitats, forests, grasslands, and so on).

In order that available resources are focused on important habitats and taxa, Scudder

(1996) recommended priorities for terrestrial and freshwater invertebrate surveys. These

included suggestions for surveys of Lepidoptera in the Nanaimo Lowlands and Okanagan

Basin, invertebrates of coastal old-growth forests, and invertebrates in caves and springs.

AQUATIC SURVEYS

Lakes

Carl (1953), Clemens et al. (1938, 1939), Rawson (1934), Robertson (1954), and

Withler (1956) surveyed insects and other benthic organism in a number of the larger lakes

in the province. Saether (1970) studied the bottom fauna of lakes in the Okanagan Valley,

paying special attention to the Chironomidae.

Scudder began surveys and detailed studies on saline lakes and other waterbodies,

including peatlands, in the Cariboo, Chilcotin and Kamloops regions in 1959. These

studies continued for 25 years. The physical and chemical limnology of most of the larger

lakes were described by Topping and Scudder (1977), and many of the smaller ponds were

characterized by Scudder (1988). Scudder (1969, 1988) listed some of the insects that

showed a differential distribution in the lakes, and the community structure in the

Coleoptera and Hemiptera was discussed by Lancaster and Scudder (1987). The

distribution of Odonata in these same lakes was examined by Cannings ef a/. (1980) and

Cannings and Cannings (1987), and the Chironomidae by Cannings and Scudder (1978).

Surveys have been an important stimulus to the development of entomological collections

in British Columbia — Rob Cannings collecting chironomid larvae during a survey of saline

lakes near Williams Lake, August 1970. Photo by Syd Cannings.

9) J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

Some insects in meromictic lakes were listed by Northcote and Halsey (1969), and the

benthic insect fauna of several lakes in the UBC Research Forest near Maple Ridge was

reported in Hindar ef al. (1988) and Rempel and Northcote (1989). Northcote et al. (1978),

Northcote and Hall (1983), Hume and Northcote (1985), Chapman et al. (1985) and

Walters ef al. (1987) also gave records of the occurrence of Chaoborus species in some

coastal and interior lakes.

Streams and Rivers

Collections of stream and river insects in BC have been made by fisheries inventory

personnel of the now-titled provincial Ministry of Sustainable Resource Management, and

members of the Federal Department of Fisheries and Oceans. Some of the streams studied,

for which samples are available, are Loon Creek near Clinton, Centennial/ Slim/Rosanne

creeks east of Prince George, Adam and Keogh rivers on northern Vancouver Island, Big

Silver Creek on Harrison Lake, Mesilinka River on Williston Reservoir, Torpy and Upper

Nechako rivers east and west of Prince George, respectively, and Takla Lake creeks north

of Fort St. James (P. Slaney, in /itt.)

Insects were also collected in two major fish/forestry interaction programs in coastal

BC: the Carnation Creek Experimental Watershed Study on the west coast of Vancouver

Island, which began in 1970 (Hartman and Scrivener 1990), and the Fish/Forestry

Interaction Program (FFIP) in the Queen Charlotte Islands, initiated in 1981. Over 30

watersheds were studied in this program (Hogan ef al. 1998). In addition, collections have

been made during environmental surveys on contract to government or private industry.

For instance, Perrin and Associates (Vancouver) worked for many years conducting

invertebrate surveys in the Nechako River for Alcan. Unfortunately, most records for

many of these studies exist only in government reports or in non-refereed documents that

are difficult to locate. The species names mentioned in such “grey literature’ may be

unreliable, since voucher specimens are seldom deposited in provincial or university

collections where they are easily accessible to taxonomic experts and where identifications

can be confirmed.

Idyll (1943) studied portions of the Cowichan River and considered it a “Trichoptera”

stream because of the density of these insects in the bottom fauna. Filmer (1964) surveyed

the mayfly fauna in the Alouette River, and Wigle and Thommasen (1990) studied this

order in the Bella Coola and Owikeno watersheds. Ricker surveyed many streams and

rivers around Cultus Lake (see aquatic insects paper), and benthic insects in the lower

Fraser Valley were listed by Northcote ef al. (1976). S. Salter (pers. comm.) has followed

Scudder’s (1996) suggestion that collections in springs should be a priority by undertaking

a preliminary inventory of invertebrates in selected warm springs and associated streams in

the province.

Reece and Richardson (2000) surveyed benthic macroinvertebrate assemblages of

coastal streams in the UBC Research Forest near Maple Ridge, continental streams in the

Merritt area, and large rivers, namely the Fraser River near Agassiz and the Thompson

River near Spences Bridge. Compared to small streams, large rivers had low invertebrate

abundance, species richness and diversity. Coastal streams were richer in species, but

Interior ones contained more individual insects.

Odonata Surveys

Despite the extensive collecting of Buckell (1938) and Whitehouse (1941) and the

collecting and small-scale inventories of the Cannings brothers (e.g., Cannings and

Cannings 1987, 1997), formal, large-scale inventories of dragonflies really were not

organized until the 1990s. From 1996 to 2001, dragonfly surveys planned by the BC

Conservation Data Centre and the Royal BC Museum (partly funded by Forest Renewal

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 23

BC, Parks Canada, and the Habitat Conservation Trust Fund) were undertaken to build

collections, to improve understanding of species status, distribution and _ habitat

requirements, and to better characterize the conservation status of species previously

considered rare. The surveys covered southern Vancouver Island and the lower Fraser

Valley; the Okanagan Valley (Cannings ef al. 1998); the Peace River-Fort Nelson

lowlands; the Columbia-Kootenay region, including the mountain National Parks

(Cannings et al. 1999); the Cariboo-Chilcotin and Prince George-Robson Valley regions;

and the Mackenzie and Omineca-Fort St. James regions. These surveys, which will

continue until the whole province is covered, have added greatly to our knowledge of the

distribution and ecology of this presumably well-known group of insects.

Burns Bog and other peatlands

The future of Burns Bog, the huge raised peatland near the mouth of the Fraser River,

became a controversial issue in the late 1990s. In 1999 a preliminary survey of the insects

of the bog was undertaken as part of the Burns Bog Ecosystem Review, an assessment of

the habitat’s value as a potential protected area. The results (Kenner and Needham 1999)

showed that some insects in the centre of the bog were obligate peatland inhabitants, liable

to be negatively affected if large areas of the bog were not conserved. Others found in the

surrounding forested habitats were more widely distributed species, including a high

proportion of introduced ones.

Insect surveys in other peatlands have mainly been associated with the dragonfly

projects mentioned above. Seven of the 23 dragonfly species of management concern in

the province inhabit bogs and fens, and surveys have focused on these habitats in the

regions under study (Cannings 1994, Cannings ef al. 1999). Much of the collecting done in

the Brooks Peninsula project occurred in coastal bogs (Cannings and Cannings 1997).

FOREST SURVEYS

The most significant, long-term survey of the forests of BC is the Forest Insect and

Disease Survey of the Canadian Forest Service. In BC, the FIDS collected specimens and

ecological data on forest insects from 1946 to 1995.

Aided by the impressive systematic monograph on the ground beetles of Canada and

Alaska by Lindroth (1961-1969), workers have undertaken several studies on carabid

diversity in various forested ecosystems in the province. The variability of this diversity

with succession and various logging and silvicultural practices has been stressed, largely

because much of the financing for these studies has come from Forest Renewal British

Columbia, a fund established in the 1990s to sustain the forest industry. Craig (1995)

investigated carabid community structure in a chronosequence in the dry east coast

Vancouver Island subzone of the Coastal Western Hemlock Zone. The litter spiders from

this same pitfall trap study were studied by Brumwell (1996). Peak diversity in these taxa

occurred in regenerating (3-8 year old) forest (Brumwell et al. 1998). The BC

Conservation Data Centre sponsored a survey by J. Bergdahl of the rare carabid beetles in

old-growth forests on Vancouver Island; results are not yet fully compiled.

McDowell (1998) examined ground beetle diversity in the Engelmann Spruce-

Subalpine Fir (ESSF) and Interior Cedar-Hemlock (ICH) zones near East Barriere Lake.

The ESSF forest sites had more individuals, but fewer species, than the ICH sites. Logging

had a positive impact on generic and species diversity, but a negative impact on total

number of individuals. Carabids in the ESSF zone were also studied by both Lemieux

(1998) and Lavallee (1999), who studied the response of the carabid community to

prescribed logging practices, in the Copper River Valley near Smithers, and at Sicamous

Creek, respectively. Both studies showed a peak diversity following regrowth after

logging. The Sicamous Creek insect surveys, largely undertaken by D. Huggerd, were part

24 J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

of a larger, interdisciplinary investigation of the effects of logging on the subalpine

ecosystem.

J. Jarrett (pers. comm.) investigated the much more diverse carabid community in the

Interior Douglas-fir (IDF) zone at Opax Mountain, studying both the wet (IDFdk) and

dryer (IDFxh) subzones. He has shown that there is much more diversity in the latter

subzone, much of it shared with the adjacent grassland habitats. S. Carlson (pers. comm.)

has documented the aerial dispersing Coleoptera fauna in the IDF zone at its northern

limits near Fort St. James. In a comparison of the beetle fauna attracted to non-pheromone

and Douglas-fir beetle pheromone baited traps, she found that a vast array of non-target

beetles are attracted to the latter. In the course of this research, many species of Coleoptera

new to BC were discovered.

Inventories of Collembola in forest soils have greatly increased our knowledge of the

diversity and status of this important group. Vlug and Borden (1973) reported that the

Collembola fauna was reduced by logging and slash burning near Maple Ridge. Marshall

et al. (1990) surveyed springtails in forest nurseries and found 22 species, 10 of which

were new to the province; observations on pest species were reported. Battigelli et al.

(1994) examined the soil fauna in adjacent stands of old-growth Western Redcedar-

Western Hemlock and Amabilis Fir-Western Hemlock forests on northern Vancouver

Island and found the relative abundance of Collembola was equal in both types. In the

Interior, Nadel (1999) studied the fauna of soils and litter in subalpine ecosystems as part

of the interdisciplinary Sicamous Creek project.

Marshall (1993) compared the soil fauna of Coastal Douglas-fir, Interior Douglas-fir,

Subalpine Fir-Engelmann Spruce and Coastal Western Hemlock forests and showed that

the highest densities of Collembola occurred in hemlock forests. Addison et al. (1998)

studied the diversity and abundance of microarthropods in successional Douglas-fir forests

on Vancouver Island. The same species tended to occur in all seres studied; differences

were mostly in relative and absolute abundance. Setaéla and Marshall (1994), Setala et al.

(1995) and Marshall er a/. (1998) studied the succession of springtails in tree stumps at the

same study sites. Seventy-two species were identified; some of these were not found in the

regeneration sere but most were either positively correlated with stand age or were

ubiquitous. Berch ef al. (2001) examined the diversity and abundance of springtails in the

wettest subzone of the Coastal Western Hemlock Zone near Franklin River on Vancouver

Island. In a comparison of habitats based on tree species, Sitka Spruce cover had the

highest average number of species (21) and the highest densities (32,000/m?).

On the Gulf Islands, Scudder surveyed insects in forested areas on the north end of

Galiano Island; sweeping, beating, and window-intercept and pitfall traps were used.

As an adjunct to the ambitious survey of the insect fauna of the Yukon, mostly in the

1980s (Danks and Downes 1997), field parties on their way to the Yukon collected

extensively in northern BC. Much of this collecting was in forested areas, but northern

grassland and aquatic sites were also sampled.

In the 1990s, working in the canopies of Sitka Spruce forests in the Carmanah Valley

on Vancouver Island, Winchester and his colleagues found more than 300 new arthropod

species, many of which are restricted to habitats found only in these ancient forest treetops

(Behan-Pelletier and Winchester 1998, S.A. Marshall and Winchester 1999, Winchester

and Ring 1999). This work has been expanded to other biogeoclimatic zones such as the

subalpine forests at Mount Cain on Vancouver Island (Winchester and Fagan 2000).

Studies continue on species life cycles and factors that influence the distribution,

abundance, organization and ecological importance of these aerial communities

(Winchester and Ring 1999). This pioneering canopy and conservation work demonstrates

that loss of unique canopy microhabitats may cause local species extinctions.

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 D5

GRASSLAND AND SHRUB-STEPPE SURVEYS

Over the years, considerable collecting has been done in the interior grasslands by

Buckell, the Cannings brothers, Guppy, Spencer and Scudder. Much of this work occurred

in the Cariboo, Chilcotin, Kamloops, Merritt and Hat Creek regions, and results have been

published in various faunistic (e.g. Scudder 1993) and systematic papers. More recently,

there has been considerable insect sampling in the South Okanagan, much of this

associated with the conservation of the potentially rare and endangered species and

habitats there.

Cannings (1989) made a nine-year study of the robber flies of Festuca grasslands near

Penticton. As part of the South Okanagan Conservation Areas Program, and sponsored by

the Royal BC Museum, Blades and Maier (1996) published the results of a survey of

grassland and montane arthropods around Mount Kobau carried out in the summer of

1991. A sampling transect from low to high elevation produced 1101 species; 12 of these

were new to Canada, 12 were new to BC and two were undescribed. Subsequently, a

collaborative study of the impact of livestock grazing on the Antelope-brush (Purshia

tridentata (Pursh) DC) community was initiated; in 1994-95, Scudder studied the ground-

dwelling arthropods at nine sites between Osoyoos and Vaseux Creek. The sites had

different livestock grazing histories, and analyses to date show that some of the species at

risk are affected by grazing, while others are not (Scudder 2000). Also, while Heteroptera

diversity varies with grazing impact, ant (Heron 2001) and orthopteroid insect (S. Liu,

pers. comm.) diversity does not.

In 1996, pitfall trap sampling of ground-dwelling arthropods was extended to the

Chopaka and White Lake areas to examine diversity in habitats potentially suitable for the

endangered Sage Thrasher. Insects were also surveyed in areas inhabited by Burrowing

Owls, and their crop pellets were analyzed; they contained a high proportion of beetle

remnants, especially parts of carabid, silphid and tenebrionid beetles.

In 1997-1998 pitfall trapping was continued in and around the South Okanagan to

document the species of management concern that were actually confined to the South

Okanagan Valley. Most were found not to occur outside the valley.

The shrub-steppe at the Desert Centre in Osoyoos has been sampled to determine

changes in the ground-dwelling arthropod fauna associated with both the removal of

livestock grazing and attempts at habitat restoration. Samples were taken during two years

of grazing as well as after livestock were removed in 1998.

Following a number of years of pitfall trapping of arthropods associated with the

Antelope-brush community on the Haynes Lease Ecological Reserve near Osoyoos, first

by S.G. Cannings and then Scudder, Scudder has studied the recovery of the arthropod

fauna in this community following its virtual destruction by fire on 9 July 1993. Scudder

(2001) reported the fallout of airborne insects onto the reserve in the first three weeks after

the fire; the highest rate was 1.768 billion/km?/24 hrs, recorded on 22-23 July.

OTHER SURVEYS

Brooks Peninsula and other Coastal Surveys

Cannings and Cannings (1997) reported on the terrestrial arthropods collected during

the RBCM’s 1981 interdisciplinary expedition to study the presumed ice age refugium of

the Brooks Peninsula, on the northwest coast of Vancouver Island. Over a two-week

period in August, 420 species of insects in 15 orders and 139 families were collected. In

addition, 34 species of spiders and 22 of oribatid mites were identified. The project found

31 species and 4 genera new to science.

The RBCM has organized other, smaller interdisciplinary surveys to the north coast. In

1950, G.C. Carl, then the Director of the Provincial Museum, along with G.A. Hardy and

26 J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

other colleagues visited the Scott Islands off northern Vancouver Island to study the

biogeography of these remote sites (Carl et a/. 1951). They collected a small number of

insects and other terrestrial invertebrates. In 1987 the RBCM visited Zayas and Dundas

islands near the Alaska border, and the Tatshenshini River drainage in 1992.

Since the 1980s, there has been some specialized collecting on the Queen Charlotte

Islands by R.A. Cannings, G.G.E. Scudder and others. D.H. Kavanaugh (California

Academy of Sciences), in particular, has studied the carabid beetle fauna of the islands,

first visiting them, accompanied by D.H. Mann, in July 1981. In 1986 he joined Scudder

and other biologists on a survey of the biota of mainland and island localities between

Vancouver and Prince Rupert. This coastal expedition aimed to confirm that the species

endemic to the Queen Charlotte Islands were actually confined to the islands; most were

found to be so restricted. In a monograph on the ground beetles of the Queen Charlottes,

Kavanaugh (1992) listed collection data and assessed the composition, affinities and origin

of the fauna.

Lepidoptera Surveys

In 1995, the BC Conservation Data Centre organized butterfly surveys to document the

status of these insects in two conservation hot spots -- southeastern Vancouver Island

(Shepard 1995) and the Okanagan-Similkameen valleys (S.G. Cannings, pers. comm.).

Subsequently, Shepard studied the butterflies of the Peace River Lowlands (Shepard 2000)

and Kondla (1999) surveyed the butterflies of south-facing slopes along the Pend d’Oreille

River.

Fischer et al. (2000) conducted a major survey of the macrolepidoptera of the Cariboo-

Chilcotin grasslands and grassland-forest interface, and reported an impressive 538

species. This is 96 per cent of the estimated total number of species in the study area. A

voucher collection of over 2500 specimens was deposited at the RBCM.

These surveys contributed to the data analyzed in the ‘Butterflies of British Columbia’

(Guppy and Shepard 2001), the definitive work on the butterflies of the province.

Cave Surveys

Caves often contain little-known, rare and endemic species and are often threatened by

groundwater changes and other disturbances. A 1995 survey initiated by the BC

Conservation Data Centre examined caves on Vancouver Island (S.G. Cannings, pers.

comm.). Results are not fully compiled. The first females of Parasimulium melanderi

Stone were discovered; these rare simuliids throw light on the origins of the black fly

family (Borkent and Currie 2001).

CONSERVATION

The history of insect conservation in BC is short. The first published reference to

endangered insects in the province may be Scudder’s (1980) symposium presentation on

the Osoyoos Arid Biotic Area, in which he listed some representative invertebrates along

with vertebrates and plants confined to this endangered ecosystem and emphasizes the

need for conservation of all of these populations. Later, he prepared a preliminary list of

the arthropod species that might be at risk in the South Okanagan (Scudder 1992).

Cannings (1990) discussed the diversity of insects on a provincial scale, outlined the

problems in determining conservation risk for them, and presented a short sample list of

potentially endangered or threatened species. Later, Guppy et al. (1994) and Guppy and

Shepard (2001) listed species of butterflies and skippers of conservation concern in the

province.

A major problem in developing defensible lists of invertebrates of conservation

concern is the lack of comprehensive inventories. Even in supposedly well-known groups

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 27

such as butterflies or dragonflies, species known from only one or two localities may be

subsequently discovered to be widespread, or at least much more common than the

previous collection records had indicated. But with very limited resources, the common

question is, where should we start?

To help address this question, Scudder was contracted by the provincial government to

develop a list of inventory priorities, and out of this work two publications emerged. One

(Scudder 1994) was an annotated list of 818 terrestrial and freshwater invertebrates that,

on the basis of limited known occurrence or restriction to obviously endangered habitats or

ecosystems, were potentially rare and/or endangered. This list included 168 species

endemic to the province (based on collection information at that time) and an additional

203 species restricted to BC in Canada. The other publication (Scudder 1996) gave a list

and discussion of inventory priorities, a discussion of sampling methods and resources

needed, a list of taxonomic experts, and a series of annotated lists that divided the species

noted in Scudder (1994) by ecoprovince.

THE RED AND BLUE LISTS

The Red and Blue lists of species of conservation concern were originally developed

for vertebrates by the Wildlife Branch in the provincial Ministry of Environment, Lands

and Parks. The British Columbia Conservation Data Centre (CDC) began in 1991 as a

cooperative venture of the BC Ministry of Environment, Lands and Parks, The Nature

Conservancy (US), and the Nature Trust of British Columbia; it is now a section within the

provincial Ministry of Sustainable Resource Management. The CDC assumed the role of

assigning provincial status ranks to not only vertebrates, but to plants, plant communities,

and invertebrates as well. The CDC assigns status ranks using a methodology that was

created originally by The Nature Conservancy (US) and is now used by conservation data

centres and natural heritage programs throughout North America and much of Latin

America. The provincial Red (endangered or threatened) and Blue (vulnerable) lists of

species and ecosystems are now translated directly from the CDC’s ranks; up-to-date lists

can be viewed or downloaded at the CDC’s website (http://srmwww.gov.bc.ca/

cdc/trackinglists/red_blue.htm).

All species of dragonflies, butterflies, and tiger beetles have been assigned ranks, and

Scudder’s (1994) list has been used to rank a number of other species where the status can

be confidently assigned. Currently, 69 species of insects are on the Red List and 74 are on

the Blue List. However, the problem of lack of inventories is so acute in most groups that

the majority of uncommonly collected species cannot be assigned useful ranks.

The Red and Blue lists offer no direct legal protection to any species; they simply

provide an account of the conservation status of species of concern within the province.

Insects and other invertebrates are not considered ‘wildlife’ under the provincial Wildlife

Act, so cannot be officially designated as Endangered or Threatened under that Act.

However, under two more recent pieces of legislation, the Forest Practices Code Act and

the Fisheries Protection Act, there is the provision for possible protection of the habitat of

certain endangered insects. Under the Forest Practices Code, listed invertebrates that are

deemed to be affected by forest or range practices may be designated as “Identified

Wildlife” and have management practices for them specified for certain areas. A number

of insects have been proposed for this designation, and have had preliminary management

accounts written for them (K. Paige, pers. comm.).

NATIONAL DESIGNATIONS

The Committee on the Status of Endangered Wildlife in Canada (COSEWIC) is a

national body made up of representatives of federal government agencies and _ all

provincial and territorial governments. COSEWIC has recently expanded its mandate to

28 J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

include Lepidoptera, the first insect order that it has considered. Status reports have been

accepted and designations made for six BC species: Euchloe ausonides (Lucas) insulanus

Guppy and Shepard (Extirpated), Plebejus saepiolus Boisduval insulanus Blackmore

(Endangered), Euphydryas editha (Boisduval) taylori (W.H. Edwards) (Endangered),

Satyrium behrii (W.H. Edwards) columbia (McDunnough) (Threatened), Euphyes vestris

(Boisduval) (western population, Threatened), and Danaus plexippus (Linnaeus) (Special

Concern). These designations come with no legal protection but, under the proposed

federal Species at Risk Act, insect species designated by COSEWIC as threatened or

endangered would be recommended to the federal cabinet for official designation under

that Act. National designations are detailed at the COSEWIC website at

http://www.cosewic.gc.ca/cosewic/default.cfm.

RECOVERY PLANS

Once a species is designated provincially or nationally, the next stage in its

conservation is the development of a recovery strategy. To date, no recovery plans have

been written for specific insect species. However, recovery plans for endangered or

threatened insects from the south Okanagan-Similkameen and southeastern Vancouver

Island-Gulf Islands areas are now being included in the work of two ecosystem recovery

teams: the South Okanagan Ecosystem Recovery Team (now part of the South Okanagan-

Similkameen Conservation Program) and the Garry Oak Ecosystem Recovery Team.

BIODIVERSITY MAPPING

One way to select areas for protection is to focus efforts on sites with concentrations of

species diversity and rare species. Scudder, in his research on these hotspots of richness

and rarity in the province, has assembled georeferenced databases for the known,

specialist-determined specimens of Odonata, Plecoptera, Hemiptera (Prosorrhyncha =

Heteroptera), Lepidoptera (butterflies), Megaloptera, Raphidioptera, and Neuroptera.

Databases on other groups such as carabid beetles, water beetles and aphids are being

prepared. Results obtained by mapping these data with WORLDMAP software show that

the provincial insect richness and rarity hotspots are in the South Okanagan and

southeastern Vancouver Island-Gulf Island areas. Concentrations of rare species and total

species numbers seem to coincide, and they also match similar concentrations mapped for

vascular plants and small mammals.

PROTECTED AREAS

Because there is little, if any, specific protection for insects and their habitats, the

general conservation of habitat in parks and other areas plays a crucial role in insect

conservation. Since 1991, the area of BC protected in ‘protected areas’ (that is, national or

provincial parks, ecological reserves, and other protected areas that fall under the

Environment and Land Use Act) increased from 5.74 million hectares to over 11 million

hectares—over 12 per cent of the province’s area (BC Ministry of Sustainable Resource

Management 2001). However, the ecological representation of these protected areas

remains less than ideal. Even though there was some initial effort to target ecological areas

that had been poorly represented in the past, 68 per cent of ecosections still have less than

12 per cent of their area protected (BC State of Environment Reporting 2000).

Furthermore, a number of ecosections where endangered species are concentrated, such as

the Lower Mainland and southeastern Vancouver Island, still have less than 2 per cent of

their area protected.

Data on rare insect species have contributed to the success of some conservation

efforts. Most of the grassland inventories noted in the present paper were undertaken in

conjunction with conservation planning and strategies for protecting threatened habitats in

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 29

the Okanagan Valley. Several of these important grassland areas (e.g., Chopaka, Mt.

Kobau, White Lake and Kilpoola Lake) have been preserved, either as parkland through

provincial government processes, or by the Nature Trust and other conservation

organizations. The results of the interdisciplinary Brooks Peninsula project were

instrumental in the decision to create a provincial park there, as were the findings of the

canopy studies in the Carmanah Valley. Scudder’s inventories on Bodega Ridge on

Galiano Island helped preserve that site, as did his aquatic surveys at Westwick and Rock

lakes, now ecological reserves in the Cariboo-Chilcotin. Inventories in Burns Bog were

designed to gather data for conservation purposes and, if present negotiations go well, this

critical peatland may be protected in the future.

ACKNOWLEDGEMENTS

We thank J. Addison, S. Berch, A. Borkent, D. Blades, B. Duncan, R. Kenner, V.

Marshall, K. Needham, T. Northcote, J. Richardson, P. Slaney and N. Winchester for help

in the production of the manuscript.

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J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 33

An overview of systematics studies concerning the insect

fauna of British Columbia

ROBERT A. CANNINGS

ROYAL BRITISH COLUMBIA MUSEUM,

675 BELLEVILLE STREET, VICTORIA, BC, CANADA V8W 9W2

GEOFFREY G.E. SCUDDER

DEPARTMENT OF ZOOLOGY, UNIVERSITY OF BRITISH COLUMBIA,

VANCOUVER, BC, CANADA VO6T 124

INTRODUCTION

This summary of insect systematics pertaining to British Columbia is not intended as

an historical account of entomologists and their work, but rather is an overview of the

more important studies and publications dealing with the taxonomy, identification,

distribution and faunistics of BC species. Some statistics on the known size of various taxa

are also given.

Many of the systematic references to the province’s insects cannot be presented in such

a short summary as this and, as a result, the treatment is highly selective. It deals largely

with publications appearing after 1950. We examine mainly terrestrial groups. Although

Geoff Scudder, Professor of Zoology at the University of British Columbia, at Westwick

Lake in the Cariboo, May 1970. Scudder is a driving force in many facets of insect

systematics in British Columbia and Canada. He is a world authority on the Hemiptera.

Photo: Rob Cannings.

34 J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

we mention the aquatic orders (those in which the larvae live in water but the adults are

aerial), they are more fully treated in the companion paper on aquatic insects in this issue

(Needham ef al.) as are the major aquatic families of otherwise terrestrial orders (e.g. the

Dytiscidae in Coleoptera, Corixidae in Hemiptera, Culicidae in Diptera, and so on).

However, when numbers of species are reported below for various orders or families,

aquatic species are included. The classification used here follows Kristensen (1991) except

that the so-called entognathous hexapods (e.g., Collembola) are treated as classes. A great

benefit of this scheme is that it is based on a cladogram and is supported by phylogenetic

discussion.

Danks (1979) summarized the insect fauna of Canada and indicated that there are about

54,000 insect species in the country, with almost half of these undescribed or unrecorded.

Cannings and Cannings (1996) estimated that there are about 35,000 species in BC, over

60 per cent of the Canadian fauna.

Spencer (1952) briefly reviewed the status of knowledge of the insect orders in the

province up to 1951. Since this, major advances have occurred in many, but not all,

groups. The systematic overview of many groups in the Yukon (Danks and Downes 1997)

is a useful basis for the study of the insect fauna of northern BC, even though collections

from the region may be scattered or lacking. In an annotated list of the potentially rare and

endangered species in BC, Scudder (1994) gave references to the available checklists,

major monographs and keys useful for the identification of our insect fauna. Nadel (1996),

Scudder (1996) and Biological Survey of Canada (1996) listed some systematic specialists

able to identify BC material.

SYSTEMATIC SURVEY

Class Protura

Based on regional distributions elsewhere, Marshall (1993) estimated that 25 species of

proturans should live in BC. However, only three species in two families have been

reported: Nipponentomon bifidum Rusek, N. kevani Rusek and Vesiculentomon marshalli

Rusek — all described from Douglas-fir forest near Shawnigan Lake (Rusek 1974). The

North American genera were keyed by Copeland and Imadaté (1990); the world fauna was

treated by Tuxen (1964).

Class Collembola

Spencer (1948a) published a preliminary list of the springtails known in BC. This was

augmented by Skidmore (1995) who, in a recent checklist of the Collembola of Canada

and Alaska, listed 145 species and 14 families from the province. This list, however,

represented species from only a small number of localities and habitats, and omitted some

recorded ones; for example, about 19 taxa listed by Battigelli and Marshall (1993) were

not included. In addition, one more family and at least 25 other species, as well as several

undescribed species, have since been collected in BC by J. Addison, H. Nadel and B.

Baumbrough (J.A. Addison, in /itt.; G. Marshall, in /itt.). Marshall (1993) estimated 200

species occur in the province and noted that there is a desperate need for basic taxonomic

and ecological studies in BC’s soil fauna. A detailed and annotated checklist of the

Collembola for the entire province is sorely needed; a good basis for such a work is the

treatment of the North American fauna by Christiansen and Bellinger (1998).

Class Diplura

Three species of Diplura in two families are recorded in BC (Spencer 1952; Marshall

1993), but more are likely to occur (V.G. Marshall, in /itt.). As well as the widely

distributed Campodea and the japygid, Evalljapyx sonoranus Silvestri, recorded from

Victoria, another japygid from the Gulf Islands and the Queen Charlotte Islands is

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 35

apparently undescribed (M.A. Muegge, in /itt.). Ferguson (1990a) provided a key to the

families of Diplura and genera of Campodeinae in the United States.

Class Insecta

Order Archeognatha

The two families of jumping bristletails, the Machilidae and Meinertellidae, both occur

in the province, but there is no current checklist. Mesomachilis canadensis Sturm

(Machilidae) and Nearctolinus auriantiacus (Schoett) (Meinertellidae) are rarely collected

species of grassland and dry forest communities in the Interior (Scudder 1994). A species

of Pedetontus is common above high tide on rocky seashores on the south coast. New

records and new species from BC have been published by Sturm (1991) and Sturm and

Bach de Roca (1992) but these were not included in the available key to the genera of the

contiguous United States (Ferguson 1990b).

Order Thysanura

Two alien species of bristletails, Lepisma saccharina Linnaeus (silverfish) and

Thermobia domestica (Packard) (firebrat) commonly occur indoors in BC (Scudder 1994).

Both are household pests. Ferguson (1990b) published a key for the identification of these

and other species.

Order Ephemeroptera

Ten families and 92 species of mayflies are recorded in BC. Needham ef al., in a

companion paper to this one, discuss systematic and ecological studies on the aquatic

insects of the province.

Order Odonata

There are 87 species of dragonflies and damselflies known in the province. These are

contained in ten families. Needham ef a/., in a companion paper to this one, discuss

systematic and ecological studies on the aquatic insects of the province.

Order Blattodea

Vickery and Scudder (1987) recorded 14 species of cockroaches in three families in

BC, but probably fewer than five of these are established. Blattella germanica (Linnaeus)

is the most common and has been reported throughout the southern part of the province

(Vickery and Kevan 1985). All our species have been introduced from elsewhere through

commerce (e.g. Belton et a/. 1986) or from laboratory cultures; none live freely outside

buildings except, perhaps, B. germanica, which is known to survive in refuse heaps.

Vickery and Kevan (1985) provided keys for their identification.

Order Mantodea

There are only two species of mantids in the province, both in the family Mantidae --

the rare, native Litaneutria minor (Scudder) and the alien, introduced Mantis religiosa

Linnaeus (R.A. Cannings 1987; Vickery and Scudder 1987). Both species are restricted to

the Okanagan Valley, although recent specimens of Mantis from southern Vancouver

Island suggest a population may be established there. Vickery and Kevan (1985) gave keys

for identification and R.A. Cannings (1987) documented the occurrence and ecology of

Litaneutria.

Order Isoptera

Three native and one introduced termite species live in BC. They are Reticulotermes

hesperus Banks (Rhinotermitidae), Zootermopsis angusticollis (Hagen) and Z. nevadensis

36 J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

(Hagen) (Termopsidae) and the alien Cryptotermes brevis (Walker) (Kalotermitidae)

(Vickery and Scudder 1987). Cryptotermes occurs only in indoor colonies here. Beall

(1931) and Spencer (1937a) wrote about the provincial species, but the most

comprehensive early work was never published — J.K. Jacob’s Masters thesis (Jacob 1938).

Vickery and Kevan (1985) provided identification keys.

Order Grylloblattodea

Buckell (1925) first collected Grylloblatta campodeiformis Walker in 1925 under logs

at 2286 metres elevation in the Selkirk Mountains near Invermere. The species was

captured a second time in BC beneath rocks on a talus slope on Mt. St. Pauls (Mt. Paul)

near Kamloops (Gregson 1938). This collection at only 427 metres elevation in dry forest

on 14 November 1936 surprised entomologists, because earlier reports associated this

unusual insect with high altitude habitats. Campbell (1949) detailed the circumstances of

its occurrence at Kamloops and in this paper Spencer speculated that the Grylloblatta at

Kamloops might be a separate race from the Rocky Mountain one.

Kamp (1973) made extensive collections of Grylloblatta throughout BC and the

western United States; he considered the Kamloops populations to be the nominate

subspecies, G. c. campodeiformis; this is also the opinion of V.R. Vickery (in Jitt.). Kamp

(1973) also extensively studied the habits, habitats, temperature preferences, and

comparative lipid composition of Grylloblatta and later described new species and

subspecies from BC, namely G. c. athapaska Kamp from Stone Mountain, G. c. nahanni

Kamp from the Cassiar Mountains and G. scudderi Kamp from Whistler Mountain in

Garibaldi Provincial Park (Kamp 1979). All of these are potentially rare in BC (Scudder

1994). Grylloblatta c. campodeiformis is widely distributed and recently was collected

commonly in both logged and unlogged terrain at high elevation forests in the Interior (D.

Huggard, in /itt.). Gregson (1996) gave a popular summary of Gry//oblatta in the province.

The species and subspecies were keyed by Vickery and Kevan (1985).

Order Dermaptera

Four alien species of earwigs in three families are reported from BC (Vickery and

Scudder 1987); they were keyed by Vickery and Kevan (1985). The species are:

Anisolabis maritima (Bonelli) and Euborellia annulipes (Lucas) (Anisolabididae), Labia

minor (Linnaeus) (Spongiphoridae) and Forficula auricularia Linnaeus (Forficulidae).

Anisolabis frequents ocean beaches on the southwest coast; Forficula is the common

earwig, an irritant to many gardeners.

Order Orthoptera

Early studies on the ecology and systematics of the Orthoptera in the province were

published by Buckell (1921, 1922, 1924, 1925), Handford (1961), and Treherne and

Buckell (1924a, 1924b). Spencer (1958a) outlined the natural control complex affecting

grasshoppers in Southern Interior grasslands. He also described the habits, larval stages

and economic importance of two nemestrinid flies that parasitize the grasshoppers in this

region (Spencer 1958b). More recently, Vickery and Nagy (1973) documented additional

ecological information on local species and Scudder and Kevan (1984) published an

updated list.

In the most recent annotated checklist of the Orthoptera in Canada, Vickery and

Scudder (1987) listed 117 species in 12 families from the province. Of the 40 species of

Ensifera (katydids, crickets) listed, 5 are adventitive (recorded but not established) and one

is an alien introduced species. Among the 77 Caelifera (grasshoppers), 2 are adventitive.

Eleven Orthoptera species in BC are possibly rare (Scudder 1994).

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 37

Vickery and Kevan (1985) published a monograph on the fauna, and Otte (1981, 1984)

provided invaluable additional information. Although the species are relatively well

studied, the identity of some is still in doubt. For example, the Jerusalem Cricket recorded

in BC as Stenopelmatus fuscus Haldeman is actually an undescribed species and the

identities of Gryllus species need clarification (D. Wiesmann, pers. comm.). In fact, the

correct identity of all material in collections in BC needs verification.

Order Plecoptera

Nine families of stoneflies containing 132 species are recorded in BC. Needham ef ai.,

in a companion paper to this one, discuss systematic and ecological studies on the aquatic

insects of the province.

Order Psocoptera

Although the bark lice have not been well studied in BC, 22 species in 13 families have

been recorded (Mockford 1993). This work supplies identification keys, but

determinations are difficult.

Order Phthiraptera

The bird lice (Amblycera) and mammal lice (Anoplura) were favourite groups of G.J.

Spencer, who published the original lists of our fauna. The Amblycera were examined by

Spencer (1928, 1948b, 1957) and Ballard and Ring (1979). The most modern treatment of

bird lice in Canada (Wheeler and Threlfall 1989) listed four families, 168 species and

subspecies and their known hosts in BC. Emerson (1972) is also a useful reference for bird

lice.

Spencer (1966) published an annotated list of the Anoplura of BC, and some of the

entries were corrected by Kim ef al. (1986), who also provided identification keys.

Twenty-six species in 8 families are known from the province. Both Spencer (1966) and

Kim et al. (1986) listed the known hosts.

Order Thysanoptera

Chiasson (1986) recorded 44 species of thrips in 3 families in BC. The fauna has not

been well studied.

Order Hemiptera

Following the many early records of Hemiptera from BC published before 1920 in the

‘Annual Report of the Entomological Society of Ontario’, Parshley (1919, 1921), Stoner

(1920, 1925), Downes (1924), and Torre-Bueno (1925) reported other species from the

province.

Downes (1927), the first true hemipterist to intensively survey the provincial bug

fauna, produced a complete checklist. Subsequently, many additions have been published,

including those by Downes (1935, 1957), Scudder (1960, 1961a, 1961b, 1985, 1986, 2000)

and Schwartz and Scudder (1998, 2000). Waddell (1952) made a list of the Hemiptera

from the Kootenay Valley, but this lacked precise locality records.

Published papers on the scale insects in the province include Venables (1939) and

Koczar et al. (1989), while Kitching (1971) listed and keyed the Psyllidae. Gillespie (1985)

published a paper on whiteflies. Following a series of 19 papers on aphids by A.R. Forbes

and C.K. Chan (with various co-workers) that appeared from 1973 to 1989, Forbes and

Chan (1989) published a list of aphids and host plants known from BC. Two more papers

(Forbes and Chan 1991; Chan and Frazer 1993) followed. The recent checklist of

Canadian and Alaskan species of Hemiptera (Maw ef al. 2000) listed 580 species of

Sternorrhyncha (aphids, psyllids, whiteflies and scales), 534 Clypeorrhyncha (cicadas,

38 J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

leafhoppers, froghoppers and treehoppers), 79 Archaeorrhyncha (planthoppers) and 815

Prosorrhyncha (true bugs), for a total of 2008 species in 72 families. Additional species

new to BC have been added since, and several new species are being described.

The Heteroptera (Prosorrhyncha) of the Montane Cordillera Ecozone were listed by

Scudder (1998), and the Clypeorrhyncha and Archaeorrhyncha by Hamilton (1998).

Several monographs on the Canadian fauna are available to aid the determination of BC

species. These include treatments of the minute pirate bugs (Anthocoridae) (Kelton 1978),

the flatbugs (Aradidae) (Matsuda 1977), the spittlebugs (Cercopidae, Clastopteridae)

(Hamilton 1982), the prairie plant bugs (Miridae) (Kelton 1980), stink bugs

(Pentatomidae) (McPherson 1982), and genera of aphids (Aphidoidea) (Foottit and

Richards 1993). References to other literature are given by Maw et al. (2000).

Order Neuroptera

Seven of the eight families of Neuroptera reported from BC are terrestrial — the

Berothidae, Chrysopidae, Coniopterygidae, Hemerobiidae, Mantispidae, Myrmeleontidae

and Polystoechotidae. The Sisyridae, whose larvae feed on sponges, are aquatic. Spencer

(1942) gave preliminary lists of all families and most have been investigated in detail since

then, resulting in a list of at least 66 terrestrial species.

Lomamyia_ occidentalis (Banks) is the only species of Berothidae known in the

province, recorded from Lytton by Spencer (1942). The green lacewings (Chrysopidae)

have been investigated in detail by Garland (1982, 1984, 1985, 2000, 2001); 18 species in

7 genera are now reported, some of which may be considered rare (Scudder 1994). The

brown lacewings (Hemerobiidae), studied by Klimaszewski and Kevan (1985, 1987, 1988,

1992), are represented by 33 species in 5 genera; some of these may be at risk (Scudder

1994). In his monographs on the Coniopterygidae, Meinander (1972, 1974) listed only four

species from the province, but the family has been little studied here.

Two species of Mantispidae occur, Climaciella brunnea (Say) and Mantispa pulchella

(Banks), the former from Vancouver Island to the Rockies, the latter only in the Okanagan

Valley. Mantispa pulchella, at least, may be at risk in the province because of its rarity and

limited distribution. In BC there are five species of antlions in three genera, but only four

species have been named with certainty; works useful in the identification of

Myrmeliontidae include Banks (1927) and Stange (1970). The family Polystoechotidae is

represented by a single rare species, Polystoechotes punctatus (Fabricius).

Order Raphidioptera

Spencer (1942) listed two species of snakeflies, but the recent world revision of the

Raphidioptera by Aspéck et a/. (1991) indicates that eight species in two families occur in

the province. Of these, Agu/la adnixa (Hagen) is the most common and widespread. Agulla

bicolor (Albarda), known only from the Osoyoos area, and A. crotchi Banks, collected

only from Summerland in BC, may be at risk.

Order Megaloptera

The Megaloptera is a small order in BC; this aquatic group includes the dobsonflies

(Corydalidae) and the alderflies (Sialidae). The former family contains three species, the

latter has five recorded in the province. Needham et a/., in a companion paper to this one,

discuss systematic and ecological studies on the aquatic insects of the province.

Order Coleoptera

The most recent checklist of the beetles of Canada and Alaska (Bousquet 1991) listed

3626 species in BC, which is about half the total number in Canada. One-hundred families

are represented; the ten most speciose are the Staphylinidae (581), Carabidae including the

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 39

tiger beetles (483), Curculionidae (261), Elateridae (194), Chrysomelidae (181),

Dytiscidae (167), Cerambycidae (145) and Scolytidae (134), Coccinellidae (94) and

Scarabaeidae (88). There are probably another 1200 or more species still unrecorded in the

province.

This list, of course, rests on many earlier works in the province. Spencer (1952) noted a

few of them, and Hatch (1952) added detail. Mentioned are the studies of Keen (1895) in

the Queen Charlottes, the long list by Auden (1925) from Midday Valley near Merritt, the

collections of Clark (1948, 1949) around Terrace and those of Hardy, especially in the

Cerambycidae and Buprestidae, on Vancouver Island (1942, 1944, 1950) and elsewhere

(1948). The large collections and studies of Stace-Smith (1929, 1930), R. Hopping (1922)

and G. Hopping (1932, 1937) were also critical. The most significant publication on the

province’s beetles remains Hatch’s monumental five-part treatise (Hatch 1953, 1957,

1962, 1965, 1971) keying and describing all the species in the Pacific Northwest

(including southern BC) known at that time. Arnett (1983) compiled North American

species, and most genera can be identified using Arnett (1968); Bousquet (1991)

highlighted generic revisions and good sources for species keys. Scudder (1994) listed 114

rare beetle species and subspecies in the province. Campbell (1979) summarized the

Canadian fauna. Anderson (1997) compiled the fauna of the Yukon; the biogeography of

many BC species, especially northern ones, is clarified by this work.

The Carabidae has been a favourite family of study in the province, and much of the

systematic and ecological work on the ground beetles has depended on the identification

power of the keys and descriptions in Lindroth (1961-1969). Wallis (1961) wrote a

monograph on the tiger beetles (Cicindelidae) of Canada and Freitag (1999) provided an

up-to-date taxonomy of the group, which is placed by many in the Carabidae (see

Bousquet 1991). A sampling of significant papers revising ground beetle groups that deal

with BC species include Ball (1966) (Pterostichus subgenus Cryobius), Bousquet (1988)

(Dyschirius), Erwin (1970) (Brachinus), Goulet (1983) (Elaphrus) and Maddison (1993)

(Bembidion subgenus Bracteon).

Kavanaugh examined the biogeography of the Carabidae, especially of the Queen

Charlotte Islands (1992) and, in the genus Nebdria, throughout northwestern North America

(Kavanaugh 1980, 1988). Spence and Spence (1988) studied the introduced ground beetles

of western Canada and the influence humans have had on their distribution. Some of the

surveys of carabids in the province are noted in the companion paper on collections,

surveys and conservation (Cannings ef a/.) in this issue.

Anderson and Peck (1985) treated the Silphidae and Agyrtidae of Canada and

Campbell (1968) revised the Micropeplidae. The extensive provincial diversity of the

Leiodidae (small scavenger beetles) is not well known, but some genera have been revised,

for example, Anisotoma (Wheeler 1979). Modern revisions of many genera of the huge

family Staphylinidae occurring in BC are available, for example, Bledius (Herman 1986),

Quedius (Smetana 1971) and Tachinus (Campbell 1973, 1988), although large gaps

remain. The largest Canadian subfamily, the Aleocharinae, is especially poorly known,

although some genera have been studied, such as Aleochara (Klimaszewski 1984).

Scudder (1994) listed several rare species; Pseudohaida rothi Hatch (Omaliinae) was

found for the first time in Canada during the canopy studies in the Carmanah Valley

(Campbell and Winchester 1993).

J. Cooper is currently revising the Scarabaeidae of Canada and Alaska (Bousquet

1991). The dung beetles of the Aphodiinae are common in the province; Gordon and

Cartwright (1988) reviewed the tribe Aegialiini. The Buprestidae of Canada and Alaska

was revised by Bright (1987); this work includes keys to, and descriptions of, all the 88

known species in the province. Everson (1978) recorded 23 species from southern

Vancouver Island. The Ptilodactylidae, with only three species in Canada, is represented in

40 J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

BC only by Ptilodactyla serricollis (Say); Cannings and Fisher (1987) recorded the species

for the first time in the province. The Elateridae are speciose in BC, but there is no overall

treatment. However, Lane (1952) summarized earlier work and produced a preliminary list

of 150 species. Becker (1956, 1979) treated several large genera, including Agriotes and

Athous. The introduction of two European species of Agriotes was reported by Vernon and

Pats (1997). The most common BC lampyrids, those in the genus Ellychnia, do not

produce light as adults, and few people have ever reported fireflies in the province.

However, there are two light-producing species recorded in the literature, Pyropyga

nigricans (Say) and Photuris pennsylvanica (DeGeer) (Bousquet 1991) and a study by

R.A. Cannings and B. McVickar has turned up one or two more.

BC has at least 94 species of Coccinellidae, more than any other Canadian province.

Some of these have been introduced for biological control, and the interaction of native

and alien species is of interest. The taxonomy of BC (and nearctic) species is rather well

known, owing to the monograph of Gordon (1985). Belicek (1976) examined the western

species and analysed the biogeographic relationships between those in Alberta and BC.

The pioneering work of Hardy in the study of BC’s Cerambycidae was mentioned

earlier. The family in the province is large; the 145 species represent 40 per cent of the

Canadian fauna. Linsley (1962a, 1962b, 1963, 1964) and Linsley and Chemsak (1972,

1976, 1985) treated the North American fauna. With 181 species in the province, the

Chrysomelidae is even larger. The Chrysomelinae were reviewed by Wilcox (1972).

Numerous genera common in BC have been revised, for example Chrysomela (Brown

1956), Cryptocephalus (White 1968) and Plateumaris (Askevold 1991).

Bright (1992) revised the Canadian curculionoid families, except Curculionidae and

Scolytidae. Anderson (1988a) documented the weevils of the Queen Charlotte Islands and

the Montane Cordillera Ecozone in the southern Interior (Anderson 1998). Some revisions

that include significant provincial taxa deal with the Rhynchaeninae (Anderson 1989), the

Cleonini (Anderson 1988b), and genera such as Dorytomus (O’Brien 1970) and Tychius

(Clark 1971). The Scolytidae have a high profile in BC forestry and are well known

taxonomically; Bright (1976) revised the Canadian species. Duncan (1987) provided an

identification guide to Dendroctonus in the province.

Order Strepsiptera

The Strepsiptera are endoparasites of Hemiptera and solitary Andrena bees, often

classified with the Coleoptera. Although there are a number of unpublished observations in

the province, only two species in the Stylopidae, Stylops advarians Pierce, and S. leechi

Bohart, are recorded by Bousquet (1991). Kenner (in /itt.) collected Stylops shannoni

(Pierce) parasitizing Andrena hippotes Robertson in a Richmond garden.

Order Hymenoptera

The excellent treatment of the Hymenoptera families edited by Goulet and Huber

(1993) keyed all the BC families of this huge order and gave many references to

systematic studies. Krombein ef a/. (1979) is the latest catalogue of the North American

fauna; it included taxonomic details and brief summaries of distribution and biology of the

species. Masner (1979) summarized the Canadian fauna. Nevertheless, no complete

checklist of species has ever been produced. The number of species, even described ones,

in the province has not been calculated, but our estimate of recorded and unrecorded

species is about 10,000 in around 70 families. The Hymenoptera is probably the largest

order in BC, and contains the largest number of unrecorded and undescribed species. The

diverse parasitic forms are especially inadequately known. Scudder (1994) listed 79 rare

species that may be of management concern.

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 4]

Spencer (1952) gave the most important early compilations of provincial species: the

lists of ants (Buckell 1927, 1932), bees (Buckell 1949, 1950, 1951), vespid wasps (Buckell

and Spencer 1950), sphecid wasps (Spencer and Wellington 1948) and ichneumonid wasps

(Guppy 1948).

The fauna of Symphyta (sawflies and relatives) in the Montane Cordillera Ecozone

(the southern half of the BC Interior) was nicely summarized by Goulet (1998). The 254

species recorded represent 69 of the 119 Canadian genera. Most of the species (95 per

cent) are native and about 17 per cent occur nowhere else in Canada. The alien fauna

arrived mainly through Pacific coastal ports and via the nursery trade. Sawfly systematics

is generally up-to-date for most BC genera. Goulet (1992) covered all the fauna at the

generic level and many other groups have recently been treated. Goulet (1986) studied the

Dolerini (Tenthredinidae) and Middlekauf (1984) examined the Orussidae. Smith

published monographs on several subfamilies of the Tenthredinidae, including the

Allantinae (Smith 1979) and revised genera such as Nematinus (Smith 1986) and Arge

(Smith 1989). Examples of other genera treated are Deda (Gibson 1980a), Fallocampa

(Wong 1977), Macrophya (Gibson 1980b) and Tenthredo (arcuata group) (Goulet 1996).

Basic taxonomic studies of the vast superfamilies Ichneumonoidea, Proctotrupoidea,

Chalicidoidea, Cynipoidea and others that relate to species in Canada and BC are meagre.

Examples of classificatory studies in the Parasitica include works on the Braconidae by

Marsh (1965), Mason (1978, 1981) and Quicke and Sharkey (1989) and on the

Ichneumonidae by Barron (1976) and Townes (1969-1971). Finlayson (1990) and

Gillespie and Finlayson (1983) studied the larvae of the Aphidiidae and Ichneumonidae,

respectively, and made significant contributions to the systematics of these groups.

Mackauer (1968), Mackauer and Campbell (1972) and Smith ef al. (1999) also examined

various aspects of the systematics of the Aphidiidae, important parasitoids of aphids, in

BC. Masner (1979) noted that the Proctotrupoidea in North America is, perhaps, the least

known of the superfamilies of parasitic Hymenoptera — about 90 per cent of the species are

undescribed or unstudied; he (Masner 1976) revised part of the Diapriidae in North

America. In the Platygastroidea, Masner (1980) keyed the genera of the Scelionidae of the

Holarctic. Yoshimoto (1984) outlined the classification and identification of the Canadian

families and subfamilies of chalcidoid wasps. Darling (1983) revised the nearctic species

of Euperilampus (Perilampidae) and keyed the New World genera of Chrysolampinae

(Pteromalidae) (Darling 1986). Heraty (1985) keyed the genera and revised the species of

Eucharitinae (Eucharitidae) in North America and Huber (1988) examined Gonatocerus in

the Mymaridae.

The hymenopterous parasitoids of various forest pests have been documented in BC:

for example, 13 species attacked the black-headed budworm on Vancouver Island (Gray

and Shepherd 1993) and 9 parasitized, or were hyperparasites in, the larch casebearer

(Andrews and Geistlinger 1969).

The aculeate Hymneoptera are diverse in the province. Omitting the bees, Finnamore

(1998) tallied 408 species in BC, about 45 per cent of the known Canadian fauna. He

calculated that although 243 species are recorded from the Montane Cordillera Ecozone in

BC (this includes all of the southern Interior), this is about 70 per cent of the true total.

About two-thirds of aculeate wasps found in the Montane Cordillera Ecozone prefer

grasslands or the dry, warm habitats found on lower south-facing slopes; 69 species (25

per cent) are restricted to the Okanagan Valley (Finnamore 1998). The superfamily

Chrysidoidea is represented in BC by four families; relevant systematic works include

Bohart and Kimsey (1982) (Chrysididae), Evans (1978) (Bethylidae), Olmi (1984)

(Dryinidae) and Olmi (1995) (Embolemidae).

The five BC families of the Vespoidea are striking and mostly familiar insects. The

Tiphiidae were treated by Allen (1965, 1968, 1971). There are few relevant works for the

42 J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

Mutillidae; only Finnamore (1998) and the early paper by Mickel (1928) dealt with the BC

species. Although there were a few preliminary lists of Formicidae from the province (e.g.,

Buckell 1932, Blacker 1992), the descriptions, keys, and distributional information

provided by Naumann et al. (1999) are the most detailed data available on the ants in BC.

The spider wasps of the family Pompilidae were, in part, dealt with by Townes (1957) and

the vespid wasps by Akre ef al. (1980), Miller (1961), Carpenter and Cumming (1985),

Cumming (1989) and others. Gerber (1990) documented the spread into BC, in the 1980s,

of the introduced yellowjacket wasp, Paravespula germanica (Fabricius) and Cannings

(1989a) recorded an Asian hornet, Vespa simillima xanthoptera Cameron, on Vancouver

Island.

The Apoidea of BC can be split into two general groups, the sphecid wasps and their

relatives, the bees. The Sphecidae are diverse, often spectacular and especially abundant in

the dry Interior; 174 of the 189 species recorded in BC are found in the Montane

Cordillera Ecozone (Finnamore 1998). Revisions relevant to BC species include those of

the tribes Sceliphronini and Sphecini (Bohart and Menke 1963) and the genera Crabro

(Bohart 1976), Cerceris (Ferguson 1984), Mimesa (Finnamore 1983) and Tachysphex

(Pulawski 1988). The genera of bees in North America were detailed and keyed by

Michener ef al. (1993). Some useful bee studies include those of the huge genus Andrena

(Andrenidae) by LaBerge (1986-1989), the Anthophorini (Brooks 1988), the genera of

New World Megachilini (Mitchell 1980) and Bombus (Milliron 1971). Although much bee

research is done in BC at M. Winston’s laboratory at Simon Fraser University, it is mostly

in fields other than systematics. The diversity of native bee species pollinating berry crops

in the Fraser Valley was examined; 13 species were recorded, most of them bumblebees

(Winston and Graf 1982).

Order Mecoptera

The genus Boreus in the Boreidae is the only known taxon of Mecoptera in BC. These

snow scorpionflies are small flightless insects that most commonly are found hopping on

the snow in winter. Penny (1977) published a monograph on the family; the five named

species in BC were included in the descriptions and identification keys. D. Blades (pers.

comm.), now studying the genus in the province, believes that there is at least one

undescribed species from the south coast. Boreus elegans was chosen as the emblem of the

Entomological Society of BC and the Society’s newsletter is named for the genus

(Cannings 1981).

Order Siphonaptera

Spencer (1936, 1937b) and Wagner (1936) published the main early papers on the fleas

in BC. Holland (1985), in his superb work on the group in Canada, Alaska and Greenland,

listed 6 families, 98 species and 6 additional subspecies in the province. There apparently

is some endemism; for example, Megarthroglossus sicamus Jordan and Rothschild is

restricted to Bushy-tailed Woodrats (Neoftoma cinerea (Ord)) in the Dry Interior. Fleas

transmit bubonic plague to mammals in the Interior. The bacterium was recorded in

Yellow-bellied Marmots (Marmota flaviventris (Audubon and Bachman)) in 1950

(Holland 1985) and in woodrats and some carnivores in 1988 (D. Nagorsen, pers. comm.)

Order Diptera

The Manual of Nearctic Diptera (McAlpine et al. 1981, 1987, McAlpine and Wood

1989) is the major single resource for information on systematics and biology of North

American (and BC) Diptera. Illustrated keys identify adult (and often immature)

specimens to family and genus, citations for generic revisions are given and phylogenetic

hypotheses for higher categories are outlined. Subsequently, significant advances in higher

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 43

classification directly relevant to BC studies were published by dipterists at the Canadian

National Collection of Insects, Arachnids and Nematodes in Ottawa (Wood 1991, Sinclair

et al. 1994, Cumming et al. 1995). The taxonomic and distributional status, to the early

1960s, of many BC species was outlined by Stone ef al. (1965). Stone (1980) also

summarized the history of North American dipterology, and included many major

publications and biographies of workers important in naming the province’s fly species.

The work of McAlpine (1979), who summarized the Canadian fauna, and Cole (1969) has

relevance to most fly families in BC, but no complete checklist of species has ever been

developed and it is unclear how many species are known for the province. Our estimate of

recorded and unrecorded species is about 8500 in almost 100 families. Scudder (1994)

listed 76 species that are possibly rare and threatened.

Spencer (1952) mentioned the pioneering work of Osburn (1908), Sherman (1920),

Garrett (1925) and Spencer (1943, 1948c). Spencer (1948c) listed the Tipulidae known at

that time. A most interesting genus in this huge family is Chionea, wingless crane flies that

walk about on the snow. These are common in the province and have been monographed

by Byers (1983); S.G. Cannings (1987) added C. macnabeana Alexander to the Canadian

list. Cramptonomyia spenceri Alexander, named after the famous University of BC

professor who discovered it, is the sole member of the Pachyneuridae in Canada; the larvae

live in dead red alder (A/nus rubra Bong.) logs. Its biology and distribution were recorded

by Vockeroth (1974) and Cannings and Cannings (1979).

The Bibionidae are by far the most common Diptera fossils in the abundant Eocene

shales of the province. Rice (1959) gave an overview of many of the species; 20 of 22 are

in the genus Plecia, which today is largely a tropical taxon. Cecidomyiidae and

Mycetophilidae are huge families in the forests of BC but remain largely unknown despite

their importance in plant and soil health. A little work has been done, however. For

example, in the Cecidomyiidae, Tonks (1974) found a species of Oligotrophus new to

Canada on junipers on Vancouver Island; Coher (2000) made some changes to the

taxonomy of the mycetophilids based on collections from Winchester’s surveys of forest

insects in the Carmanah Valley.

Curran (1927) and McFadden (1972) listed a number of the Stratiomyidae in the

province. Although many of its species develop in wetlands, we have not included the

Tabanidae in the aquatic insect chapter, but deal with the family here. The deer and horse

flies are of great importance because the females suck mammalian blood. Both Teskey

(1990) and Turner (1985) are useful for identifying BC species. Teskey (1985) also dealt

with some of the immature stages. Irwin and Lyneborg (1980) described and keyed the

nearctic genera of the Therevidae. The single member of the Apioceridae in Canada,

Apiocera barri Cazier, one of the rarest of the province’s flies from the sandy shrub-

steppes of the South Okanagan, was included in a revision of the genus by Cazier (1982).

The same locations support another rare fly, Nemomydas pantherinus (Gerstacker), the

sole species of the Mydidae in BC and one of only two in Canada. In the Asilidae,

revisions of large genera such as Cyrtopogon (Wilcox and Martin 1936), Efferia (Wilcox

1966), Lasiopogon (Cole and Wilcox 1938) and Dioctria and related genera (Adisoemarto

and Wood 1975) included references to species in BC. The various taxonomic works of

Curran, for example, the designation and summary of the genus Eucyrtopogon (Curran

1923) also are relevant. Foxlee's (1942) intensive collecting around Robson in the

Columbia Valley of the West Kootenay region resulted in specimens that are still the main

source of our knowledge for that region. Adisoemarto (1967), in his overview of the

Asilidae of Alberta, included records from the province. Cannings (1994) updated the

species list for the region and published an account of the species found in a grassland

typical of mesic sites at low elevations in the southern Okanagan Valley (Cannings

1989b). He has studied the taxonomy and biogeography of Rhadiurgus (Cannings 1993)

44 J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

and the large genus Lasiopogon (Cannings 2002). Cannings’ (1997) work on the fauna of

the Yukon gave a picture of the probable robber fly fauna of BC’s northern regions.

Cannings (1998) outlined the fauna of the Montane Cordillera Ecozone, covering much of

the Interior. There remains much work to be done on the provincial fauna of the two very

diverse families of the Empidoidea. Spencer (1943) made an early list of the

Dolichopodidae, and Pollet and Cumming (1998) revised Achalcus. Sinclair studied the

higher classification of the Empididae and reviewed Trichoclinocera (Sinclair 1994).

There is much more known about the large family Syrphidae in the province. Osburn

(1908) made an early list, added to by Allan (1969) and Morgan and Arrand (1971).

Vockeroth revised a number of genera including Paragus (1986) and Platycheirus (1990)

and published a monograph of the large subfamily Syrphinae (1992) — all vital for an

understanding of the province’s flower flies. Also in the Achiza, other genera have

received relevant and useful revisions by students elsewhere; examples include

Gymnophora (Phoridae) (Brown 1987) and Pipunculus (Pipunculidae) (Skevington and

Marshall 1998).

Few acalyptrate families have been treated from a provincial perspective, let alone a

Canadian one. Smith (1959) produced a preliminary list of the Conopidae of the province.

In the Sphaeroceridae, Marshall and others have revised a number of genera, for example,

Spelobius (Marshall 1985). The Canadian fauna of a handful of families has been analysed,

including the Agromyzidae (Spencer 1969), the Micropezidae (Merritt and Peterson 1976)

and the Piophilidae (McAlpine 1977). Likewise, in the calyptrate Cyclorrhapha, reference

to the BC fauna is found in a number of works such as Hall (1948) for the Calliphoridae

and Wood (1985) for the blondeliinine tachinids.

Order Trichoptera

There are 279 species of caddisflies in 15 families recorded in BC. Needham ef al., ina

companion paper to this one, discuss systematic and ecological studies on the aquatic

insects of the province.

Order Lepidoptera

The authoritative list of the Lepidoptera of North America that has been the basis of

systematic study in BC for almost 20 years is Hodges et al. (1983). Munroe (1979)

summarized the Canadian fauna. The recently published ‘Butterflies of British Columbia’

(Guppy and Shepard 2001) is the major single resource on the butterflies of the province.

It thoroughly covers the description, distribution, taxonomy and status of the 187 species

recorded in BC and provides an exhaustive bibliography of publications dealing with the

systematics and biology of the fauna. The authors described eleven new subspecies and

estimated that nine more peripheral species will be added to the provincial list. There are

several older publications dealing with the butterflies on a broader geographical scale; the

most useful is the recent ‘The Butterflies of Canada’ (Layberry ef al. 1998). There is no

modern treatment of the moths of the region, although the ongoing and detailed ‘Moths of

North America north of Mexico’ (selected monographs are cited below under specific

families) covers some important groups. Studies on the moth families of the province at

the species level are sorely needed. Approximately 4000 species of Lepidoptera in about

60 families occur in BC. Scudder (1994) listed 61 species that are possibly rare and

threatened. Guppy ef al. (1994) and Guppy and Shepard (2001) gave details of the

butterflies and skippers that are of conservation concern.

In Guppy and Shepard (2001), Shepard wrote an excellent account of the early history

of Lepidoptera collections and systematics in BC, a history that goes back to 1850. The

discussion focuses on butterflies and skippers, but it is a good source of historical papers

on all BC Lepidoptera. The monumental three volume work by Edwards (1868-72, 1874-

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 45

84, 1887-97) was the earliest North American butterfly work that included some of the BC

fauna; it is still of great taxonomic significance. Anderson (1904) published the first

synthesis of BC material, followed by several provincial and regional lists culminating in

Blackmore (1927). This was followed 24 years later by Llewellyn Jones’ (1951)

‘Annotated Check List of the Macrolepidoptera of British Columbia’, which not only

summarized the known distribution of each species, but included flight periods and larval

food plants.

G.A. Hardy documented the Lepidoptera fauna of southern Vancouver Island in the

1950s and 1960s, and published a number of studies of the larval stages and life histories

of both butterflies and moths in the ‘Proceedings of the Entomological Society of British

Columbia’ (e.g., Hardy 1957, 1963). Also in the ‘Proceedings’ and the ‘Journal’ that

succeeded it, a long series of annotated lists documented the forest insects of the province,

and Lepidoptera played a large role (Ross and Evans 1957; Sugden 1968). Guppy (1956)

studied the macrolepidoptera of Vancouver Island. Underhill surveyed the fauna of

Manning Provincial Park (Harcombe and Underhill 1970) and Threatful (1989) studied the

butterflies of Mount Revelstoke and Glacier National Parks. Kondla et al. (1994)

documented the butterflies of the Peace River district and Kondla (1999) reported on the

species he collected in the Pend d’Oreille Valley. Fischer et al. (2000) recorded the

macrolepidoptera of the Chilcotin. Significant range extensions, including additions of

butterfly and moth species to the province’s fauna, have been recorded in the annual ‘Field

Season Summary’ of the Lepidopterists’ Society for over 50 years. Other recent inventory

efforts are discussed in the paper by Cannings ef a/. on insect collections and surveys in

this issue. The affinities of Yukon Lepidoptera were examined by Lafontaine and Wood

(1997), throwing light on the biogeography of many BC forms. The introduction of alien

Lepidoptera is documented in publications such as Gillespie and Gillespie (1982), which

recorded 48 plant feeding species and their history in the province.

Numerous taxonomic and biogeographic studies have been published by BC butterfly

specialists. Shepard published on the taxonomy of Bo/oria (Shepard 1975), Parnassius

(Shepard and Manley 1998) and other taxa. Guppy (1986) studied geographic variation in

the wing melanism of the butterfly Parnassius phoebus (Fabricius) and Troubridge and

Parshall (1988) reviewed the Oeneis polixenes (Fabricius) complex. All this butterfly work

is encapsulated in Guppy and Shepard (2001).

The moths of BC need much work, but many treatments of taxa of various sizes are

scattered through the literature; those listed here are only a small sample. In the huge

Gelechioidea, with many BC species, Landry (1991) has studied the North American

Scythrididae and Hodges (1974) the Oecophoridae. The typical leafrollers of the

Tortricoidea are represented in the province by the large, economically important

Tortricidae; much of the work of interest deals with the population dynamics of spruce

budworm taxa and the fight against codling moth, best discussed in other papers in this

volume. The Pyraloidea is dominated by the family Pyralidae; much of the North

American fauna was revised by Munroe (1972-1976). The coneworms of the genus

Dioryctria, important in BC forests, were reviewed by Mutuura ef a/. (1969), Mutuura and

Munroe (1972-1973) and Sopow ef al. (1996).

In BC the Geometridae makes up most of the superfamily Geometroidea, and the

family is abundant in provincial collections. Many taxa were reviewed by McGuffin (e.g.,

McGuffin 1988) and he also made important contributions to the description of larvae (e.g.

McGuffin 1958). Bolte (1990) revised the genus Eupethecia. In the Bombycoidea, the tent

caterpillars of the Lasiocampidae have received much attention, and were studied by

Franclemont (1973). The Saturniidae includes some of our largest insects, the giant

silkmoths, which are noticed by everyone who comes across them. The BC species were

included in the revision by Ferguson (1971-1972) and in Tuskes ef al. (1996). Cannings

46 J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

and Guppy (1989) recorded Hyalophora gloveri Strecker for the first time in BC and

Morewood (2000) studied the colour pattern dimorphism in the more common H. euryalis

(Boisduval). The hawkmoths of the Sphingidae in the superfamily Sphingoidea also attract

the attention of the general public. The BC fauna is small (16 species) but spectacular.

They are described and illustrated in Borkent and Greenway (1997). The nearctic species

are treated by Hodges (1971).

The superfamily Noctuoidea is probably the largest in the order; its largest family, the

Noctuidae, alone has about 2000 species in Canada (Munroe 1979). Hardwick (1970)

studied the genera of the nearctic Heliothidinae and Lafontaine (e.g., 1987, 1998) revised

parts of the family. He also discussed in detail the biogeographic history of Euxoa in

western North America (Lafontaine 1982). The northern and Asian affinities of Beringian

noctuids were examined by Lafontaine and Wood (1988); this puts the distributions of

many BC species into perspective. Troubridge and co-workers have recently described a

number of noctuid species from BC, including several Oncocnemis (Troubridge and Crabo

1998).

ACKNOWLEDGEMENTS

We thank J. Addison, D. Blades, S. Cannings, C. Guppy and V. Marshall for help in the

preparation of this paper.

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J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 6]

Aquatic insects in British Columbia: 100 years of study

GEOFFREY G.E. SCUDDER, KAREN M. NEEDHAM,

REX D. KENNER

SPENCER ENTOMOLOGICAL MUSEUM, DEPARTMENT OF ZOOLOGY,

UNIVERSITY OF BRITISH COLUMBIA, VANCOUVER, BC, CANADA V6T 124

ROBERT A. CANNINGS

ROYAL BRITISH COLUMBIA MUSEUM,

675 BELLEVILLE STREET, VICTORIA, BC, CANADA V8W 9W2

and SYDNEY G. CANNINGS

BRITISH COLUMBIA CONSERVATION DATA CENTRE,

MINISTRY OF SUSTAINABLE RESOURCE MANAGEMENT,

PO BOX 9344 STN PROV GOVT, VICTORIA, BC, CANADA V8W 9M1

INTRODUCTION

Aquatic habitats in British Columbia are diverse and numerous. BC is home to large

rivers, small ponds, and lakes which vary tremendously in size, depth, and chemical

composition (Northcote and Larkin 1963), as well as a variety of fens, marshes, and bogs

(Rosenberg and Danks 1987). Not surprisingly, studies in aquatic insects have a long

history in our Province. The status of entomological knowledge from 1901 to 1951 in BC

was summarized by Spencer (1952), including short paragraphs on each of the aquatic

orders. The history of many of the entomologists involved and their collections are

highlighted in Hatch’s (1949) charming summary of a century of entomology (1835-1948)

in the Pacific Northwest.

In the past, provincial studies have emphasized the importance of freshwater fisheries,

and these have been accompanied by benthic invertebrate surveys. In recent years,

forestry-fisheries interactions have come to the fore. The role of insects in this interaction

is creating a new interest in aquatic entomology. The current concerns over water quality

and groundwater pollution will undoubtedly involve more study of aquatic insects as

environmental indicators.

EPHEMEROPTERA

Since the outstanding and prolific work of J. McDunnough in the 1920s and 1930s on

the mayflies of Canada, the only BC-wide survey of Ephemeroptera was conducted by

Filmer (1964). Most records since then have been obtained incidentally during other faunal

studies. A checklist of BC species was produced by Scudder (1976). McCafferty and

Randolph (1998) published a Canadian mayfly compendium that added seven new species

to Scudder’s 1976 list, as well as containing two species that Scudder (1976) omitted as

probable erroneous records, for a total of 92 species in BC. Since this new list was

compiled without examining specimens from the Royal British Columbia Museum, the

Spencer Entomological Museum, or the Fisheries Branch of the Ministry of Water, Land

and Air Protection, these collections may contain more new records for the Province.

Locally, Wigle and Thommasen (1990) collected and recorded ecological notes on 26

species of mayflies in the Bella Coola and Owikeno Lake watersheds of mid-coastal BC.

62 J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

One of these, Epeorus nitidus Eaton, was a new record for the Province. Another new

record for BC, Acentrella turbida (McDunnough), was found there later (McCafferty et al.

1994). Zloty (1996) and Zloty and Pritchard (1997) collected at various southwestern BC

locales in search of Ameletus species, and to date have found one, Ameletus pritchardi

Zloty, new to the Province and to science. S. Salter (pers. comm.) recently collected a new

mayfly species for the Province, Caenis youngi Roemhild, during some invertebrate

surveys at Liard Hot Springs in northern BC. In their comparison of the macroinvertebrate

assemblages of coastal and continental streams and large rivers in southwestern British

Columbia, Reece and Richardson (2000) found that Paraleptophlebia temporalis

McDunnough predominated in coastal streams.

Heise (no date) in a study of the effect of logging on the fauna of streams in the

Penticton Creek and Sicamous Creek watersheds found that mayflies were very sensitive

to logging. Richardson and Kiffney (2000) studied the response to metal pollutants of

macroinvertebrate fauna in experimental streams off Mayfly Creek in the UBC Research

Forest near Maple Ridge, and found that Ame/etus, Baetis, and Paraleptophlebia (mainly

P. temporalis) were very sensitive to Cu, Zn, Mn, and Pb, typical components of urban

runoff. Rempel et a/. (2000) in a study of the macroinvertebrates along gradients of

hydraulic and sedimentary conditions in a 10-km reach of the Fraser River near Agassiz,

found that the distribution of several genera of mayflies was correlated with hydraulic

conditions. Rempel e¢ a/. (1999) also showed that the densities of Baetis and Rhithrogena

were highest at a depth of 1.5 m before flooding, but shifted to depths of 0.2-0.5 m at peak

flow.

Keys to genera of BC mayflies can be found in Edmunds et al. (1976), Merritt and

Cummins (1996), and Needham (1996). Keys to species have only been completed for a

few genera (e.g. Ameletus, Zloty (1996), Zloty and Pritchard (1997); Caenis, Provonsha

(1990); Tricorythodes, Alba-Tercedor and Flannagan (1995)).

ODONATA

Spencer (1952) in his brief summary of the status of the knowledge of Odonata in

British Columbia did not cite what is perhaps the most significant work of the period,

Whitehouse’s (1941) delightful and detailed treatment of the provincial fauna. He also

neglected Walker’s important monographs on two of our most speciose genera, Aeshna

(Walker 1912) and Somatochlora (Walker 1925); although these dealt with the entire

North American fauna of these genera, they included much information on the species in

British Columbia.

Walker’s major opus on the Odonata of Canada and Alaska (Walker 1953, 1958) led

off the second half of the century and was completed by the international Odonata expert

P. Corbet (Walker and Corbet 1975). It listed many of the locality records known to this

time and summarized ecological information.

Since the mid 1970s the study of the Odonata in BC has been continuous. It has been

helped by work in adjacent areas, particularly the Biological Survey of Canada’s Yukon

insect project (S.G. Cannings and R.A. Cannings 1997; S.G. Cannings et al. 1991) and the

extensive studies of Paulson (e.g. 1997, 1999) to the south in Washington State. This

followed the annotated checklist produced by Scudder ef al. (1976) and the book written

by R.A. Cannings and Stuart (1977). The latter published the first distribution maps for the

species of the Province.

Robert and Sydney Cannings, periodically helped by their brother Richard, have been

the driving force in odonatology in the Province during the past 25 years. They organized

numerous dragonfly inventories across the Province, from the Brooks Peninsula on the

outer coast of Vancouver Island (R.A. Cannings and S.G. Cannings 1983), to the

grasslands of the Chilcotin (R.A. Cannings and S.G. Cannings 1987), to the Rocky

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 63

F.C. Whitehouse, a pioneering odonatologist, collecting specimens from his caravan at

Harrison and Cultus Lakes in the summer of 1936.

Mountains (R.A. Cannings eft a/. 2000) and the North (R.A. Cannings ef a/. 2001). Others

who have made significant contributions to these collections and inventories include G.

Hutchings, L. Ramsay, C. Guppy, R. Kenner, D. Blades, and H. Nadel. In addition, the

Cannings, along with their colleagues, have documented new species to the Province (R.A.

Cannings 1988, 1997; Kenner 2000a) and published much faunistic (R.A. Cannings 1978,

1996; R.A. Cannings ef a/. 1980; S.G. Cannings 1980; S.G. Cannings and R.A. Cannings

1994) and ecological work (R.A. Cannings 1980, 1982a; R.A. Cannings ef al. 1980;

Paulson and R.A. Cannings 1980). Significant in the taxonomic work done on the Odonata

in British Columbia and Yukon over the past 25 years is the description of four previously

unknown larvae (R.A. Cannings 1981; S.G. Cannings and R.A. Cannings 1980; R.A.

Cannings and Doerksen 1979; Kenner e¢ a/. 2000) and the redescription of several others,

along with improved keys to this important life stage (R.A. Cannings 1982b; Kenner

2001).

G.P. Doerksen had begun to examine local dragonflies intensively (Doerksen 1980)

when he was killed by a Grizzly Bear while on a research trip to Liard River Hot Springs

in 1981. His wonderful dragonfly photographs, willed to the Royal BC Museum, are a

valuable research and interpretative resource. Peters (1998), while on a visit from

Germany, published a useful study on variability in Aeshna. R. Kenner (Kenner 1996,

2000b, 2000c; Kenner and Lane 1997; Kenner and R.A. Cannings 2001) and I. Lane (Lane

2000) have improved our knowledge of the fauna of the Lower Mainland, including the

documentation of the first Canadian breeding site of Tanypteryx hageni Selys.

Few experimental studies on the Odonata have been undertaken in the Province. G.

Pritchard, University of Calgary, has studied the ecology and development of Odonata,

especially Argia vivida Hagen, of warm springs in British Columbia and adjacent regions

(Pritchard 1989, 1991). At the University of BC, P. Pearlstone (1973) examined the food

64 J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

of larval Enallagma boreale Selys and R. Baker (1983) studied the larval competition and

feeding behaviour of /schnura cervula Selys.

An important recent development has been the listing of species of management concern

by the BC Conservation Data Centre (Ramsay and S.G. Cannings 2000; R.A. Cannings

2001). At the end of the 2001 field season, 23 species were listed. This highlighting of rare

species spurred research into their status and habitats, and stimulated regional inventories.

Odonata are now the best known aquatics insects in the Province. Of course, there is

still much to learn, especially regarding the ecology of the fauna and the distribution of

northern species. Spencer noted 78 species in 1952, but the most accurate published record

up to that time was from Whitehouse (1941), who recognized 74 species in 23 genera. By

1977, R.A. Cannings and Stuart could report 80 species and 23 genera. In 2001, 87 species

in 27 genera are known from the Province (there are 201 recorded in Canada). Perhaps the

most striking addition to the provincial list in the past 50 years was the discovery of

Calopteryx aequabilis Say at Christina Creek in 1998 (R.A. Cannings et al. 2000). This

added a new family to British Columbia, the Calopterygidae, making a total of 10.

All available records for the Odonata in BC are databased, georeferenced, and mapped,

ready to serve as baseline data for the faunal changes of the new millennium. Useful works

for identification, in addition to Walker (1953, 1958), Walker and Corbet (1975), and R.A.

Cannings and Stuart (1977) are Westfall and May (1996), Needham ef al. (2000), and

Dunkle (2000).

PLECOPTERA

Ricker (e.g. 1943, 1954) conducted early systematic and distributional studies on

stoneflies. An annotated checklist of the Plecoptera of BC was published by Ricker and

Scudder (1976). S.G. Cannings (1989) also provided a number of new records, especially

for capniids. An up-to-date checklist can be extracted from the North American stonefly

list provided on the internet by Stark (2001). Currently, 132 species are known from the

Province. More detailed systematic studies that are relevant include Nelson and

Baumann’s (1989) work on Capnia, Stanger and Baumann’s (1993) work on Taenionema,

and Stark and Nelson’s (1994) work on Yoraperla. Kenneth Stewart (University of North

Texas) and Mark Oswood are actively writing "Stoneflies of Alaska and Northwestern

Canada," which will include history, keys, illustrations, new locality records, maps, species

accounts, and biological notes on the species documented for Alaska, British Columbia,

Yukon, and Northwest Territories; publication is anticipated in 2003.

D.B. Donald and R.S. Anderson of the Canadian Wildlife Service (Edmonton)

surveyed the stoneflies of the Rocky Mountain National Parks. The lentic stoneflies of

these collections were summarized in Donald and Anderson (1980). Donald and Patriquin

(1983) related the wing length of Rocky Mountain lentic capniids to their inferred

postglacial recolonization history.

A number of studies investigating stonefly population dynamics, foraging ecology,

species interactions, and life history in coastal streams have been undertaken by J.

Richardson and his colleagues at UBC (Reece and Richardson 2000; Rempel ef a/. 2000;

Richardson 2002; Soluk and Richardson 1997). Muchow and Richardson (2000) reported

the occurrence of Plecoptera in small headwater streams in the UBC Research Forest near

Maple Ridge. They found the fauna differed in intermittent and continuous flow streams.

While Paraleuctra vershina Gautin and Ricker and Visoka cataractae (Neave) occurred

only in continuous flow streams, Alloperla pilosa Needham and Claassen, Despaxia

augusta (Banks), Moselia infuscata (Claassen), Ostracerca foesteri (Ricker), Pteronarcys

californica Newport, Soyedina producta (Claassen), Zapada cinctipes (Banks), and Z.

oregonensis (Claassen) occurred in both. Even in small streams (less than half a metre

wide) with intermittent flow, the univoltine Moselia infuscata, Soyedina producta, and

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 65

Zapada cinctipes were found to emerge in periods when no flow was perceptible.

Despaxia augusta, with a 2-year life cycle, was also able to complete its development in

the periodically ‘dry’ channels and reached its highest densities in intermittent streams.

The authors suggest that suitable refugia exist for this species and others in the wet

sediments of these habitats despite the periodic disappearance of detectable surface flows.

All available, reliably identified records are now databased, georeferenced, and

mapped. References useful for identification include Baumann et al. (1977), Harper and

Stewart (1984), Jewett (1959), Ricker (1943), Stark et a/. (1986), and Stewart and Stark

(1988).

HEMIPTERA

In addition to records in the early literature of Hemiptera in British Columbia, Scudder

(1977) published an annotated checklist of the aquatic and semi-aquatic bugs in the

Province. In the recent checklist for Canada (Maw ef al. 2000), a total of 46 species of

aquatics (Infraorder Nepomorpha: Families Belostomatidae, Nepidae, Gelastocoridae,

Corixidae, Notonectidae) and 14 species of semi-aquatics (Infraorder Gerromorpha:

Families Mesoveliidae, Hebridae, Hydrometridae, Gerridae, Veliidae) are recorded.

Over the past 40 years, detailed studies of the biology of both Corixidae and Gerridae

have been undertaken, much of this associated with a study of the saline lakes of the

British Columbia interior. Scudder (1965a, 1969a, 1969b) investigated the distribution of

two closely related species of Cenocorixa in these lakes, and found that while C. bifida

(Hung.) was evidently a freshwater species, C. exp/eta (Uhler) was more saline tolerant.

Extensive study of the osmotic and ionic balance in these two corixid species (Scudder

1971a; Scudder et al. 1972; Szibbo and Scudder 1979; Needham 1990) and an

investigation of their cuticular permeability (Oloffs and Scudder 1966; S.G. Cannings

1981; S.G. Cannings et al. 1988) was accompanied by detailed histological and

ultrastructural studies of their excretory organs (Jarial and Scudder 1970) and other organs

that were evidently involved in osmotic and ionic regulation (Jarial et a/.1969; Lo and

Acton 1969; Jarial and Scudder 1971). Although the two species differed in their osmotic

and ionic regulatory abilities at high salinities, both were able to regulate equally well in

low salinity waters. Indeed, the more saline tolerant C. exp/eta, which in the field does not

live in freshwater lakes, was successfully reared in freshwater (S.G. Cannings 1978).

These studies subsequently led to much more detailed research on the coastal C. blaisdelli

(Hung.) (Cooper et al. 1987, 1988, 1989) and other corixid species (Scudder 1987).

Both of the interior species of Cenocorixa, and other Corixidae, were found to be

predaceous (Jansson and Scudder 1972; Reynolds 1975), which permitted laboratory

rearing and led to other studies. Once species had been reared and the immature stages of

both C. bifida and C. expleta described (Scudder 1966a), studies were carried out on their

life cycle (Jansson and Scudder 1974), flight muscle development (Scudder 1971b), and

flight muscle polymorphism (Scudder 1964; Acton and Scudder 1969; Scudder and

Meredith 1972; Scudder 1975).

Jansson (1972, 1973, 1974a, 1974b), in his extensive study of stridulation in these

Cenocorixa species, found species-specific mating signals and hence an effective pre-

mating isolating mechanism. Further, in an intensive study of their feeding biology, using

serological and other techniques, Reynolds and Scudder (1987a, 1987b) studied both the

fundamental and realized feeding niches of C. bifida and C. expleta.

In a review of the factors governing the distribution of C. bifida and C. expleta, Scudder

(1983), following studies on mite parasitism of these species (Smith 1977), suggested that

C. expleta was excluded from freshwater lakes by mite parasitism. This suggestion has

been substantiated by more research (Bennett and Scudder 1998), which showed that for

66 J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

C. expleta saline lakes are enemy-free space. This research has been cited as an example of

a parasite-structured animal community (Minchella and Scott 1991).

In the Gerridae, several species were found to coexist in BC lakes (Maynard 1969;

Scudder 1971c). Following the rearing and description of the immature stages of these

(Scudder and Jamieson 1972; Spence and Scudder 1978), detailed studies were undertaken

of their food consumption and predatory behaviour (Jamieson and Scudder 1977, 1979),

growth patterns (Spence ef al. 1980a), life cycles (Spence and Scudder 1980; Rowe and

Scudder 1990), mating and other behaviours (Spence et al. 1980b; Rowe 1992, 1994).

Spence (1981, 1983, 1989) was then able to undertake a detailed analysis of habitat

selection, life cycle strategy, species-packing, and coexistence in these pond skaters.

Subsequently, Spence (1990) discovered introgressive hybridization in two species of

Limnoporus within British Columbia and Alberta, and has gone on to look at the mating

system of these (Spence and Wilcox 1986; Wilcox and Spence 1986), and some of the

genetic factors accompanying this hybridization between non-sister species (Spence and

Maddison 1986; Sperling and Spence 1990, 1991; Sperling et a/. 1997).

Seven species of Notonectidae occur in BC (Scudder 1965b, 1977). Notonecta borealis

Hussey was discovered to be primarily a flightless species (Scudder 1966b). Ellis and

Borden (1969) have investigated the effect of temperature and other environmental factors

on N. undulata Say.

MEGALOPTERA

Spencer (1942) treated the Megaloptera, which includes the dobsonflies and the

alderflies, within his listing of the Neuroptera, and recorded 4 species from British

Columbia. A systematic review of the Sialidae by Ross (1937) gives a key to the species of

Sialis; 5 species are now know to occur in the Province. Munroe (1951, 1953) sorted out

the identity of material from British Columbia that Spencer (1942) reported as Neohermes

disjunctus, and now 3 species of Corydalidae are known from BC.

While Dysmicohermes disjunctus (Walker) is widely distributed, both Chauliodes

pecticornis L. and Protochauliodes spenceri Munroe may be at risk, as they have a

restricted distribution in areas of the Province subject to heavy human impact. Chauliodes

pecticornis is known from Cloverdale and Cowichan, while Protochauliodes spenceri is

restricted to south-east Vancouver Island in Canada. All known records of Megaloptera in

the Province are databased, georeferenced, and mapped.

NEUROPTERA

Of the 8 families of Neuroptera reported from British Columbia, only the family

Sisyridae has aquatic larvae. In the Sisyridae, although Spencer (1942) recorded only

Sisyra vicaria (Walker) from BC, based upon a specimen from Agassiz determined by

F.M. Carpenter, it is now known that another species of spongilla fly, Sisyra fuscata

(Fab.), also occurs in the Province. While S. vicaria is widely distributed, S. fuscata is only

known from Kaslo and Seton Lake. A key to the species of Sisyridae is given in Parfin and

Gurney (1956). All known records of aquatic Neuroptera in BC are now databased,

georeferenced, and mapped.

TRICHOPTERA

Some of the earliest records of Trichoptera from British Columbia come from N.

Banks, who worked on the group from the early 1900s to the early 1940s. H.H. Ross also

contributed significantly to these early records, collecting in the 1930s, 1940s, and 1950s

throughout North America. The first attempt to pull all of these scattered records together

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 67

into one publication was made by Ross and Spencer (1952), who created a preliminary list

of the Trichoptera of BC. This list was added to significantly by Nimmo and Scudder

(1978) when they published their annotated checklist containing 248 species, 43 of which

were new records for BC. At this time, they stated that “there has been no concerted

Trichoptera collecting in British Columbia as a whole, so the fauna must still be regarded

as incompletely known.”

To partially rectify this, A. Nimmo spent the spring, mid-summer, and fall of 1979

exploring much of the Province in search of caddisflies (Nimmo and Scudder 1983). He

added many range extensions to known species, and also recorded 36 new species for BC

(Nimmo and Scudder 1983), thus bringing the provincial total to 279 species (they

removed 5 species from the 1978 list). However, Nimmo admitted that the northern half of

the Province, the Queen Charlotte Islands, and many alpine areas still remain uncollected

(Nimmo and Scudder 1983).

Also in the 1970s, G. Wiggins published his invaluable and beautifully illustrated

identification guide to the larval caddisflies of North America (Wiggins 1977). A second

edition in 1996 recognized some 1400 species of Trichoptera in North America by the end

of 1993 (Wiggins 1996). More specific to our region are the keys provided by Schmid

(1980) to the genera of Trichoptera in Canada and adjacent States. An English version of

this Insects and Arachnids of Canada (Part 7) publication has also been recently made

available (Schmid 1998). Unfortunately, keys to caddisflies of BC at the species level are

still lacking.

Collections of caddisflies from specific areas of our Province can be found in Clemens

et al. (1939) for Okanagan Lake, Hardy (1955) for the Forbidden Plateau area of

Vancouver Island, Nimmo (1971, 1974, 1977) for eastern BC, Rawson (1934) for the

Kamloops region, Schmid and Guppy (1952) for southern Vancouver Island, Scudder

(1969b) for the Fraser Plateau, and Winterbourn (1971a,b) for Marion Lake, BC.

On the biology of caddisflies, Richardson (1991) found Lepidostoma roafi (Milne) to

be univoltine. In a study contrasting the fauna of coastal and continental streams and large

rivers in southwestern BC, Reece and Richardson (2000) found that Onocosmoecus

unicolor (Banks) was unique to coastal streams, while Brachycentrus occidentalis Banks

was unique to larger rivers. B. americanus (Banks) was unique to continental streams in

the Merritt area. In Marion Lake, Trichoptera larvae, in particular Banksiola crotchi

Banks, were the most important food item of rainbow trout (Efford and Tsumura 1973).

The effects of the mosquito larvicide Bacillus thuringiensis on caddisflies was reported by

Duckitt (1986).

Rempel et al. (1999), in an investigation of the effect of flooding on benthic

invertebrates in the Fraser River near Agassiz, noted that Hydropsyche was most abundant

at 1.5 m in all months of the years, but the location shifted laterally over a distance of 30 m

through the flood cycle. They evidently migrated to the shore zone with the rise in water

level, so maintaining a suitable hydraulic microhabitat. Heise (no date), in a study of the

effect of logging on the fauna of streams in Penticton Creek and Sicamous Creek

watersheds, found that while some caddisfly larvae showed a constant pattern of

distribution in response to logging, Psychoglypha, a shredder, declined after logging and

Ecclisocosmoecus, an algal feeder, increased after logging.

Caira and Scudder (1985) reported parasitism of Psychoglypha alascensis Banks

(Trichoptera: Limnephilidae) by Pseudoallocreadium alloneotenicum (Wootton) (Digenea:

Allocreadiidae). Caira (1981) investigated parasitism of Trichoptera by Bunodera

mediovitellata Zimbaluk & Roytman (Digenea: Allocreadiidae) and the encapsulation

response. This hemocyte encapsulation response in P. alascensis was studied later using

Epon implants (Caira and Scudder 1987).

68 J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

COLEOPTERA

British Columbia has a very rich aquatic beetle fauna. For example, BC has more

species of Dytiscidae, Amphizoidae, Haliplidae, and Hydraenidae than any other Province

(Larson ef al. 2000; Bousquet 1991). Yet, beetles appear to be one of the more neglected

groups of aquatic insects in BC. Early lists of Coleoptera from various parts of the

Province (e.g. Keen 1895, 1905; Stace-Smith 1929, 1930) do contain some aquatic species,

but there has been no published provincial checklist, although a preliminary list was

developed by Scudder (unpublished). Most of the works dealing with aquatic beetles in BC

have been as parts of studies of particular taxa over larger geographical areas. References

to keys for aquatic beetles are in Bousquet (1991) and Arnett and Thomas (2001). Species

lists for BC are in Bousquet (1991) and Larson ef a/. (2000).

The most important person to study aquatic beetles in BC was H.B. Leech. Leech was

born in Kamloops, received much of his early education in Vernon, and graduated from

UBC in 1933. From 1930-1947, he worked at the Forest Entomology Laboratory in

Vernon where he collaborated with R. Hopping. In 1947, he joined the California

Academy of Sciences. He described his first new species in 1937, Agabus vancouverensis

Leech (Leech 1937), and subsequently described over 50 beetle taxa (Kavanaugh and

Arnaud 1981). His collection of over 30,000 specimens of aquatic beetles, many from BC,

was donated to the California Academy of Sciences in 1947. Several species of aquatic

beetles found in BC are named after Leech: Haliplus leechi Wallis, Helophorus leechi

McCorkle, and Cymbiodyta leechi Miller, which is known from Washington and probably

occurs in BC (Smetana 1988).

Several other people are associated with aquatic beetles in BC mostly as collectors; for

example, G.J. Spencer, R. Hopping, and G. Stace-Smith. M.H. Hatch described several

species of aquatic beetles found in BC and is best known for his “Beetles of the Pacific

Northwest” (Hatch 1953-1971). Each of them has at least one aquatic beetle from BC

named in his honor.

In more recent times, Scudder (1969b) reported some species from interior saline lakes,

and he, his students, and associates have collected aquatic beetles as part of more general

surveys and added to our knowledge of the distribution of various species, especially in the

Gulf Islands and in the northern part of the Province. In addition, J. Lancaster has

conducted an ecological study of aquatic beetles in a comparison of the community

structure in a series of saline lakes on the Chilcotin Plateau (Lancaster and Scudder 1987).

Kenner (2000d) described the distribution of two species of gyrinids in BC, the Yukon,

and Alaska.

DIPTERA

The Manual of Nearctic Diptera (McAlpine 1981, 1987) is a wonderful resource for

information on systematics and biology of North American Diptera. Illustrated keys

identify immature and adult specimens to family and genus. The taxonomic and

distributional status, to the early 1960s, of many British Columbia species is outlined in

Stone et al. (1965).

The Mountain Midges (Deuterophlebiidae) are peculiar, little-known flies found in

streams in the western mountains. Courtney (1990) documented the systematics of the

Nearctic species, including those known from BC.

In the Chaoboridae, Northcote (1964) and Teraguchi and Northcote (1966) studied the

vertical distribution and vertical migration of Chaoborus larvae in BC lakes. Taxonomic

works on the Phantom Midges that relate to the provincial fauna include Saether (1970)

and Borkent (1979, 1993).

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 69

Early work on the Culicidae in BC includes that of Hearle (1926, 1927) and Curtis

(1967). Belton (1978) studied the mosquitoes of Burnaby Lake, and more recently

produced a monograph on the 46 species known in BC (Belton 1983). This work, together

with Wood et al. (1979), can be used to identify these species.

In his study of the fauna of saline lakes on the Fraser Plateau, Scudder (1969b) found

that Aedes campestris Dyas and Knab occurred in the most saline of them. The osmotic

and ionic regulation in the larvae of this mosquito is therefore of interest; it has been

studied by Phillips and Meredith (1969a). They also investigated the active transport of

sodium and chloride by the anal papillae of the larvae (Phillips and Meredith 1969b) and

the electron microscopic structure of these organs (Meredith and Phillips 1973a, 1973b).

Regulation and active transport of magnesium by the Malphighian tubules of these larvae

was described by Kiceniuk and Phillips (1974) and Phillips and Maddrell (1974). They

also discovered that these tubules could actively transport sulphate ions (Maddrell and

Phillips 1975). This work led to comparative studies on other saline-water mosquito larvae

including Aedes dorsalis (Meigen) and A. togoi (Theobald) (Meredith and Phillips 1973c;

Strange et al. 1982, 1984; Strange and Phillips 1984), both of which occur in the Province.

Important reviews resulted from this research (Bradley and Phillips 1977; Phillips er al.

1978; Phillips and Bradley 1978; Strange and Phillips 1985; Ng and Phillips 1985).

Gammarid predation on Aedes togoi at Horseshoe Bay was reported by Hossack and

Costello (1979). Oviposition of Culex pipiens L. in water at different temperatures was

described by Gillespie and Belton (1980). Ishii and Belton (1984) provided evidence for

autogenous egg development in this species and Belton (1982) described the cuticular

vestiture of Aedes communes (DeGeer) and A. nevadensis Chapman and Burr.

The comprehensive treatment of the Black Flies (Simuliidae) of North America by P.

Adler, D. Currie, and D.M. Wood is nearing completion; when published, this work will

deal with all aspects of the systematics of BC species. Early studies by Curtis (1954) and

Shewell (1957, 1959) were followed by research by Peterson (1970), Mahrt (1982), Currie

and Adler (1986), and Corkum and Currie (1987). Currie’s work in Alberta (1986) and the

Yukon (1997) has direct application to the BC fauna. Studies on Parasimulium (Borkent

1992; Borkent and Currie 2001) have led to a better understanding of the phylogenetic

origins of the Simullidae. Currie and Walker (1992) documented the use of fossil black fly

larvae in paleoecological studies. Cytological research into the giant chromosomes of

simuliids has been undertaken in the genus Prosimulium (Basrur 1962), in Stegopterna

(Madahar 1969), in the Simulium (Eusimulium) vernum group (Hunter and Connolly

1986), and in the S. (E.) aureum group (Leonhardt 1985).

McMullen (1978) surveyed Ceratopogonidae in the Okanagan area of BC. Light trap

catches of Culicoides in the Fraser Valley were described by Costello (1982). Anderson

and Belton (1993) examined populations of Culicoides obsoletus Meigen in the lower

mainland of BC and hypersensitivity of horses to Culicoides bites was reported (Anderson

et al. 1988, 1991, 1993, 1996). A. Borkent of Enderby is a world authority on

Ceratopogonidae and some of his systematic studies directly relate to BC (Borkent and

Grogan 1995; Borkent 1998).

Both freshwater and saline-tolerant species of the Chironomidae inhabit saline lakes in

the BC interior (Scudder 1969b). At the University of Victoria, Morley and Ring (1972a,

1972b) studied the life histories, ecology, and identification of the intertidal chironomids

Paraclunio alaskensis Coquillett, Saunderia_ clavicornis (Saunders), S. marinus

(Saunders), and S. pacificus (Saunders). The detailed study of the occurrence of midges in

the littoral zone (R.A. Cannings and Scudder 1978) resulted in the discovery of a new

species of Chironomus (R.A. Cannings 1975a) and many species new to BC and Canada

(R.A Cannings 1975b). Subsequently, the physiology of this and other Chironomus species

was investigated (Sargent 1978) and the phenology compared (R.A. Cannings and Scudder

70 J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

1979). The ecology of one of these, C. tentans Fab., which has distinct chromosomal races

in the Province (Acton and Scudder 1971), was researched by Topping (1971, 1972) and,

experimentally, Topping (1969) and Topping and Acton (1976) examined the influence of

environmental factors on the frequency of certain chromosomal inversions. Eastern Brook

Trout were used as predators in this research. The chironomids of Marion Lake in the

Fraser Valley were studied by Hamilton (1965); Schultze and Northcote (1972)

documented chironomids as fish food in another coastal lake. Mundie (1971) studied their

diel drift in Robertson Creek, an artificial spawning channel and secondary outflow of

Great Central Lake on Vancouver Island, and found that larval drift rates increased almost

two-fold in darkness.

Brillia retifinis Saether is a multivoltine species, able to track shifts in resource

abundance in its habitat by virtue of its short generation time (Richardson 1991). The

influence of Brillia on the decomposition of conifer leaves in a coastal stream was reported

by Summerbell and R.A. Cannings (1981). Richardson and Kiffney (2000), in their study

of the response of aquatic insects to metal pollutants, found that chironomids (mainly B.

retifinis) showed no significant decrease in densities with increasing doses of copper, zinc,

and manganese, typical urban pollutants. Parkinson and Ring (1983) examined more

physiological adaptations of P. alaskensis to the marine environment. As part of a

provincial government attempt to control the introduced aquatic weed, Myriophyllum

spicatum L., in warm lakes and waterways, potential biological control agents were

screened. The chironomids found in this study were documented by Kangasniemi and

Oliver (1983). One of them, an unnamed, native species that feeds on the plant’s apical

buds, was subsequently described as Cricotopus myriophylli by Oliver (1984). Borkent’s

(1984) systematic work on the Sfenochironomus complex relates to the BC fauna. Ian

Walker and his students and colleagues (Walker et a/. 1991, 1993; Heinrichs et al. 1999;

Riick et al. 1998; Palmer 1998) have used chironomid fossils in lake sediments to

reconstruct paleoenvironments in British Columbia. The larvae of Canadian chironomids

can be identified to genus using Oliver and Roussel (1983).

CONCLUSION

The above clearly documents that there has been a dramatic increase in our knowledge

of the aquatic insects in British Columbia over the past century. However, much remains

to be done, with the Ephemeroptera in particular needing more attention. Additionally,

many areas of the province remain to be surveyed, such as the far North and the numerous

islands off the west coast. Many more interesting discoveries can be expected during the

next 100 years.

ACKNOWLEDGEMENTS

The authors thank the following people for their contributions to this paper: Peter

Belton, Art Borkent, Doug Currie, Tom Northcote, John Phillips, and John Richardson.

Thanks also to Peter Belton for his thoughtful and meticulous review of the manuscript.

His comments greatly improved our submission.

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 71

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Spiders (Araneae) and araneology in British Columbia

ROBERT G. BENNETT

BC MINISTRY OF FORESTS,

7380 PUCKLE ROAD, SAANICHTON, BC, CANADA V8M 1W4

“.. . spiders are ruthless storm troops in the matriarchal anarchy that is the arthropod

world: theirs is the most diverse, female-dominated, entirely predatory order on the face of

the earth. As such, spiders are key components of all ecosystems in which they live.”

Bennett 1999

THE SPIDERS...

In large part because of its cool temperate climate and the significant amount of

Pleistocene time much of it spent under a series of transcontinental ice cubes, Canada is

sadly lacking in arachnid diversity. Although six of the dozen or so orders of extant

arachnids are represented in the national fauna, only two (Acari, mites and ticks; and

Araneae, spiders) are significantly diverse in Canada and these contain only a small

proportion of the world acarine and spider species.

Canada contains 1,400-1,500 members of the world’s described fauna of nearly

40,000 spider species (Bennett 1999). With its tremendous range of biogeographical

diversity, British Columbia (BC) provides habitat for nearly half of these — over 650 spider

species are known to occur here (Table 1) and many are found nowhere else in Canada.

Canada has representatives of both mygalomorph (tarantulas and their kin) and

araneomorph (“true”) spiders. Mygalomorphs are most diverse in the subtropics and

tropics; few are found in northern hemisphere cool temperate climates and always as

northern populations of species more widespread to the south. Three of the four

mygalomorph families occurring in Canada are restricted to BC. A tiny funnel-web

weaving diplurid is known from a single locality near Creston. One small sheet-web

weaving mecicobothriid occurs in disjunct populations in remnant old Vancouver Island

wet forests. Two large antrodiaetid trapdoor spiders occur in BC: one is common in a wide

range of coastal habitats from old, wet forest to dry Garry oak meadows and suburban

lawns and gardens; the other is uncommonly encountered in the dry southern interior

valleys.

Similarly, haplogyne (“primitive”) araneomorphs are much more diverse and common

south of Canada. Of five families with Canadian species, two (telemids and segestriids) and

most of a third (pholcids) are found only in BC. The other Canadian haplogynes are

cosmopolitan synanthropes. A single, tiny telemid is rarely encountered but likely locally

abundant in deep litter in old forests. Extreme south-western BC is home to Canada’s

single tube-web weaving segestriid. Two swift, small and cryptic pholcids are uncommonly

encountered on the undersurface of rocks and other objects (often in apparent association

with Latrodectus widow spiders) in the hottest, driest parts of southern BC. A third

pholcid, the common and familiar “daddy-long-legs” cellar spider is found in older homes

throughout much of Canada. One feisty synanthropic dysderid makes meals of isopods in

yards and basements across southern Canada.

The world araneofauna (especially in the Holarctic) is dominated by entelegyne

(“higher”) araneomorphs. Among BC’s 643 known entelegynes, the crab-like philodromids

and thomisids, orb-web weaving araneids, cob-web weaving theridiids, and active and

84 J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

highly visually orienting salticid jumping spiders and lycosid wolf spiders are familiar to

most people with basic knowledge of natural history.

Table 1

Classification of spider families with Canadian and British Columbia representatives

(modified from Bennett 1999).

Approx. No. of

no. of known

species in _—_— species in

Canada BC

ORDER ARANEAE

Suborder Opisthothelae

Infraorder Mygalomorphae

Fornicephalae

Atypidae | 0

Antrodiaetidae 2 2

Tuberculotae

Mecicobothriidae ] l

Dipluridae l l

Infraorder Araneomorphae

Neocribellatae

Araneoclada

Haplogynae

“Scytodoidea”

Scytodidae l 0

Telemidae l l

Pholcidae 3 3

Dysderoidea

Segestriidae l l

Dysderidae l l

Entelegynae

Palpimanoidea

Mimetidae 6 2

Eresoidea

Oecobiidae l 0

“RTA Clade”

Lycosoidea

Lycosidae 110 47

Pisauridae i l

Oxyopidae 2 l

Dictynoidea

Dictynidae 75-80 29

Cybaeidae 12 1]

Hahniidae 16 15

Dionycha

Anyphaenidae i! 2

Liocranidae 18 5

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 85

Clubionidae 35 15

Corinnidae 1] )

Gnaphosidae 100 49

Zoridae l l

Philodromidae 47 33

Thomisidae 68 32

Salticidae 110 43

Amaurobioidea

Amaurobiidae 30 10

Titanoecidae 4 2

Agelenidae 1] 9

Orbiculariae

Deinopoidea

Uloboridae 3 l

Araneoidea

Nesticidae 2 l

Theridiidae 100 Bp

Theridiosomatidae l 0

Mysmenidae l l

Pimoidae 2 pi

Linyphtidae >500 230

Tetragnathidae 23 12

Araneidae 74 33

TOTAL ~ 1,400 653

Uncommon in Canada, palpimanoids are represented in BC by two species of

araneophagic mimetids found in the south. All other BC entelegynes are either

orbicularians (orb-weavers and their kin) or members of the “RTA clade” (spiders with a

distinctive process on the male pedipalpal tibia).

RTA clade diversity in Canada is dominated by dionychan (two-tarsal-clawed spiders)

and lycosoid (true wolf spiders and their relatives) hunters. The other RTA groups

(dictynoids and amaurobioids) are reasonably diverse web weavers but tend to be

overlooked because members are mostly cryptic, litter inhabitants and many are tiny

(except for a small number of amaurobiids and some synanthropic, introduced agelenids).

Lycosid wolf spiders often are the most abundant (but not necessarily most diverse)

ground dwelling predators in any open habitat from forest openings and coastal shorelines

to bogs and alpine meadows. Pitfall traps may be inundated, often by a single species. A

good sign of spring is the first appearance of myriad small, dark Pardosa spiders in

meadows as soon as the snow disappears. Nearly half of Canada’s lycosid species have

been found in BC. Of Canada’s seven known pisaurids (nursery-web or fishing spiders),

only one is found west of the Rockies. This is BC’s loss — two eastern pisaurids are

Canada’s largest spiders and contribute greatly to the excitement of shoreline life east of

the Rockies. One of Canada’s two oxyopid lynx spiders is common in some

agroecosystems and similar open habitat in southern BC.

Nearly half of Canada’s approximately 400 species of dionychans occur in BC.

Gnaphosid ground spiders, philodromid and thomisid crab spiders, and salticid jumping

spiders predominate with a combined total of well over 150 species in the province.

Gnaphosids are common ground dwelling hunters. Generally nocturnal and cryptic,

gnaphosids are infrequently encountered by general collectors but may be as abundant as

86 J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

lycosids in many habitats. A few are well-known synanthropes. In contrast, salticids are

often conspicuously coloured and diurnal. Although difficult and confusing taxonomically,

they are well known by the general public and regularly serve as photogenic subjects of

natural history essays. Thomisid and philodromid crab spiders are sit-and-wait, generally

cryptic, diurnal hunters. Thomisids are rather bristly, slow moving, ground dwelling and

very crab-like; philodromids are faster, hairier, less crab-like and mostly encountered

above the ground layer.

The remaining dionychans form a diverse assemblage of primarily nocturnal hunters

occupying a variety of habitats. Clubionids and anyphaenids are active on foliage while

corinnids and liocranids are ground dwellers. Some corinnids and liocranids are ant

mimics. Restricted to BC in Canada, Zoridae is a recent addition to our fauna. One species

of these relatively small, ground dwelling, lycosid-like spiders has expanded its range

northward into southern Vancouver Island and the south Okanagan Valley.

Most of Canada’s dictynoids (dictynids, cybaeids, and hahniids) and amaurobioids

(amaurobiids, titanoecids, and agelenids) weave more or less reduced sheet webs. Nearly

all cybaeids are restricted to BC in Canada. All cybaeids are forest floor litter inhabitants

and some are dominant but infrequently collected species where they occur. (It is

interesting to note that few cybaeids are found in eastern Nearctic forests. Coelotine

amaurobiids dominate cybaeid-type habitats there but are absent from western Nearctic

forests.) Half of Canada’s hahniids are restricted to BC. Like cybaeids, hahniids are

generally small, cryptic litter inhabitants and often abundant but infrequently collected.

Dictynidae is a large group of small to tiny, often exceedingly difficult to identify spiders.

Many are arboreal and produce distinctive cribellate silk; their small, “hackled” webs may

be abundant among conifer branch tips. Ecribellate dictynids are primarily litter inhabitants

and, not surprisingly, often abundant but overlooked.

Canada is home to a range of cribellate and ecribellate amaurobiids; a third of these

(all cribellate) occur in BC. Four are found nowhere else in Canada. Most amaurobiids are

forest floor spiders. A couple of beefy Ca//obius species are common under bark or in bark

crevices of coastal BC conifers; the exceedingly small and rare members of the genus

Zanomys may be ground dwelling rodent associates. Agelenids construct large, distinctive,

sheet-like funnel webs in a variety of habitats. In Canada, most species are widespread and

often abundant. Three closely related introduced agelenids are well known BC

synanthropes: one (Tegenaria domestica) is cosmopolitan, another (7. duellica) rivals the

eastern pisaurids as Canada’s largest spider and is very abundant around buildings,

disturbed areas, and beaches in south western BC; the third (7. agrestis, the undeservedly

infamous “hobo spider”) is rare but locally abundant at various sites in southern BC.

Canadian titanoecids are a small group of wide ranging, nondescript, small forest floor

species. Half occur in BC.

The remaining BC spiders (fully half the known fauna) are orbicularians. Uloborids

spin cribellate, radial sector webs and are uncommon in southern Canada. One species has

been found in BC. Nesticids are cryptic, rare, often troglobitic “ccomb-footed” cobweb

weavers. One of Canada’s two species is restricted to southern Vancouver Island, the other

is a European introduction in eastern provinces. Mysmenids are minute, rarely encountered

deep litter inhabitants and troglobites. Canada’s single mysmenid appears restricted to

southern Vancouver Island. In Canada, the long spiny-legged pimoids also are restricted to

BC. Two species hang underneath tangled sheet webs in relatively dark, humid places in

southern BC: one is common on the south coast, the other is rare in the southern interior.

The remaining orbicularians are widespread and much more diverse.

About half of Canada’s nearly 100 species of orb-web weaving araneids (garden

spiders) and tetragnathids (long-jawed orb weavers) are found in BC. Spinners of the

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 87

radially symmetrical sticky webs familiar to most people around the world, individuals of

certain species may be very abundant in fields or around homes. Theridiids are the

quintessential cobweb-weavers. Over half of Canada’s 100 theridiid species are found in

BC in a wide variety of habitats. Black widow (Latrodectus sp.), false black widow

(Steatoda spp.), and brown house (Achaearanea spp.) spiders are familiar to most British

Columbians. Individuals of the latter two genera are common in and on homes throughout

the province. Black widows are locally abundant in some areas of southern BC. The most

interesting theridiids often are small, inconspicuous soil and deep litter inhabitants.

Linyphiid species and individuals dominate the northern hemisphere, especially the

Holarctic region. Over a third of Canada’s araneofauna are linyphiids. At least 230 species

are known to occur in BC. Linyphiids may be exceedingly abundant in certain habitats,

especially meadows and old fields. Most are tiny to minute and famously difficult to

identify.

For more detailed information on Canadian and Nearctic spiders, see Roth (1993) and

Bennett (1999) and references therein.

... AND THE STUDY OF THEM...

Given that nearly half the total number of spider species known to occur in Canada are

found in (and often only in) British Columbia (Table 1) and the obvious importance of

spiders to all terrestrial and many aquatic ecosystems, one wonders why such fascinating

creatures have received little scientific attention in the province. No professional

arachnologist is employed anywhere in Canada currently and, in BC, few researchers have

ever seriously considered spiders or other arachnids. In BC, araneology has largely been

the realm of a few dedicated amateur natural historians.

A notable exception to this is A.L. Turnbull (Fig. 1). A pioneer of ecology in Canada,

Turnbull’s scientific career spanned most of the latter half of the 20" Century. His

professional interest in spiders was sparked by mid-century encounters with “vast numbers

of spiders” during spruce budworm (Choristoneura sp., Tortricidae) work in the vicinity of

Lillooet, BC. He subsequently studied spider ecology as an Oxford graduate student and

learned the arcane art of spider taxonomy at the American Museum of Natural History

under Willis Gertsch, one of the granddaddies of modern arachnology. Turnbull’s interests

in spider ecology and taxonomy were honed during a stint at the federal lab in Belleville,

Ontario. There C.D. Dondale, who went on to become Canada’s premier professional

arachnologist, eventually joined him. Turnbull returned to BC in the late 1960s to join the

faculty of the nascent Simon Fraser University. During his tenure there he published his

landmark summary review of spider ecology (Turnbull 1973). Following his retirement in

1982, failing eyesight has kept him from contributing further to araneology.

The difficulties and drawbacks of spider ecological studies noted by Turnbull (1973)

are still relevant today: ecologists necessarily are reliant upon sound taxonomy; the work of

taxonomists and systematists is often excellent but much taxonomy is deplorably produced

and presented (where are the taxonomy police when we need them?); taxonomic studies

should be undertaken at inclusive supraspecific levels; assessing species distributions 1s

relatively simple but quantifying populations and communities and drawing meaningful

ecological conclusions are exceedingly difficult to do well; spider ecology papers often are

of questionable merit and; authors tend (understandably) to overlook relevant papers not

published in their native language. Ecologists and taxonomists will do well to keep

Turnbull’s observations in mind.

88 J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

Figure 1. Bert Turnbull, a pioneer of spider ecology in Canada, was a long-time faculty

member of Simon Fraser University and is now in retirement in the Vancouver area.

In recent years at least one student at each of the Universities of Victoria, British

Columbia, and Northern British Columbia and Simon Fraser University have worked on

some aspect of local spider diversity or the ecology of single species. However, most of

this work is unpublished and all the students have gone on to other things.

Most published knowledge on BC spiders, other than species descriptions, is in the

form of faunal lists (Thorn 1967; West et al. 1984, 1988; Blades and Maier 1996). Much

of the raw data for these came from the undirected efforts of a small number of amateur

collectors working in only a few areas of BC. Thus, the spider fauna of large areas of BC

remains unknown.

Two early spider collectors in BC were English expatriates Reverend John H. Keen

and Marianne Hippesley-Clark. Better known as beetle collectors, both made important

spider collections. Around the end of the 19" Century, as an Anglican missionary Keen

collected in the vicinity of Massett and Metlakatla on the north coast of BC. His spiders

apparently went to Nathan Banks. Subsequently, J.H. Emerton described several new

species from these specimens including at least one, Diplostyla keeni, named in Keen’s

honour. Hippesley spent much of her life near Terrace. A challenge for spider collectors is

to come up with new specimens of some of the tiny linyphiids unseen since collected by

her.

George and Elizabeth Peckham collected BC jumping spiders late in the 19" Century.

Emerton made at least one foray into the colonial wilds of BC, apparently making it as far

west as the Yoho area. The famous duo of Ralph Chamberlin and Wilton Ivie collected in

various areas of BC in the 1930s and described a number of new species from BC material.

In later years, other well known arachnologists such as Takasuma Kurata, Boris Malkin

and, most recently, C.D. Dondale made important collections of BC spiders.

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 89

A relatively small, good collection of about 4,400 vials of identified BC spiders is

maintained at the Royal BC Museum. A large proportion of these specimens were amassed

by S.L. Neave from the Kyuquot region of Vancouver Island and donated to the Museum

in the late 1950s. The Museum collection has been considerably augmented by specimens

collected from various BC localities by Rick West (primarily in the 1980s) or during

studies conducted or directed by Dave Blades or Neville Winchester (1990s). The

Universities of Victoria and British Columbia possess smaller collections. Geoff Scudder

and some of his students have collected substantial numbers of spiders from the south

Okanagan and south coastal BC. Robb Bennett, Don Buckle, Walter Charles, Robert

Holmberg, Lee Humble, Douglas Knight, Robin Leech, Jeff Lemieux, Malcolm Martin,

and others also have produced significant collections of BC spiders.

These collection efforts resulted in the publication of a series of BC provincial spider

checklists. Thorn (1967) compiled the first comprehensive list of 212 species from Royal

BC Museum holdings. These represented a relatively small number of collection localities,

dominated by Kyuquot (Neave) and Masset/Metlakatla (Keen) records. Subsequently,

efforts led by Rick West boosted the number of known BC spiders to 433 (West ef al.

1984) and then 570 (West et a/. 1988). Recent large collections by Winchester and others

increased the list to 653 (Bennett et a/. unpublished data) by the time of this writing.

Apparently only one BC regional list has been produced: Blades and Maier (1996) listed

spiders and other arachnids collected in their general survey of south Okanagan arthropods.

Large areas and many specific habitats of BC remain uncollected and no doubt many

list additions are still to come, especially from northern areas and the deep south of BC. No

effort has been made to produce a comprehensive, habitat-specific spider inventory for any

area in BC. That new records can be made with relative ease is suggested by the following

examples: hundreds of specimens of a gnaphosid previously only known from a couple of

Washington specimens turned up in a simple pitfall study in Burnaby (see cover of Journal

of the Entomological Society of BC, Vol. 96, 1999), the first specimen of a new family

record for Canada came from the carpet of a provincial government office (Bennett and

Brumwell 1996), and a new species record for BC came from the bathtub of an Osoyoos

motel (Bennett unpublished data) in 2001.

Araneology in Canada at the start of the 21" Century is in a dismal state. Advancement

of our knowledge of spiders depends upon the uncoordinated work of amateurs and

occasional students provincially and, since the retirement of C.D. Dondale, nationally.

Ecologists seeking expert identification of their data points depend upon the volunteer

services of a small number of qualified amateurs — the majority of people with arachnid

expertise in North America receive little or no payment for application of their knowledge

(Coddington ef al. 1990). This may change with growing interest in biodiversity, non-

vertebrate endangered species, and habitat protection and restoration in Canada. But I’m

not holding my breath.

ACKNOWLEDGEMENTS

I thank Dave Raworth for suggesting this topic; Peter Belton for putting me in touch

with Bert Turnbull; and Bert Turnbull, Don Buckle, Dave Blades, Rick West, and Rob

Cannings for relevant discussions and information. For inspiration, I thank all those who

pursue non-economically important avenues of natural history research. I believe and hope

the ecological wealth of nations will, some day, be considered of equal or greater value

than their economic health.

90 J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

REFERENCES

Bennett, R.G. 1999. Canadian spider diversity and systematics. Newsletter of the Biological Survey of

Canada (Terrestrial Arthropods) 18 (1): 16-27.

Bennett, R.G. and L. Brumwell. 1996. Zora hespera in British Columbia: a new spider family record for

Canada (Araneae, Zoridae). Journal of the Entomological Society of British Columbia 93: 105-109.

Blades, D.C.A. and C.W. Maier. 1996. A survey of grassland and montane arthropods collected in the

southern Okanagan region of British Columbia. Journal of the Entomological Society of British

Columbia 93: 49-74.

Coddington, J.A., S.F. Larcher and J.C. Cokendolpher. 1990. The systematic status of Arachnida,

exclusive of Acari, in North America north of Mexico (Arachnida: Amblypygi, Araneae, Opiliones,

Palpigradi, Pseudoscorpiones, Ricinulei, Schizomida, Scorpiones, Solifugae, Uropygi). Pp. 5-20 In:

M. Kosztarab and C.W. Schaefer (Eds.), Systematics of the North American Insects and Arachnids:

Status and Needs. Virginia Agricultural Experiment Station Information Series 90-1. Blacksburg.

Roth, V.D. 1993. Spider Genera of North America, with Keys to Families and Genera, and a Guide to

Literature, 3™ ed. American Arachnological Society, Gainesville, 203 pp.

Thorn, E. 1967. Preliminary distributional list of the spiders of British Columbia. Report of the Provincial

Museum of Natural History and Anthropology of British Columbia. 17 pp.

Turnbull, A.L. 1973. Ecology of the true spiders (Araneomorphae). Annual Review of Entomology 18:

305-348.

West, R., C.D. Dondale and R.A. Ring. 1984. A revised checklist of the spiders (Araneae) of British

Columbia. Journal of the Entomological Society of British Columbia 81: 80-98.

West, R., C.D. Dondale and R.A. Ring. 1988. Additions to the revised checklist of the spiders (Araneae)

of British Columbia. Journal of the Entomological Society of British Columbia 85: 77-86.

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 9]

Arthropod introductions into British Columbia

— the past 50 years

DAVID R. GILLESPIE

PACIFIC AGRI-FOOD RESEARCH CENTRE

P.O. BOX 1000, AGASSIZ, BC, CANADA VOM 1A0

Introduced arthropods have affected, and continue to affect many different aspects of

life in BC. Social concerns arise from public reactions to both epidemics of introduced

pests, and to government actions against these pests. Economic losses to both individuals

and the provincial economy occur as a result of these introduced species. Finally,

ecological impacts of potentially far-reaching nature inevitably occur as a result of these

new additions. The extent of the impacts of these introductions spans forestry, agriculture,

urban landscapes, and human health, and is too great to allow detailed treatment in these

few pages. The effects of introduced species tend to develop over years, if not decades.

Some species that were introduced over 100 years ago are still with us. Some of these have

undergone important range extensions. Changes in agriculture or forest practices, and

shifts in quarantine policies have made these species more or less important. Some species,

i.e. natural enemies, have been deliberately introduced for biological control of pests.

Changing perceptions of the importance of the impacts of these introduced natural enemies

on native plants and arthropod communities has altered the way in which we view and

value these introductions.

My objectives in writing this were not to provide an exhaustive account of alien

insects in the province. Rather, I hope to touch on some of the significant events

surrounding introductions of insects over the past 50 years.

ALIEN SPECIES IN BIODIVERSITY

An important question one might ask is to what extent has the fauna of the province

been modified by the presence of introduced species? In their checklist of the Hemiptera of

Canada, Maw ef al. (2000) identify the introduced species of each genus, by province. In

the Cicadellidae, 42, or about 9%, of the 485 species present in BC are introduced. The

family Miridae has been particularly well studied in BC by Geoff Scudder and colleagues.

Among the subfamilies of the Miridae, 2 of 16 Bryocorinae, 0 of 28 Deraocorinae, 10 of

142 Mirinae, 7 of 67 Orthotylinae, and 12 of 79 Phylinae are introduced. Within the

Miridae as a whole, 9% are introduced. Among the Aphididae, 19 of 41 Aphis, 5 of 8

Acrythosiphum, and 8 of 10 Myzus are introduced, but only 2 of 31 Macrosiphum are of

exotic origin. Among the aquatic families, none of the Gerridae, Corixidae, or

Notonectidae are introduced.

Scudder and Kevan (1984) provided a checklist of the Orthopteroid insects of British

Columbia. Ten of the 130 species listed are identified as alien to the province. These

include cockroaches, such as Blatta orientalis L. and Periplaneta spp., and the praying

mantid Mantis religiosa religiosa L. Interestingly, all four of the species in Dermaptera are

alien, but none of the 74 species in the Order Orthoptera are introduced. To this list can be

added the Surinam cockroach, Pycnocelis surinamensis L. (Belton et al. 1986).

An estimate that from 8 to 10% of the insect species of BC are be alien might be

relatively accurate. The pest and beneficial species accidentally or deliberately introduced

into the province certainly have economic impacts. However, not all alien species are

pests. Since these aliens now form a substantial part of the provincial fauna, they are

92 J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

almost certainly having impacts in a wide variety of habitats in the province. These

impacts will have long-term environmental consequences and implications for

conservation.

ALIEN SPECIES AS PESTS

In a report on the status of economic entomology in British Columbia, R.C. Treherne

(1914), remarked that few of the pests troubling other agricultural areas of North America

occurred in British Columbia. He attributed this to the application of quarantine to restrict

the importation of these pests. Glendenning (1952), in a review of fifty years of

entomology in the Fraser Valley, noted that the pest complex had changed dramatically

following Treherne’s time, and that British Columbia agriculture was then plagued by

many insect pests. Most of these had been introduced into British Columbia since

Treherne’s (1914) report. The fifty years following Glendenning’s remarks have also seen

a dramatic change in the insect pest fauna in the province. The pages of the Journal of the

Entomological Society of British Columbia, and its antecedent, the Proceedings, are

liberally sprinkled with reports of new pest insects, and with comments on management of

recently introduced species. These pests range from species that have had dramatic impacts

on the face of agriculture and forestry in the province, to those whose presence is scarcely

of significance except to those directly affected. Some of the more noteworthy, interesting,

or just plain curious examples are treated in the following paragraphs.

The oriental fruit moth, Grapholita molesta (Bsk) (Lepidoptera: Tortricidae) was

introduced into the Okanagan valley in 1956. This serious pest of peaches was rapidly

quarantined and eradicated within 2 years (Touzeau and Neilson 1957, 1958). The oriental

fruit moth continues to be a pest in Washington and Oregon, to the south, but ongoing

quarantine measures have apparently prevented its re-invasion. It could be argued that this

was the single most significant introduction into the province in the past 50 years, since it

demonstrated the opportunities for quarantine and eradication provided by the geographic

isolation of the province. Eradication of the grape phylloxera, Phylloxera vitifoliae Fitch

(Hemiptera: Phylloxeridae), first detected in 1961 in the Okanagan, was attempted then

abandoned (Morgan et al. 1973). The application of sterile insect release methods against

the codling moth, Cydia pomonella (L.) and its eradication in the Similkameen Valley

(Proverbs and Newton 1975; Proverbs et a/. 1976) also took advantage of the geographic

isolation of the fruit-growing regions of the province. In the current iteration of sterile

insect release techniques to control codling moth in the Okanagan Valley, re-introduction

from outside of the treated regions, and thus quarantine, remains one of the most serious

problems for the program. The apple maggot, Rhagoletis pomonella (Walsh) (Diptera:

Rhagoletidae), continues to be conspicuous in its absence from the province despite its

presence in Washington State. Whether this is because of sound quarantine or good fortune

cannot be determined.

The most controversial attempt at control by eradication in this province is certainly

that against the gypsy moth, Lymantria dispar (L.) (Lepidoptera; Lymatriidae). The first of

the recent introductions into the province occurred in 1978, and proposals to eradicate it by

widespread application of pesticides to an urban area were met with considerable

resistance. Since then, a program of detection of new infestations by pheromone trapping

and eradication by application of Btk has been followed. The value of this program is

controversial, since some introductions appear to have become extinct without treatment,

and the potential for gypsy moth to cause economic impact through defoliation continues

to be disputed (Myers et al. 1998). A study of the potential range of gypsy moth in the

province (Hunter and Lindgren 1995) suggested that this pest would be restricted to the

southwest through a combination of climate effects and host availability, and that the garry

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 93

oak, Quercus garryana Douglas (Fugaceae) zone would be the most affected. The most

important impacts of the introductions of gypsy moth into BC have been the generation of

a vocal opposition to all eradication attempts, and a general erosion of the public

perception of both entomologists and biological control in the province.

The winter moth, Operophtera brumata L. was first found in British Columbia in

1977 in Victoria, BC, but had apparently been introduced some years earlier (Gillespie e¢

al. 1978). Two exotic parasitoids, Cyzenis albicans (Fall.) (Diptera: Tachinidae) and

Agrypon flaveolatum (Gravely) (Hymenoptera: Ichneumonidae) were introduced (Embree

and Otvos 1984). This pest was brought under control by the action of endemic generalist

predators, but is regulated under non-outbreak conditions by the exotic parasitoids (Roland

1994; Roland and Embree 1995). The winter moth has since spread to the lower mainland,

where it is a pest of highbush blueberry crops (Fitzpatrick et al. 1991; Sheppard eg al.

1990).

Severe defoliation of willows (shown here) and many other wild and domesticated trees

and shrubs was caused by winter moth, Operophtera brumata in the Victoria, British

Columbia area in the 1970s. Invasive pests such as this have long-term economic and

environmental consequences. Photo by N.V. Tonks (deceased)

One of the significant legacies of the introductions of the gypsy moth and winter moth

was the creation of the BC Plant Protection Advisory Council (BCPPAC). This inter-

agency group, comprising representatives from provincial and federal departments,

industry and universities, has facilitated communication and joint action on pests and pest

threats, and also provides a forum for review of proposals for introductions of biological

controls of weeds.

The European crane fly, Tipula paludosa Mg. (Diptera: Tipulidae) was first observed

as a serious pest problem in BC in 1964, and was probably introduced some years earlier

(Wilkinson and MacCarthy 1967). These authors noted heavy infestations causing

extensive damage to pasture and horticulture crops, and consequent extensive use of

organo-chlorine insecticides. Although releases of the parasitoid Siphona geniculata De

Geer (Diptera: Tachinidae) established successfully, there is no evidence that the decline in

T. paludosa numbers was due to the parasitoid (Wilkinson 1984). In all probability, the

94 J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

decline was due to the combined action of several diseases (Wilkinson 1984). Recently, a

second exotic cranefly species, 7. oleraceae L. has been found in the Fraser Valley

(Costello 1998) which may add to injury to horticultural and pasture crops in the area.

The European pine shoot moth, Rhyacionia buoliana (Schiff.) (Lepidoptera:

Tortricidae) and the balsam wooly adelgid, Adelges piceae (Ratzeburg) (Hemiptera:

Adelgidae) were both first reported in the 1960s and were considered serious threats to

forestry in the province (Harris and Wood 1967, Wood, 1968). Although A. piceae has

restricted the replant of Abies spp. (McMullen and Skovsgaard 1972; Carrow 1973), no

direct losses seem to have occurred. Similarly, R. buoliana does not appear to attack Pinus

contorta, the major economic pine species in the province. The oak-ribbed casebearer,

Buccalatrix ainsliella Murtf. (Lepidoptera: Lyoneliidae) has established on red oak,

Quercus rubra L. (Gelok et al. 1998) planted as roadside trees in Vancouver. Larvae

pupate on almost any surface available, regardless of make or model, causing some

consternation in upscale neighborhoods. Although this species does not constitute an

economic threat, it and other invasive oak herbivores pose a problem for conservation of

garry oak meadows. Of the current threats to BC forests from introduced pests, the

importation of wood-boring beetles in dunnage and crating seems to be the most dire (e.g.

Humphreys and Allen 1999; Wulf 1999). Based on literature searches, no introductions

appear to have established in BC forests.

Other pest introductions of note include the lettuce aphid, Nasonovia ribisnigri

(Mosley) (Homoptera: Aphididae) (Forbes and MacKenzie 1982), an asparagus aphid

Brachycolus asparagi Mordivillo (Homoptera: Aphididae) (Forbes 1981), the strawberry

tortrix, Acleris comariana (Zeller) (Lepidoptera: Tortricidae) (Cram 1973), and the

German yellowjacket wasp, Paravespula germanica (F.) (Hymenoptera: Vespidae)

(Gerber 1990). Recently, the viburnum leaf beetle, Pyrrhalta viburni (Paykull)

(Coleoptera: Chrysomelidae) and a hemerocallis gall midge, Contarinia quinquenota

(Diptera: Cecidomyiidae) were reported for the first time in BC (Anon 2001).

Accidental introduction of exotic species is not the only way in which introduced

species arrive. Deliberate introductions of biological control agents have also added to the

fauna. Biocontrol introductions in Canada, including those made into BC, have been

reviewed (Kelleher and Hulme 1984, Mason and Huber, in press). However, these works

do not necessarily account for the accidentally introductions and range extensions of

putatively beneficial species. Harmonia axyridis Pallas and Coccinella septempunctatum

L. (Coleoptera: Coccinellidae) come to mind as dramatic examples in the latter category,

but others have occurred recently, for example aphid parasitoids (Mackauer and Campbell

1972) and a staphylinid beetle (Puthz 1972). Range extensions have also resulted in

invasions of new pests, for example the western grape leafhopper, Erythroneura elegentula

Osborne (Hemiptera: Cicadellidae), (Philip 1998), and a tentiform leafminer,

Phyllonorycter mespilella (Hiibner) (Lepidoptera: Gracillariidae) (Cossentine and Jensen

1992);

The past 50 years of introductions should not be discussed in isolation from the entire

complex of alien species in the province. In fact, many of the most serious of our

agricultural pests arrived very early in the previous century, and continue as pests to this

day. Because of their long residence in the fauna of the province, it might even be argued

that these species are now endemic. Redistribution and range extensions of pests such as

European wireworms, Agriotes lineatus L. and A. obscurus L., (Coleoptera: Elateridae)

introduced at the turn of the previous century (Wilkinson ef a/. 1976; Vernon and Pats

1997), underscore the long-term costs of management of introduced pests.

Impacts of beneficial species introduced either deliberately or accidentally, on native

fauna are becoming a serious issue in pest management and conservation, yet there is no

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 95

comprehensive inventory of these species in our province, let alone the resources to

consider such important questions as current geographic and host range. With trends to

globalization of world trade in the last two decades of the 20th century, I expected to see

an explosion of reports of new pest species and exotic introductions in the scientific

literature of the past two decades. The reverse seems to be true — there appears to be a

noticeable decline in reporting of new pests in scientific journals in the 1990s. Four of the

species noted in the past 5 years have been reported in trade magazines and in-house

publications, not in mainstream scientific literature. I can imagine many reasons for these

trends. Is society becoming so "globalized" that there is no alarm over the appearance of

new pests, which are simply viewed in the context of pest management, not from the

perspective of potential disruption of native ecosystems? Have resources available to

entomologists in this province become so restricted that free publication in trade and in-

house magazines is the only option available? Have taxonomic and systematics resources

declined to the point that it is impossible for economic entomologists to obtain

authoritative consultations and identification on species of concern? Has the size of the

entomological community in the province finally declined to a point where we are no

longer able to recognize and address new threats to our health and food, fibre and forest

production? In view of the continuing loss of entomology professionals in the province,

the loss of resources for public-good research, and the emphasis on results-driven research

for clients, I fear that all of the above have some vestige of truth. Given the continued

threats that further introductions of alien species pose to the economic and social well-

being of the province, it is essential that professional and amateur entomologists continue

to place these issues in front of the public and argue for the much needed resources.

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Belton, P., G.S. Anderson and G.L. St. Hilaire. 1986. A record of the Surinam cockroach in Vancouver.

Journal of the Entomolgical Society of British Columbia 83: 73-74.

Carrow, J.R. 1973. Establishment and survival of balsam woolly aphid on second growth amabilis fir at

intermediate elevations. Bi-monthly-Research-Notes 29 (2): 10-11.

Cossentine, J.E. and L.B. Jensen. 1992. Establishment of Phyllonorycter mespilella (Hiibner)

(Lepidoptera: Gracillariidae) and its parasitoid, Pnigalio flavipes (Hymenoptera: Eulophidae) in fruit

orchards in the Okanagan and Similkameen Valleys of British Columbia. Journal of the

Entomological Society of British Columbia 89: 18-24.

Costello, R. 1998. New Leatherjacket/crane fly in BC. Crop Protection news 20 (2): 3-4.

Cram, W.T. 1973. Occurrence of the strawberry tortrix, Ac/eris comariana (Zeller), a new pest in British

Columbia (Lepidoptera: Tortricidae). Journal of the Entomological Society of British Columbia 70:

17.

Embree, D.G. and 1.8. Otvos. 1984. Operophtera brumata L., winter moth (Lepidoptera: Geometridae). In:

J.S. Kelleher and M.A. Hulme (Eds.) Biological Control Programmes against Insects and Weeds in

Canada 1969-1980. Commonwealth Agricultural Bureaux Farham Royal, Slough, England.

Fitzpatrick, S.M., J.T. Troubridge and B. Peterson. 1991. Distribution of European winter moth,

Operophtera brumata L. and Bruce spanworm, O. bruceata (Hulst), in the lower Fraser Valley,

British Columbia. Journal of the Entomological Society of British Columbia 88: 39-45.

Forbes, A.R. 1981. Brachycolus asparagi Mordvilko, a new aphid pest damaging asparagus in British

Columbia. Journal of the Entomological Society of British Columbia 78: 13-16.

Forbes, A.R. and J.R. Mackenzie 1982. The lettuce aphid, Nasonovia ribisnigri (Homoptera: Aphididae)

damaging lettuce crops in British Columbia. Journal of the Entomological Society of British

Columbia 79: 28-31.

Gelok, E., R. McGregor, D. Henderson and L. Poirier. 1998. Seasonal occurrence and parasitism of

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Gerber, H.S. 1990. Note on the occurrence of Paravespula germanica (Hymenoptera: Vespidae), in the

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87: 73-74.

Gillespie, D.R., T. Finlayson, N.V.Tonks and D.A. Ross. 1978. Occurrence of the winter moth,

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Humphreys, N. and E. Allen. 1999. Eight-spined spruce bark beetle - /ps typographus. Exotic forest pest

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Kelleher, J.S. and M.A. Hulme (Eds.). 1984. Biological Control Programmes against Insects and Weeds in

Canada 1969-1980. Commonwealth Agricultural Bureaux Farham Royal, Slough, England.

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McMullan, L.H. and J.P. Skovsgaard. 1972. Seasonal history of the balsam wooly aphid in Coastal British

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Myers, J.H., A. Savoie, and E. van Randen. 1998. Eradication and pest management. Annual Review of

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and fruit containers as a source of reinfestation. Journal of the Entomological Society of British

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Proverbs, M.D., J.R. Newton, D.M. Logan and F.E. Brinton. 1976. Codling moth control by release of

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Roland, J. 1994. After the decline: what maintains low winter moth density after successful biological

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thuringiensis Berliner ss. kurstaki on larvae of winter moth, Operophtera brumata (L.) (Lepidoptera:

Geometridae) attacking blueberry in the Lower Mainland of British Columbia. Journal of the

Entomological Society of British Columbia 87: 25-29.

Touzeau, W.D. and C.L. Neilson, 1957. Plans to eradicate oriental fruit moth in the Okanagan Valley, B.C.

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Entomological Society of British Columbia 4: 67-71.

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Wilkinson, A.T.S. 1984. Tipula paludosa Meigen, European cranefly (Diptera: Tipulidae). Pp. 85-88 In:

J.S. Kelleher, and M.A. Hulme (Eds.). Biological Control Programmes against Insects and Weeds in

Canada 1969-1980. Commonwealth Agricultural Bureaux Farham Royal, Slough, England.

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Agriotes obscurus L. in corn in British Columbia. Journal of the Entomological Society of British

Columbia 73: 3-5.

Wilkinson, A.T.S. and H.R. MacCarthy. 1967. The marsh cranefly, Tipula paludosa Mg., a new pest in

British Columbia (Diptera: Tipulidae). Journal of the Entomological Society of British Columbia 64:

29-34.

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the Entomological Society of British Columbia 65: 13-15.

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J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 99

Research in adaptations of arthropods

in British Columbia

RICHARD A. RING

BIOLOGY DEPARTMENT, UNIVERSITY OF VICTORIA,

VICTORIA, BC, CANADA V8W 3N5

One of the major problems in reviewing this topic is defining the term adaptation.

Unfortunately, biologists have often used the word adaptation to mean various, quite

different things. However, for the purpose of this paper I will use the definition accepted

by the majority of modern ecologists i.e. an adaptation is any morphological,

physiological, sensory, developmental or behavioural character that enhances survival and

reproduction success of an organism (Lincoln ef al. 1998). This subject matter was not

covered in any of the review papers of Volume 48 of the Journal in 1951 (50 years of

entomology in BC), so I will take it upon myself to cover the last 100 years of research on

arthropod adaptations in BC — not that there is much to report from before the 1960s!

Since many aspects of adaptation research are likely to be covered in the other invited

contributions to this issue, for example in forest entomology, insect population ecology,

behavioural and chemical ecology, etc. I will concentrate my efforts in the field of eco-

physiology which is, after all, the closest to my primary area of research. The paper will

include not only work done by BC entomologists but also work done on BC insects by

BC and other entomologists, and it will be written in chronological order, as far as is

possible.

The earliest contribution that I can find that pertains to eco-physiological adaptations in

insects of BC is the brief communication from Cockle (1917), where he explains some of

the behavioural adaptations of “snow fleas” (Mecoptera; Boreus californicus) and five

species of spiders to alpine conditions in the Kaslo area. In particular, he observed active

feeding behaviour at temperatures down to between —2° and -3°C. From there we jump 22

years to Gregson (1939) who, in another brief paper, describes some of the adaptive

features of that most Canadian of all insects, Grylloblatta campodeiformis. In the

Kamploops area, these grylloblattids appear on the surface of rocky ground at relatively

low altitudes (430-820 m) after the first frosts in November. They are active all winter,

feeding on hibernating moths, ladybugs, wasps, bugs and spiders and even on active

thysanurans and collembollans (in captivity they can be fed cockroaches!). They only

appear on the surface at or around 0°C. Above and below this temperature they seek shelter

in rock crevices, as they do also during summer and other warmer months of the year.

Their optimal foraging temperature is, in fact, 3.7°C with cold prostration setting in at —

6.2°C and heat prostration at 27.8°C. They die very soon at room temperatures around

22°C. Gregson thus demonstrated that these rock crawlers are adapted to a very narrow

temperature range for their feeding and other activities.

In 1944, R.W. Salt, working out of the Dominion Entomological Laboratory (now

Agriculture Canada) in Lethbridge, Alberta, studied the behavioural adaptations and

tolerance to cold in the larvae and pupae of the warble fly, Hypoderma lineatus, a serious

pest of cattle in BC and Alberta. Most of the experimental insects came from Kamloops,

BC. He found that both larvae and pupae were what we now call “freezing intolerant” i.e.

they avoid freezing by supercooling to low temperatures, in this case to between —20° and

~24°C, indicating that early—dropped warble grubs would be frozen and killed. However,

his behavioural studies showed that early-dropped grubs were immature forms and

incapable of reaching full maturity. Fully mature grubs were never observed to drop before

the danger of cold weather was past. These findings had important implications for

100 J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

stockmen regarding whether or not to use early chemical sprays to control warble fly

populations. R.W. Salt went on to become one of the “founding fathers” in the discipline

of insect cryobiology, and, in 1961, published his paper on the “Principles of insect cold-

hardiness” which remains the foundation in this field of study to the present day. Then

follows a paper by Morgan (1952), published in the same volume as the 50" anniversary

volume of the Proceedings of the Entomological Society of BC (Vol. 48). He describes the

effects of low winter temperatures on four species of orchard mites in the Okanagan and

Kootenay valleys after the coldest winter on record in this region of Western Canada (-28°

to —36°C). His experimental protocol involved collecting winter eggs from the field and

incubating them at temperatures above 0°C. His results showed that mortality increased

from 40% at -27°C to 100% at -40°C. A few years later, Downes (1956) investigated the

effects of the longest period of drought in southern BC — a continuous period of 95 days in

summer with not more than a trace of water — on the insects of southern Vancouver Island

and the adjacent mainland, particularly on some Hemiptera and Lepidoptera. Most

populations of insects declined, although aphids increased their activities on Garry oak —

probably due to the decline in populations of hemerobiids, chrysopids, coccinellids and

syrphids. The main problem, however, was the associated decline in insect food plant

production. The paper goes on to explain the harmful effects of the combination of heat

and desiccation stress, but populations of these insects recovered, more or less, within 3

years.

The late 1960s was a relatively active period, particularly in studies of adaptations in

forest insect pests. These studies, like most of the others described in the paper, were

aimed at elucidating the “inherent” adaptations of each species, that is the built—in ability

of the insect to adapt to its environment over time — including responses to changes in the

environment within its own life cycle. Ross (1966) examined the ability of the introduced

European pine shoot moth (Rhyacionia buoliana) to survive low winter temperatures in the

Vernon area, and demonstrated that overwintering larvae can survive -4° F (-20°C) which

would be sufficient to allow this exotic pest to survive winters in most parts of the

Okanagan Valley. In the meantime, Niholt (1967, 1969) was studying the depletion of fat

deposits during the long 7-month hibernation period of adult beetles of 7rypodendron

lineatum, an ambrosia beetle. Although his studies of fats were quantitative rather than

qualitative, he did provide insights into how fat reserves affect the vigour of populations of

these beetles during hibernation and the subsequent flight period and brood establishment.

At the same time, Dyer (1969, 1970) and Gray and Dyer (1972) were elucidating the

adaptive processes involved in diapause, cold tolerance, flight muscle degeneration and

behaviour in the overwintering survival of bark beetles, Dendroctonus spp. There was not

much further activity in this area of research during the remainder of the 1970s, although

VanderSar (1977) wrote a very brief account demonstrating the ability of spruce beetle

larvae (Pissodes strobi) to overwinter successfully in Sitka spruce leaders in coastal BC

(Port Renfrew, Vancouver Island). Across most of its range, Pissodes strobi normally

overwinters in the duff at the base of host trees and is thus protected from low winter

temperatures by an insulating blanket of snow. Later, Safranyik and Linton (1991, 1998)

studied the adaptations to low overwintering temperatures of the mountain pine beetle,

Dendroctonus ponderosae, and the pine engraver beetle, /ps pini, under both field (1991)

and experimental conditions (1998). Mortality to winter cold is one of the main factors

determining the distribution and abundance of mountain pine beetle in BC, and it has been

shown that overwintering larvae can withstand short exposures to —-38°C. However, during

autumn and spring, larvae are very susceptible to extreme cold, so unseasonably low

temperatures (lower than —26°C) at these times of year can cause widespread mortality.

Under these conditions, mountain pine beetle larvae as well as the pine engraver beetle can

only survive extreme low winter temperatures at the base of trees or among the duff where

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 101

they are protected by a blanket of snow. In 1998, Li and Otvos, in attempting to provide a

plentiful laboratory supply of western spruce budworm pupae (Choristoneura occidentalis)

for research, reported their findings on adaptations to cold. Normally, western spruce

budworm overwinters in obligatory diapause in the second larval instar in the field.

However, a non-diapausing colony of this species can be induced in the laboratory and

reared on an artificial diet. This has certain advantages for mass-rearing of a species for

research purposes, and perhaps, for cold storage of beneficial biocontrol agents. Their

results indicated that western spruce budworm pupae could be stored at low temperatures

(2°C) for up to 1 week without deterioration in subsequent adult quality.

Similar motives were involved in the work of Gillespie and Ramey (1988) and

Morewood (1992a). Based on Morgan’s (1952) work, these authors studied the cold

hardiness and the cold storage potential of predatory mites (Amblyseius cucumeris and

Phytoseiulus persimilis) as biocontrol agents that could be used as the controls for

phytophagous mites and the western flower thrips, Frankliniella occidentalis. Gillespie

and Ramey (1988) were able to show that A. cucumeris could survive at 9°C for 10 weeks

(63% survival) whereas only 1.2% survived 10 weeks at 2°C. Similarly, Morewood’s

(1992b) techniques showed that although both species could survive short periods (e.g. 30

min) at —12.5°C, all individuals died within 75 min of exposure to that temperature. He

also calculated that under cold storage conditions, A. cucumeris could survive 2-3 weeks at

7.5°C (the optimal temperature) whereas P. persimilis could survive 4-6 weeks under the

same conditions. Gilkeson (1990) also worked on the problem of cold storage with the

predatory midge, Aphidoletes aphidimyza, an important insect predator of aphids in

greenhouses. The main aim of her cold storage program was to facilitate the balance

between supply and demand in the biocontrol agent market.

The contributions from my laboratory at the University of Victoria have made many

important advances in the field of insect cold-hardiness and _ eco-physiology

(environmental physiology) in the last 20 years or so. With my graduate students and co-

workers, we have attracted two international symposia on insect cold tolerance to the

University of Victoria (in 1985 and 2000), the annual meeting of the Society for

Cryobiology (1980) (where there was a special section on this topic), and a session at the

International Congress of Entomology in Vancouver (1988). Most of this work is in the

area of adaptations to low temperatures and desiccation resistance. Although many of these

studies were conducted in the Canadian Arctic, some include insects from BC, such as the

thimbleberry gall wasp (Diastrophus kincaidi) (Ring 1981), a willow leaf-gall sawfly

(Pontania sp.) (Ring 1981), the cabbage-root maggot (Delia radicum) (Turnock ef al.

1998), a pythid xylophagous beetle from high altitudes (Pytho deplanatus) (Ring 1981),

the introduced European winter moth (Operophtera brumata), (Hale 1989; Ring and

Danks 1994, 1998), the aphid, Myzus persicae (O’Doherty and Ring 1987) and several

indigenous species of intertidal insects (Morley and Ring 1972; Parkinson and Ring 1983;

Topp and Ring 1988a and 1988b; Ring 1989). During this time, I was also a collaborator

in a comparative study carried out on the sub-antarctic island of South Georgia with the

British Antarctic Survey, studying the adaptations of two endemic perimylopid beetles

(Perimylops antarcticus and Hydromedion sparsutum) to low winter temperatures and

relatively short summer growing seasons (Block ef a/. 1988; Ring ef al. 1990).

Some of the major contributions from my laboratory have been: (1) identification of a

multi-component cryoprotective system in the successful overwintering of insects,

including a combination of glycerol, trehalose and sorbital (Ring 1977; Ring and Tesar

1980, 1981); (2) the discovery of the lowest supercooling point ever recorded for an insect,

Pytho americanus, in the Western Canadian Arctic (Inuvik) at -—61°C; and (3)

identification of the various anomalies that exist in northern insects, such as not only being

freezing tolerant but also having a very low supercooling point (Ring 1982a). There are

102 J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

several other anomalies awaiting elucidation, such as the winter survival in the arctic of

coccinellid beetles, which, apparently, lack any known cryoprotectant molecules (Ring

1982b). Humble (1987) made an important attempt to tease apart the co-evolutionary

problems of cold versus desiccation tolerance and/or resistance. In arctic sawflies, he

demonstrated that their abilities to survive low winter temperature and desiccation stress

are co-adapted, that is, they are overlapping adaptations (see also Ring and Danks 1994,

1998). Similarly, Morewood (1999) studied the life history strategies and temperature

versus development relationships of the high arctic woolly bear caterpillars (Gynaephora

spp.) and their parasitoids at Alexandra Fiord, Ellesmere Island, Nunavut. Other

contributions made in my laboratory towards an understanding of the adaptations of

insects to the extreme temperature conditions of the arctic were by Winchester (1984) on

arctic trichopteran larvae of the Tuktoyaktuk Peninsula in the western arctic, by Humble

and Ring (1985) and Humble (1987) who studied the overwintering behaviour and

adaptations of arctic willow sawflies and their parasitoids, and by DeBruyn and Ring

(1999) on the overwintering behaviour and habitats of two species of diving beetles,

Hydroporus spp. (Dytiscidae) in ponds at Alexandra Fiord, Ellesmere Island.

In the miscellaneous category, we find contributions from a variety of sources. At the

top of the list I place G.G.E. Scudder, University of British Columbia (who has made three

other contributions to this volume) and his collaborators who have made many insights

into the ecological adaptations of water boatmen (Corixidae) that have successfully

inhabited the saline lakes of Interior BC (Scudder 1976). With his students, he recognizes

that many insects can, indeed, overcome the osmotic problems associated with these

highly saline lakes — many of which are more saline than the sea — once again giving rise

to that perennial entomological question “why have insects not re-invaded the sea?”

Certainly not for the lack of osmoregulatory adaptations, according to this research. At the

same time at UBC, J. Phillips and P. Hochachka were working on “pure” insect

physiology. Nevertheless, much of their research also contributed knowledge towards our

understanding of insect adaptations in the area of eco-physiology (see Hochachka and

Somero 1973; Phillips 1975).

Ring (1978) studied the adaptive significance of spiracular gills in the pupae of the

intertidal crane fly, Limonia marmorata (Tipulidae) (Fig. 1). This is a spectacular

morphological adaptation which allows the pupae to respire both above and below water —

an obvious advantage for any insect that lives in the intertidal zone. Other work that has

taken place in the Ring laboratory on insect adaptations to the intertidal habitat is that of

Morley and Ring (1972) on life history characteristics of intertidal chironomids, Parkinson

and Ring (1983) on the osmoregulatory adaptations of these marine chironomids, and

Topp and Ring (1988a, 1988b) on the adaptations of staphylinid beetles to both rocky

shores and sandy beaches of the marine environment (Fig. 2). There then follows a paper

(Ring 1991) that deals with insects that live in the natural hot springs of Hotspring Island,

BC. These insects not only have to deal with wide temperature fluctuations but also with

the associated osmoregulatory problems — the water of these natural springs is 2.5 times

more saline than the sea. The chironomid midge larvae (Thalassosmittia pacifica) survive

by being osmotic regulators in the surrounding medium rather than osmotic conformers.

Other adaptations of marine chironomid larvae involve the up-take of dissolved organic

nutrients from the surrounding seawater (Ring 1989). This is probably one of the least

resolved but yet most intriguing questions in marine invertebrate zoology. My research has

shown that these intertidal chironomids can utilize dissolved organic nutrients such as

glucose, amino acids, etc. to enhance their nutrition. This is done by absorption through

the cuticle of specialized areas in the intersegmental membranes of the larvae. This

phenomenon may be more widespread among arthropods than was formerly recognized

(see Chapman 1981).

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 103

Figure 1. Morphological adaptation — the spiracular gills of the pupa of the intertidal

tipulid, Limonia marmorata. This structure allows the pupa to respire both above the water

(low tide) and below the water level (high tide).

Figure 2. A typical view of the rocky intertidal zone on the west coast of British

Columbia. Many insects exploit this habitat and show a whole suite of adaptations for

survival in this extreme, inhospitable environment.

104 J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

Another unexpected but interesting structural adaptation found among herbivorous

insects going about their daily lives of eating plants, is the fact that many of them (if not

all) incorporate metals into their already sclerotized mouthparts (mandibles) and claws. A

good example of this can be found in Fontaine et a/. (1991) where they demonstrated that

zinc is prominent in the mandibles and claws of the mountain pine beetle (D. ponderosae),

beetles which tunnel through bark as adults and as larvae mine tissues of the inner bark.

Similar results were obtained from several species of coneworms (Lepidoptera: Pyralidae)

where larvae mine cones and feed on developing seeds — all plant structures which have

developed considerable toughness.

In summary, this paper provides, to the best of my knowledge, an up-date on the last

100 years of research in the adaptations of arthropods (mainly insects) in BC. It chronicles

those papers dealing with various aspects of eco-physiology, but also includes work

carried out on morphological/structural and behavioural characteristics. About one-third of

the papers have been published in the Journal (or Proceedings) of the Entomological

Society of BC. Please excuse me, reader, if I have not included your seminal work in this

article!

REFERENCES

Block, W., L. Somme, R.A. Ring, P. Ottesen and M.R. Worland. 1988. Adaptations of arthropods to the

sub-antarctic environment. British Antarctic Survey Bulletin 81: 65-67.

Chapman, P.M. 1981. Evidence for dissolved glucose uptake from seawater by Neocalanus plumchrus

(Arthropoda, Copepoda). Canadian Journal of Zoology 59: 1618-1621.

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J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 107

Insect population ecology in British Columbia

J.H. MYERS

DEPARTMENT OF ZOOLOGY, UNIVERSITY OF BRITISH COLUMBIA

6270 UNIVERSITY BLVD, VANCOUVER, BC, CANADA V6T 1Z4

D.A. RAWORTH

AGRICULTURE AND AGRI-FOOD CANADA,

PACIFIC AGRI-FOOD RESEARCH CENTRE, AGASSIZ, BC, CANADA VOM 1A0

The heydays of population ecology in British Columbia occurred during the 1970s and

1980s. Significant work was conducted by many people, and at several institutions, including:

the University of British Columbia; Institute of Animal Resource Ecology; Simon Fraser

University; Agriculture Canada, Vancouver Research Station; and the Canadian Forest

Service, Pacific Forestry Centre. This paper reviews some of the work, from the viewpoint of

the authors. The first section relates to agricultural insects and the second focuses on forest

Lepidoptera in BC.

Population ecology of agricultural insects

Many researchers in BC have studied various aspects of population ecology. Rather than

list these people and their contributions, this section focuses on the work of one research group

— allowing detailed development of a story. The group was selected because it treated

population ecology as a subject in its own right. This approach to ecological studies was

prevalent in the 1960s-1980s, and is quite distinct from the approach that currently dominates,

in which commercial problems drive much of the research. The title of this section reflects the

current approach; the group being considered took quite a different view.

The research group consisted of Neil Gilbert (Associate Professor, Institute of Animal

Resource Ecology), Andrew Gutierrez (Visiting Professor from Purdue University; later

Professor, University of California, Berkeley), Bryan Frazer (Research Scientist, Agriculture

Canada, Vancouver Research Station), post-doctoral fellows Rhonda Jones and Penny Ives,

University of British Columbia graduate students, and summer students.

The researchers selected organisms with characteristics that facilitated quantitative study of

ecological relationships. They then used a reductionist approach, quantitatively defining

relationships and processes at lower scales and combining those quantitative relations in

simulation models to explain population trends (Gilbert e¢ a/. 1976). A characteristic of the

work was the constant interplay between experiments in the laboratory, sampling, observations

and experiments in the field, and modeling, to achieve a realistic, dynamic picture of natural

systems. The approach was logical, it worked, and it still applies today. However, the book,

Gilbert et a/. (1976) met with considerable criticism (Lawton 1977), largely because it took to

task current and past approaches to population ecology. In particular, Gilbert et al. (1976)

objected to ecological studies that focused on the organism rather than underlying ecological

relationships that can be generalized among organisms. They also objected to laboratory-

centred population ecology and theoretical approaches that are not tested under field

conditions.

One of the first organisms selected by the group was the aphid Masonaphis maxima

(Mason) (Frazer and Forbes 1968). This is a single-host aphid that appears on thimbleberry in

sheltered locations in April, goes through 3-5 generations parthenogenically depending on

108 J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

plant quality and weather, and produces overwintering eggs in July. Because a special morph is

needed to produce sexuals, and sexuals are necessary to obtain eggs, the aphid must be able to

respond to changes in plant quality two generations in advance. A very good understanding of

the population dynamics of the aphid has been achieved (Gilbert and Gutierrez 1973; Gilbert

1980), and the work - based on this non-commercial system - provided the seeds for many of

the applied results obtained by Andrew Gutierrez and co-workers on cotton, coffee, alfalfa,

cassava and other crops (e.g. Gutierrez and Baumgartner 1984). In addition, the aphid, due to

a peculiarity in its life cycle, provides a convincing proof for one function of sex. Sex is

required to maintain dimorphic fundatrix aphids (Gilbert and Raworth 1998), and the

dimorphism, a differential production of sexual forms at the end of the season, adapts the

aphid to its heterogeneous and unpredictable environment. This is as yet, the only proven

function for sex.

Movement is a key element in the population dynamics of an insect. To study movement,

the group, which included Rhonda Jones, Michael Guppy and Vince Nealis chose to study the

cabbage butterfly, Pieris rapae (L.) (Jones 1977). Rules were obtained for short-distance

movement, and these rules were then used to predict long-distance movement (Jones ef al.

1980). The butterfly also proved amenable for the study of other aspects of population ecology

and resulted in a series of papers on the control of fecundity (e.g. Gilbert 1988) and a series of

papers on insects and temperature, one of which, Gilbert and Raworth (1996), presents a

general theory.

The effect of predation on population dynamics was studied using the pea aphid,

Acyrthosiphon pisum (Harris) on alfalfa (lucerne) and ladybird beetles, Coccinella spp. The

resulting Gilbert-Frazer predator-prey model represented a major advance in population

ecology, and the work produced one of the first cases in which the population dynamics of an

insect in the field were explained by quantitative assessment of predation processes at lower

levels (Frazer and Gilbert 1976). A key result was that, contrary to current theory, the

population dynamics of the aphid are intrinsically unstable, being determined largely by

weather via its effects on predation. Having considerable experience with the system, the

group conducted an interesting applied study when Visiting Scientist, Eric Charnov arrived in

the mid 1970's. The researchers set up several types of fisheries within an alfalfa field (four

different, constant rates, and two increasing rates, of exploitation) - the pea aphid played the

part of the fish. Analyses of catches in the various fisheries showed how compensatory

mortality within the system can affect catch-versus-effort data and result in overestimation of

potential catches based on those data (Charnov et al. 1976). Penny Ives joined the group in the

late 1970s conducting detailed work on the estimation of coccinellid numbers, beetle

movement in the field, and the relationship between feeding rate and egg production. The work

culminated in a collection of nine papers covering the complete predator-prey relationship

(Baumgartner et al. 1981).

In order to generalize about population processes it was necessary not only to compare

studies of different organisms, but also the same organism in different parts of the world. This

was done with many of the studies of P. rapae. In addition, David Raworth undertook studies

of the population dynamics of the cabbage aphid, Brevicoryne brassicae (L.) (e.g. Raworth

1984), an aphid that had been intensively studied in Australia by Dick Hughes at CSIRO; and

Vince Nealis undertook comparative studies of the ecology of Cotesia rubecula, a parasite of

P. rapae, in Vancouver, BC and Canberra, Australia (Nealis 1985).

What were the circumstances that led to this group? The work arose from the interests and

dedication of the people involved. Inspiration came from the three lead researchers: Neil

Gilbert, who trained at Cambridge as a biometrician under R.A. Fisher; Andrew Gutierrez who

trained in population ecology at UC Berkeley under R. van den Bosch; and Bryan Frazer, who

trained in entomology at UC Berkeley under R. van den Bosch. It was Neil Gilbert’s unique

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 109

view of population ecology, his insight that focused on fundamental ecological questions, his

rigorous approach in ‘consulting the animals and plants’, and his drive, that guided much of

the work. C.S. (Buzz) Holling (Director of the IARE) recognized the importance of Neil’s

views and supported him in a research position that required little teaching - although many

students will attest to the influence Neil had on their work. The group was influenced by other

workers who were making important advances in quantitative population ecology at the time,

for example: C.S. Holling had taken a quantitative, reductionist approach to predation

processes (Holling 1965); and R.F. Morris produced a series of papers on the quantitative

assessment of the population dynamics of fall webworm, Hyphantria cunea Drury (e.g. Morris

1971). Advances in computer technology very much facilitated the work. Government-funded

summer employment programmes provided the many hands that are necessary to conduct field

studies in population ecology. Finally, the research was driven predominantly by scientific

considerations — this required a unique economic, and administrative climate.

The work is not finished. Long-term, science-directed studies in quantitative population

ecology are still very much needed to gain further insights into the complex relationships and

processes that result in the various scale-dependent patterns of insect distribution and

abundance.

Population ecology of forest Lepidoptera

The history of forest entomology in British Columbia can be organized around the species

of insect under study. In the mid-1950s W.G. Wellington moved to the Pacific Forestry

Laboratory and began working on the western tent caterpillar, Malacosoma _ pluviale

californicum (Dyar). In his previous work Wellington developed his ideas on the interactions

between insects and weather. In BC he was welcomed to an outbreak of tent caterpillars on the

Saanich Peninsula of Vancouver Island between 1955-57. Here Wellington monitored changes

in both the numbers and frequency of tent types; elongate tents were thought to characterize

active groups and compact tents sluggish groups (Wellington 1960, 1964, 1965). He described

how simple behavioral tests could distinguish the “quality” of individuals and how the

composition of families in terms of how many active or sluggish individuals occurred, could

influence their success. He also related this to the impacts of climate, parasitoids, and disease.

During the 1970s Wellington and colleagues at the Institute of Animal Resource Ecology at

UBC (Ilan Vertinsky and Bill Thompson) explored the potential interactions between

heterogeneity in larval quality, climate, and population density of tent caterpillars using

simulation models. A recent review of the tent caterpillar work (Myers 2000) looks at which of

the early observations have been repeatable in subsequent population cycles. Tent shape does

not seem to be a good predictor of population condition, but a common observation is a

sudden invasion of new habitat sites associated with peak populations. Work on the viral

disease that occurs in peak populations has continued, as has work on the impacts of weather.

Wellington was a Strong force in encouraging people to recognize that all individuals in a

population are not the same and that those in increasing populations may be quite different

from those in declining populations. The actual causes of population decline may be less

important than the condition of individuals within the population. These ideas have important

ramifications for the simple density-dependence approach to population ecology. In addition

he emphasized how geographic variation in weather patterns could have long-term influences

On insect populations and distributions. These ideas take on new relevance with the current

interest in global climate change.

During the 1970s, outbreaks of both the eastern, Choristoneura fumiferana (Clemens), and

western spruce budworms, C. occidentalis Freeman, attracted the attention of insect ecologists

110 J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

across the continent. C.S. Holling and Don Ludwig (Ludwig, et al. 1978) spearheaded

modeling efforts to explore the dynamics of these species (McNamee, ef al. 1981). This

involved the development of both very complicated simulation models and very simple

deterministic models. At this time there was considerable interest in catastrophe theory and the

idea that there could be multiple stable states at low and high densities, with very rapid

transitions between. Part of this idea was that insects could be maintained in a low-density

“predator pit” until good conditions allowed them to break out and move to a new, high-

density equilibrium.

Other work on the western spruce budworm was initiated from the Pacific Forestry Centre

by Roy Shepherd. In the mid-1970s a prolonged outbreak of populations in the vicinity of

Hope led to calls for a spray program. Like many spray programs this was controversial

because the insect ecologists considered the populations to be on the verge of collapse. The

newspapers were filled with controversial articles and Shepherd was on the hot seat.

Shepherd’s group did considerable work on the western spruce budworm including trials with

viral sprays and Bt.

Douglas Fir Tussock moth, Orgyia pseudotsugata (McDunnough), is another forest

defoliator with cyclic population dynamics, and a periodic pest in BC. Imrie Otvos and

Shepherd initiated trials with a virus spray that showed that it was possible to stop an outbreak

(Shepherd, et a/. 1984). Viral sprays are still in the toolbox of potential controls for Tussock

moth. These trials are excellent examples of applied insect ecology. Shepherd also carried out

considerable work on monitoring tussock moth including developing pheromone systems to

allow predictions of impending outbreaks.

The introduced winter moth, Operophtera brumata (L.), made a mark on insect ecology in

BC starting in the late 1960s. By the early1970s what was thought to be an outbreak of the

native Bruce’s spanworm, O. bruceata (Hulst), continued to cause unsightly defoliation of oak

trees in Victoria. Finally, studies by Dave Gillespie and Thelma Finlayson (Gillespie e¢ al.

1981) on the parasitoids of the caterpillars led to the realization that this was an exotic species

and therefore lacking in parasitoids. Winter moth had been successfully controlled in Nova

Scotia by the introduction of a parasitoid fly, Cyzenis albicans (Fall.) and the experiment was

replicated in Victoria. Within 5 years of the original fly introductions winter moth populations

declined. Jens Roland did his Ph.D. research on this successful biological control program and

showed through his experiments and observations how ground predators and parasitoids

interact to maintain winter moth at densities higher than in their native habitat but much lower

than during their initial outbreak (Roland 1988, 1994; Roland and Embree 1995). Studies

continued when winter moth became a pest of blueberries on the lower mainland (Horgan et al.

1999). Imrie Otvos was given the keys to the city by an appreciative Victoria city council for

the successful biological control of winter moth.

Gypsy moth, Lymantria dispar (L.), has also presented challenges to insect ecologists in

BC. This exotic, European species, which is well established in eastern North America

continues to show up in BC. The first discovery of the species was in the early 1900s when the

Asian strain was found. However, it wasn’t until 1978 that a major introduction was

recognized in Kitsilano. The challenge of this species to insect ecologists has been to detect

introductions and coordinate spray programs. Many introductions have either gone extinct or

have been eradicated with Bt sprays (Myers and Rothman 1995). Again these spray programs

have been controversial and have involved considerable attention of insect ecologists.

One of the fine aspects of population studies of forest insects was the Forest Insect and

Disease Surveys that were done annually from the 1930s to the mid-1990s. These data bases

provided valuable long-term information on population trends of many of British Columbia’s

forest caterpillars. That they have stopped just as the world is becoming increasingly

interested in the impacts of global change is a shame.

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 11]

ACKNOWLEDGEMENTS

Thanks to Andrew Gutierrez for reading the section on the population ecology of

agricultural insects and contributing additional information.

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Roland, J. and D.G. Embree. 1995. Biological control of the winter moth. Annual Review of Entomology

40:475-492.

Shepherd, R., I. Otvos, R. Chorney and J. Cunningham. 1984. Pest management of Douglas-fir tussock moth

Orgyia pseudotsugata (Lepidoptera: Lymantriidae): prevention of an outbreak through early treatment with a

nuclear polyhedrosis virus by ground and aerial applications. The Canadian Entomologist 116: 1533-1542.

Wellington, W.G. 1960. Qualitative changes in natural populations during changes in abundance. Canadian

Journal of Zoology 38: 289-314.

Wellington, W.G. 1964. Qualitative changes in populations in unstable environments. The Canadian

Entomologist 96: 436-451.

Wellington, W.G. 1965. Some maternal influences on progeny quality in the western tent caterpillar,

Malacosoma pluviale (Dyar). The Canadian Entomologist 97: 1-14.

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 113

Behavioral and chemical ecology in British Columbia

B. ROITBERG AND G. GRIES

CENTRE FOR ENVIRONMENTAL BIOLOGY,

DEPARTMENT OF BIOLOGICAL SCIENCES, SIMON FRASER UNIVERSITY,

8888 UNIVERSITY DRIVE, BURNABY, BC, CANADA V5A 1S6

Behavior can be broadly defined as the response of an individual to a change in its

environment. During the past 50 years many ESBC members have studied insect behavior

both directly and indirectly. Below we chronicle two major schools of behavior, (1) basic

studies and (2) research on aspects of pheromone production and response.

Behavior studies are represented in almost every volume of the past 50 issues of

JESBC yet they are never common. In fact, excluding the pheromone-based studies, papers

that deal directly with behavior make up Just 6% (27/452) of all the papers published

between the years 1968 and 2000. This low frequency may not be unique to JESBC. A

survey of The Canadian Entomologist between 1980 and 1983 gives a nearly identical

frequency of behavior-based papers (29/505).

Our survey focuses on papers that study behavior for its own sake. Many other papers

have behavioral components or evaluate phenomena that are driven by behavior (e.g. trap

captures - Vernon and Gillespie 1990). Either way, behavior-based studies that are

published in the JESBC are drawn from a wide variety of insect taxa including, Hempitera,

Homoptera, Thysanoptera, Hymenoptera, Diptera, Lepidoptera and Coleoptera. The topics

range from oviposition preference in pear psylla (Stuart et a/. 1989) to predator avoidance

in fireworms (Fitzpatrick et al. 1994).

Behavioral ecology

There are five significant developments during the past fifty years in behavioral

entomology in British Columbia. First, in the mid-1960s, Bill Wellington and C.S. (Buzz)

Holling arrived at the University of British Columbia (UBC) from the Canadian Forest

Service. Wellington’s studies on the maternal effects on the behavior of forest caterpillars

are classic while Holling’s derivation (Holling 1966) of the functional response has

spawned a veritable cottage industry. Additional UBC faculty with behavior based

programs include, Ken Graham (behavior of bark beetles), Geoff Scudder (behavior of

various hemipterans), Judy Myers (behavior underlying population processes), Bob Elliot

(grasshopper feeding), Murray Isman (impact of natural products on insect feeding), John

McLean (behavior of forest insects) and Bill John Richardson (behavior of aquatic

insects).

The second major development was the formation of the Pestology Centre at Simon

Fraser University (SFU) in 1967. Several members of the centre focused on behavior

including, Bert Turnbull (predators), John Borden (host and mate-seeking behaviors), Peter

Belton (acoustic and oviposition behaviors in mosquitoes), Manfred Bryan Beirne

(behavior of biocontrol agents). Later additions to this group were Mark Winston (bee

behavior), Bernie Roitberg (behavioral ecology) and Gerhard Gries (pheromone-based

behaviors).

Third, the development of technology for identifying and synthesizing insect

pheromones has had tremendous impact on research programs at BC universities and

government research stations. A detailed history 1s provided below.

Fourth, the Behavioural Ecology Research Group (BERG) was established at SFU in

the mid 1980s. Several members of the BERG used principles from evolutionary biology

to study behavior of a range of organisms. These individuals include Bernie Roitberg,

114 J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

Mark Winston, Bernie Crespi (thrips), Larry Dill (aphids, phantom midge larvae and water

striders) and Ron Ydenberg (bees).

Fifth, the 1990s will be remembered for the awakening of the biodiversity

consciousness in this province. Led by Geoff Scudder and Richard Ring, the habits of

lesser known and endangered species were studied in their native habitats. Of particular

concern was the issue of habitat fragmentation and its impact on insect colonization and

perpetuation. To get a handle on these important issues requires a good understanding of

how insect behavior shapes habitat use.

Finally, behavior has featured in the research programs at government labs throughout

the province (Table 1). In many cases, there has been a conscious attempt to link behavior

to pest population dynamics. This approach is exemplified by Bryan Frazer and Neil

Gilbert’s (1976) seminal studies on the role of predator and prey behavior in the

Table 1

Government scientists and university researchers who have worked on insect behavior in

British Columbia but were not mentioned by name above.

NAME

Nello Angerilli

Brad Anholt

Rene Alfaro

Robb Bennett

Gerry Carlson

Alan Caroll

Joan Cossentine

Bob Costello

Collin Curtis

Don Elliott

Doug Finlayson

Henry Gerber

Linda Gilkeson

David Gillespie

Jack Gregson

Staffan Lindgren

Deborah Henderson

Leland Humble

HR (Mac) MacCarthy

Dave McMullen

Lorraine Maclauchlan

Vince Nealis

Imre Otvos

David Raworth

Les Safranyik

Ward Strong

Robert Traynier

Fred Wilkinson

Jerry Weintraub

Paul Wilkinson

Ian Wilson

Neville Winchester

_Bob Wright

AFFILIATION

Agriculture Canada

U Victoria

Forestry Canada

Ministry of Forests

Phero Tech Inc

Forestry Canada

Agriculture Canada

BC Government

Agriculture Canada

Private

Agriculture Canada

BC Government

Agriculture Canada

U Northern BC

Private

Forestry Canada

Agriculture Canada

Forestry Canada

Agriculture Canada

Forestry Canada

Ministry of Forests

BC Research

Agriculture Canada

Phero Tech Inc

U Victoria

BC Research

STUDY ORGANISM OR AREA

OF STUDY

Orchard pests

Aquatic insects

Forest insects

Orchard pests

Mosquitoes, greenhouse insects

Mosquitoes

Parasites, predators

Root maggots

Bees, wasps

Biocontrol

Parasitoids, greenhouse insects

Ticks

Forest insects

Parasites, predators

Forest insects

Aphids

Orchard pests

Forest insects

Insect predators

Forest insects

Mosquitoes, turf pests

Wireworms, biocontrol

Warble flies

Ticks

Forest insects

Tree canopy insects

Mosquitoes

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 115

population dynamics of pea aphids. In the next section, we discuss the role of behavior in

chemical ecology studies on insect pests.

Chemical ecology

Studies in insect chemical ecology are devoted to promoting an_ ecological

understanding of the origin, function, and significance of semiochemicals (message-

bearing chemicals) that mediate interactions within and between organisms. Such

relationships, often adaptively important, comprise the oldest and likely most prevalent

communication systems in terrestrial and aquatic environments.

Research in Chemical Ecology in British Columbia began in the 1960s when Ken

Graham (1968) at UBC explored the primary attraction of ambrosia beetles to Douglas-fir

logs undergoing anaerobic metabolism. This research culminated in 1970 with the

discovery by Henry Moeck of the Pacific Forestry Center (PFC, Victoria) that ethanol was

the compound responsible for this phenomenon. Another pioneer was John Chapman

(1966) of PFC who demonstrated in 1966 that a pheromone produced by female striped

ambrosia beetles, 7rypodendron lineatum (Olivier), was responsible for mediating

secondary attraction to and mass attack of host logs.

Since the arrival of John Borden at SFU in 1966, studies in insect chemical ecology

have been closely linked with, but not restricted to, faculty members at SFU. In 1981,

interdisciplinary research in chemical ecology was formalized by establishing the

Chemical Ecology Research Group (CERG) comprised of the entomologist John Borden,

chemists Keith Slessor and Cam Oehlschlager, apiculturist Mark Winston and plant

pathologist Jim Rahe. In 1992 Gerhard Gries joined the group. Group members have

specialized on identification and development of pheromones to manipulate economically

important insects in forestry, agriculture and stored products.

The list of achievements by CERG members is long and impressive. Particularly

significant are the identification of bark and ambrosia beetle pheromones and their

strategic deployment to alleviate the beetles’ economic impact (Borden and McLean 1981;

Borden 1990). Major accomplishments by CERG members and their students also include

the identification of the honey bee queen’s mandibular gland pheromone (Slessor ef ai.

1988), elucidation of pheromone biosyntheses (Plettner et a/. 1996) and deployment of

synthetic pheromone to manipulate the bees’ behavior within and outside the hive

(Winston and Slessor 1992). Gerhard and Regine Gries have used coupled gas

chromatographic-electroantennographic detection (GC-EAD) techniques to identify new

pheromones and kairomones for many insects, including the exotic orange wheat blossom

midge (Gries et al. 2000).

Two ‘semiochemical companies’ are spin-offs of CERG’s research activity. Phero

Tech Inc., founded by SFU graduates in 1981, launched its commercial enterprise by

offering a pheromone-based management program for ambrosia beetles. Today, Phero

Tech offers a wide variety of semiochemical products, traps and services for North

American and world-wide markets. Phero Tech Inc. also provides first employment for

many graduates of the Master of Pest Management program at SFU. ChemTica

Internacional, founded by Cam Oehlschlager and his wife Lilliana Gonzales in 1991 in

Costa Rica, started out by marketing pheromones of palm weevils and rhinoceros beetles

identified in CERG’s laboratories. Like Phero Tech Inc., ChemTica now offers diverse

semiochemical products, particularly in tropical regions.

Following the pioneering work of Chapman and Moeck, government-based scientists

have also developed considerable expertise in the chemical ecology of insect pests in

forestry and agriculture. In one of the earliest applications of GC-EAD technology (Struble

and Arn 1984), Dean Struble (Agriculture Canada, Lethbridge Research Station)

elucidated a number of lepidopteran pheromones. Roy Sheperd and Tom Gray of PFC

116 J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

(1985) developed synthetic moth pheromones as a tool to monitor population densities of

lepidopteran forest defoliators and to determine incipient outbreaks. Mike Hulme and Tom

Gray of PFC (1994) successfully used pheromone-based mating disruption to control an

infestation of the Douglas-fir tussock moth. Research by Sheila Fitzpatrick et al. (1998) of

the Pacific Agri-Food Research Centre (PARC, Agassiz) on pheromone-based control of

the blackheaded fireworm, a pest of cranberry, lead to 3M Canada’s registration of the

pheromone by the Pest Management Regulatory Agency. Gary Judd of PARC

(Summerland) developed a research program aimed at integrating semiochemicals into

orchard IPM systems and deciphering the mechanisms mediating pheromone-based insect

control (e.g. Judd et al. 1997; Evenden et al., 2000). Most recently, Bob Vernon of PARC

(Agassiz) has implemented a mass-trapping program for the European wireworm in the

Fraser Valley in an ambitious effort to save the local potato growing industry.

Current and future research by ESBC members in the fields of behavioral and chemical

ecology of insects will continue to advance our basic knowledge about insects and to

improve management of pest insects in commercial settings.

ACKNOWLEDGEMENTS

We thank Peter Belton and John Borden for sharing information and for comments on

an earlier version of this paper.

REFERENCES

Borden, J.H. 1990. Use of semiochemicals to manage coniferous tree pests in western Canada. Pp. 281-314

In: R.L. Ridgeway, R.M. Silverstein and M.N. Luscoe (Eds.), Behaviour-modifying chemicals for

insect management: application of pheromones and other attractants. Marchel Dekker, New York.

Borden, J.H. and J.A. McLean. 1981. Pheromone-based suppression of ambrosia beetles in industrial

timber processing areas. Pp. 133-154 In: E.R. Mitchell (Ed.) Management of insect pests with

semiochemicals. Plenum Press, New York.

Chapman, J.A. 1966. The effect of attack by the ambrosia beetle, Trypodendron lineatum (Olivier) on log

attractiveness. The Canadian Entomologist 98: 50-59.

Evenden, M.L., G.J.R. Judd and J.H. Borden. 2000. Investigations of mechanisms of pheromone

communication disruption of Choristoneura rosaceana (Harris) in a wind tunnel. Journal of Insect

Behavior 13: 499-510.

Fitzpatrick, S., J. Troubridge and C. Maurice. 1994. Parasitoids of blackheaded fireworm (Rhopobota

naevana Hbn.) larvae on cranberries, and larval escape behaviour. Journal of the Entomological

Society of British Columbia 91: 73-74.

Fitzpatrick, S.M., J.T. Troubridge and C. Maurice. 1998. Pheromone-mediated mating disruption as a pest

management strategy for Rhopobota naevana Hubner, the blackheaded fireworm of cranberries.

Second International Symposium on Insect Pheromones in Wageningen, The Netherlands.

Frazer, B.D. and N. Gilbert. 1976. Coccinellids and aphids: A quantitative study of the impact of adult

ladybirds (Coleoptera: Coccinellidae) preying on field populations of pea aphids (Homoptera:

Aphididae). Journal of the Entomological Society of British Columbia 73: 33-56.

Graham, K. 1968. Anaerobic induction of primary chemical attractancy for ambrosia beetles. Canadian

Journal Zoology 46: 905-908.

Gries, R., G. Gries, G. Khaskin, G.G.S. King, O. Olfert, L.-A. Kaminski, R. Lamb, I. Wise and R. Bennett.

2000. Sex pheromone of the orange wheat blossom midge. Sitodiplosis mosellana.

Naturwissenschaften 87: 450-454.

Holling, C.S. 1966. The functional response of invertebrate predators to prey density. Memoirs of the

Entomological Society. of Canada 48: 1-86.

Hulme, M. and T. Gray. 1994. Mating disruption of Douglas-fir tussock moth (Lepidoptera: Lymantriidae)

using a sprayable bead formulation of (Z)-6-heneicosen-1 1-one. Environmental Entomology 23: 1097-

1100.

Judd, G.J.R., M.G.T. Gardiner and D.R. Thompson. 1997. Control of codling moth in organically-managed

apple orchards by combining pheromone-mediated mating disruption, post-harvest fruit removal and

tree banding. Entomologica experimentalis et applicata 83:137-146.

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 A ley

Moeck, H.A. 1970. Ethanol as the primary attractant for the ambrosia beetle, 7rypodendron lineatum

(Coleoptera: Scolytidae). The Canadian Entomologist 102: 985-995.

Plettner, E., K.N. Slessor, M.L. Winston and J.E. Oliver. 1996. Caste-selective pheromone biosynthesis in

honeybees. Science 271: 1851-1853.

Shepherd, R.F., T.G. Gray, R.J. Chorney and G.E. Daterman. 1985. Pest management of Douglas-fir

tussock moth: monitoring endemic populations with pheromone traps to detect incipient outbreaks.

The Canadian Entomologist 117: 839-848.

Slessor, K.N., L.-A. Kaminski, G.G.S. King, J.H. Borden and M.L. Winston. 1988. Semiochemical basis

of the retinue response to queen honey bees. Nature 332: 354-356.

Struble, D.L. and H. Arn. 1984. Combined gas chromotography and electroantennogram recording of

insect olfactory responses. Pp. 161-178 In: H.E. Hummel and T.A. Miller (Eds.), Techniques in

pheromone research. Springer-Verlag, New York.

Stuart, L., B. But and R. Bell. 1989. Effect of host phenology on ovipositional preference of winter form

pear psylla (Homoptera: Psyllidae). Journal of the Entomological Society of British Columbia 86: 34-

38.

Vernon, R.S. and D.R. Gillespie. 1990. Responses of Frankliniella occidentalis and Trialeurodes

vaporariorum to fluorescent traps in cucumber greenhouses. Journal of the Entomological Society of

British Columbia 87: 38-41.

Winston, M.L. and K.N. Slessor. 1992. An essence of royalty: honey bee queen pheromone. American

Scientist 80: 347-385.

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 119

Arthropods that attack man and domestic animals

in British Columbia (1951 — 2001)

PETER BELTON

DEPARTMENT OF BIOLOGICAL SCIENCES, SIMON FRASER UNIVERSITY,

8888 UNIVERSITY DRIVE, BURNABY, BC, CANADA V5A 1S6

ART BORKENT

1171 MALLORY RD., ENDERBY, BC, CANADA VOE 1V3

BOB COSTELLO

BC MINISTRY OF AGRICULTURE, FOOD & FISHERIES,

1767 ANGUS CAMPBELL RD., ABBOTSFORD, BC, CANADA V3G 2M3

INTRODUCTION

In 1951, the Dominion Livestock Insect Laboratory, built in 1938 on 32 acres of

Mission Flats on the western outskirts of Kamloops, was the centre for Medical and

Veterinary Entomology in the Province (Fig. 1). J.D. Gregson was in charge and ran the

very successful tick laboratory that did much to reduce the incidence of paralysis of

livestock, and the occasional human, caused by bites of the rangeland tick, Dermacentor

andersoni. L.C. Curtis was in charge of the Household and Medical Entomology Unit until

he retired in 1969. He was heavily involved in testing new post-war insecticides and

repellents and many of our Society’s photographs show him with a portable sprayer or

fogger. G.P. Holland, who had been involved with Gregson in the biology and control of

fleas, mosquitoes, warble flies and ticks, had just left the laboratory in 1948 to head the

Systematics Unit at the Central Experimental Farm, Ottawa. He had been responsible for

identifying many thousands of fleas sent in by the Plague (Yersinia pestis) Survey carried

out during and after World War 2.

aide a

aii OO al ae

Figure 1. Left, Professor George Spencer, Ivor Ward and G. Allen Mail. Right, newly

completed Kamloops Laboratory. In the 1940s, Mail had just taken over from Spencer as

Officer-in-charge of the Lab. Ward shared an interest in grasshoppers with Spencer and

later became Provincial Entomologist until his untimely death in 1947.

120 J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

All three left milestone publications of their Federal work (Gregson 1956, Curtis 1967,

Holland 1949). Holland was partly replaced by J. Weintraub who acquired a lasting

interest in insects as a summer student at the Dominion Parasite Laboratory in Belleville.

He was appointed in 1949 and quickly became a world expert on the ecology and

physiology of warble flies. G.B. Rich also joined the Livestock Laboratory that year as a

student assistant from UBC. He worked with Weintraub on the flight range of warble flies,

and with Curtis surveying mosquitoes and controlling them at their breeding sites. He later

became well known for his work with lice, and devised some of the early trials of systemic

insecticides to control lice and warble grubs. His careful large-scale experiments showed

that cattle grubs and lice could be eradicated over as much as 200 square miles of

ranchland.

TICKS

Research on our ticks was continued by P.R. Wilkinson who studied their management,

resistance, life histories and host relationships until the Livestock Insect Section was

closed in 1971. Wilkinson then joined Weintraub who transferred to the larger Federal

Research Station at Lethbridge, AB in 1953.

Ticks are potentially the most important vectors of human diseases in the Province. In

addition to causing the paralysis that was responsible for at least 27 human deaths by the

1950s (Gregson 1956), they can transmit Powassen and Colorado tick fever viruses, Rocky

mountain and Q fever rickettsias as well as the spirochaete bacteria that cause relapsing

fever and Lyme disease. Relapsing fever was first reported as an outbreak in the

Kootenays in the 1930s (Palmer and Crawford 1933) and bites from the fast-feeding tick

Ornithodoros hermsi probably were, and continue to be, responsible (Gregson 1956).

Lyme disease, which made a sudden appearance in an epidemic at Old Lyme in the

eastern United States in 1977, has a curious history. By 1990, 11 cases had been reported

in British Columbia as well as 84 in Ontario and Manitoba (Anon.1991). The western

vector, /xodes pacificus, was a severe problem in the 1940s in West Vancouver, the

Malahat on Vancouver Island and Harrison Bay and Cultus Lake in the Fraser Valley. At

that time, however, it was not associated with disease but was known for its painful slow-

healing bites that could be exacerbated if the long mouthparts broke when the tick was

being removed. There is still some doubt whether Lyme disease is established in our

Province. People travel freely across the continent and modern and extremely sensitive

chromosomal diagnostic techniques may exaggerate its prevalence but, on the other hand,

before these techniques were used, real cases of Lyme disease may have been

misdiagnosed as arthritis from other causes.

Tick research in BC is still largely concerned with livestock and a Ministry of

Agriculture entomologist at the Kelowna office, H. Philip, in collaboration with T. Lysik

in Lethbridge is currently looking at immune responses to tick bites.

SPIDERS

In the last 50 years two types of spider bites have been reported. Western black widow

spiders (Latrodectes hesperus) are not uncommon in southern BC. The females are large,

about 2cm across the legs, and have a shiny black abdomen, with ventral red markings on

most specimens. They can inject a neurotoxic venom with their bites which occasionally

require medical treatment, but G.A. Mail, Gregson’s predecessor at the Kamloops

Laboratory found that the venom had no effect when injected into the leg muscle of a

guinea pig. Perhaps encouraged by this, Mail tested it on his own arm, which became red,

swollen and painful for about 4 days. When he heard of it, the Dominion Entomologist,

Arthur Gibson told Mail firmly to restrict his experiments to the laboratory animals

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 121

(Riegert 2000). The symptoms can usually be treated successfully with calcium injections.

The other type of spider bite seems to cause slow healing wounds. One of the earliest

medical investigations of it in the Province was reported from Kamloops (Davies 1963).

The patient was bitten on the thigh, she believed “during the night” and necrotic

arachnidism was diagnosed although the spider was not found. Such bites are often

attributed to Loxosceles reclusa, the brown recluse of the southern US, but R.G. Bennett

(2002) has recently emphasised that this species, whose bite can cause serious necrotic

injury, has not been recorded in BC (nor in Canada). Since the early 1980s there has been

an increasing number of reports of such bites being blamed on the introduced agelenid

hobo spider (Tegenaria agrestis) which can be found at various localities, and often in

homes, in southern BC. In 1999 Dr.G.Willis of the Vancouver Poison Control Centre

received 132 calls (but still less than 1% of her annual total) relating to spider bites.

Bennett notes that reports of such bites claimed to be from Tegenaria species are seldom

accompanied by the actual spider and are better explained by other factors. Binford (2001)

gives some of these explanations and shows convincingly that the proteins in the venom of

these introduced spiders are not significantly different from those found in Europe where

necrotic bites from them have never been reported. A search for necrotizing enzymes like

the sphingomyelinase D found in recluse spiders (Goddard 1999) might be worthwhile.

BITING MIDGES

A group of biting flies that were just considered a nuisance, the Ceratopogonidae

(biting midges or no-see-ums), became important in 1975 when Bluetongue, a

quarantinable virus disease of sheep and cattle, was reported for the first time in Canada

near Osoyoos. Seven species including one thought to be a major vector, Culicoides

variipennis, were known in the Province (Curtis 1941) but none had been collected in the

south Okanagan valley. Agriculture Canada organized an extensive survey led by R.D.

McMullen. They identified another 9 species of mammal-biting midges but no virus was

detected in the many adults of C. occidentalis and other potential vector species tested

(McMullen 1978). It is now certain that C. sonorensis is the major vector of Bluetongue

virus and that it was previously misidentified in BC as both C. variipennis and C.

occidentalis (Holbrook et al. 2000). A second surge of interest in these insects followed

the recognition of their association with Sweet Itch, an allergic reaction of horses to midge

bites. Fifteen cases, the first in Canada, were reported in southwestern BC (Kleider & Lees

1984) and a later survey by Anderson et al. (1988) showed that up to 26% of horses in the

Province were affected becoming unrideable and unworkable. She trapped two new midge

species bringing the total in BC to 18 (Anderson ef al. 1993). Borkent (unpublished 2001)

has now identified 31 species of these pests from the Province and suggests that another

11, found just across State and Provincial borders, may also be here.

BLACK FLIES

The start of our second half century was marked by an outbreak of another group of

biting flies, the Simuliidae or black flies, in Cherryville in the Shuswap region (Curtis

1954). In 1952 the Provincial Entomologist, C.L. Nielson, was asked to investigate their

control. He enlisted the help of L.C. Curtis who found many black fly larvae in Cherry

Creek in March and used a control program based on the one perfected by F.J.H. Fredeen

in Saskatoon (Riegert 1999). Treatment with 0.lppm DDT cleared Cherry Creek, Eight-

Mile Creek and the Shuswap River of black flies much to the satisfaction of local ranchers.

The fly, at first thought to be a new species, was later identified by Fredeen as Simulium

defoliarti, a large-mammal biting species less toxic to cattle than the prairie biter, S.

arcticum. Nevertheless it seems to have caused some cattle deaths although our Province

Loo J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

appears to be less affected than Alberta and Saskatchewan where S. arcticum can still

cause toxic and allergic reactions and fatal stampeding.

MOSQUITOES

Mosquitoes are second only to ticks as carriers of human diseases. Two viruses,

Western equine encephalomyelitis (WEE) and Snowshoe Hare (SSH), in the California

group, have or can potentially cause clinical disease in BC. Two human deaths from WEE

occurred in the Interior in 1971 and cases are reported in unvaccinated horses, presumably

brought north by virus-infected migratory birds. SSH is endemic in the north but does not

seem to cause disease in humans west of Ontario (McLean 1975). Some viruses with very

high concentrations in the host’s blood can be transmitted on the mouthparts of almost any

biting insect. Fortunately mosquitoes have never been found to transmit Human

Immunodeficiency Virus in this way but the fatal Myxoma rabbit virus is thought to be

transmitted thus by mosquitoes in the western US and is a potential threat here. Human

malaria has not been transmitted so far in BC by our indigenous Anopheles species

although one of them, An. freeborni is a capable vector in the southwestern US.

Mosquitoes can also transmit parasitic nematodes but the only one of concern in the

Province is a heartworm restricted to dogs and the odd cat, perhaps only established in the

dry interior (Slocombe 1999).

Curtis (1967) described 42 mosquito species in BC and listed five more that might be

expected. Since then, two species new to Canada, Aedes togoi and Ae. nevadensis, and two

of the species Curtis expected, Ae. melanimon and Culiseta minnesotae, have been found

in the Province bringing the total to 46 (Belton 1983). Aedes togoi, first collected in

Horseshoe Bay, is thought to have arrived by boat from Japan and is now known from rock

pools in Cortes Island south to Fidalgo Island, WA, in the south and west to Bamfield on

the Pacific coast, all of them close to harbours. In Asia, Aedes togoi is a known vector of

both Japanese B Encephalitis and filarial nematodes.

OTHER BITING PESTS

In 1955 the Provincial Entomologist, C.L. Neilson, with the help of the Federal

entomologists in Kamloops, kept agricultural workers up to date on the recommended

control practices for other biting livestock insects with a duplicated “Handbook of the

Main Economic Insects of British Columbia”. He mentions female horse and deer flies that

can consume their own weight in blood at each feeding and transmit tularemia in the

Interior of the Province. He knew of 24 species in BC, all of which bite, most of them

between June and September. Teskey (1990), who recently updated this number to 60

Species, names another seven diseases of mammals that they can transmit in North

America and pointed out that their painful bites make them likely to be dislodged while

feeding and thus more likely to transmit infections to their next host. Snipe flies

(Rhagionidae) like the horse flies to which they are related, can sometimes be troublesome

human pests in woodland, even in greater Vancouver. Neilson also deals with horn flies,

stable flies, keds, biting and sucking lice and poultry mites in his handbook and all of them

can be found as biting pests of domestic animals today.

MANAGEMENT

In the 1950s and 60s DDT and methoxychlor were thought to have a relatively low

toxicity but Neilson (1955) warned agricultural workers that DDT was eliminated very

slowly by mammals and would “build up if not used wisely.” Some of his

recommendations were for rotenone, derris and pyrethrum — “almost non-toxic for man

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 13

and animals.” Some of the techniques he suggested were designed to limit the broadcast

use of pesticides, for example setting up self-applied back-rubbers in pastures and

paddocks to control lice and flies.

Curtis (1967) recognised that the removal of larval habitat was “the most important and

efficient” method of mosquito abatement and this has been going on, consciously or not,

since the first immigrant settled in the Province. He promoted larviciding with pesticides

or by encouraging predators and parasites as the next best management technique “‘as the

larvae are confined to their native water and cannot escape”. He described adulticiding as

“the method of last resort” to be used when larval treatment was not possible or when

adults invaded from distant breeding areas. This advice can hardly be bettered although

there is valid concern nowadays for the preservation of wetlands. Curtis wrote that at that

time “the most spectacular, popular and expensive method” of controlling adults was by

aerial spraying, covering large areas in a short time. Using persistent insecticides this

technique provided immediate relief and often also a barrier to further invasion.

Unfortunately in the mid-1960s there were several incidents where pesticides were

misused. In Kamloops, for example, 45 gallon drums of DDT and 2-4-D were confused

and an airspray defoliated most of the urban trees leaving a large population of adult

mosquitoes unharmed. Adding insult to injury, many of the trees, which probably would

have recovered, were cut down and replaced. Several such incidents lead to the drafting of

new regulations for the old Pharmacy Act in 1969. Courses were taught and examinations

required for licenses to resell and to use pesticides and for certification of individuals to

dispense and apply them.

At that time a strong environmental lobby group, pioneered by SPEC, the Society for

Pollution and Environmental Control, developed in the Province and there was

considerable opposition to aerial adulticiding particularly in Coquitlam. Malathion, a

short-lived organophosphate and Baygon, a longer lasting carbamate insecticide were

registered for aerial application but in 1971 some of the residents showed their objection to

this procedure by flying balloons in the path of the spray plane.

The New Democratic Party came into power in 1972 and ordered a Royal Commission

on Pesticides and Herbicides. The commission recommended (14-III) “Aerial spray for

mosquito (sic) should only be permitted if there is a real threat to human health or

livestock. The decision to allow aerial spraying for adult mosquitoes should lie solely with

the Minister of Health Services and Hospital Insurance. Aerial distribution of chemical

larvacides should only be undertaken under permit from the proposed Pesticide Control

Branch.” It also recommended that Provincial Government leadership should be provided

“to establish rational ongoing mosquito control programmes.” At that time the Ministry of

Agriculture employed a medical entomologist, R.A. Costello, but until 1974 he was in a

Ph.D. programme at Simon Fraser University and unable to devote much time to mosquito

control. However in 1976 the Minister of Agriculture, Don Phillips, approached P. Belton,

an impartial academic with an interest in mosquitoes, to chair a Provincial Mosquito

Advisory Committee. The committee included Costello, two medical doctors, a pesticide

chemist, representatives from Environment Canada, Provincial Fish and Wildlife, a private

consultant and (a masterstroke) Mrs. M. Doucette, the chair of SPEC. For just over 10

years, and with a few changes in membership, the Committee amicably prepared and

revised a Mosquito Control Guide and generally advised the Minister of Agriculture on

policies and procedures. In 1978 the new Provincial Pesticide Control Act came into force

and by 1987 the Ministry of Environment had taken over the regulation of pesticides,

following the recommendations of the Royal Commission, and the Ministry of Agriculture

and Food no longer published the Guide. For several seasons mosquitoes had not been a

great problem, partly because of low river levels associated with slow melting snow.

Another reason was the improved management of the larvae of human-biting species, with

124 J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

better mapping of breeding sites and the intelligent use of the very effective and selective

Bacillus thuringiensis, serotype H14 larvicide. In 1988 the Mosquito Advisory Committee

quietly dissolved.

In the 21“ century, most active control of biting flies is done by larviciding. Highly

effective and specific bacterial toxins are applied to breeding sites, which can be mapped

and located using satellite techniques. Door and window screens and repellents are often

recommended as a first line of defence, but in most regions adults may be controlled on

request, often requiring unanimous groups of local residents. One of the original

controversial organophosphates, malathion, is still being used applied from the ground to

control adult mosquitoes when their numbers warrant it. Unless there is a medical

emergency, the malathion is applied as a low volume spray from trucks which can avoid

residents who object to spraying and others likely to be affected, such as beekeepers.

CONCLUSIONS

Biting insects and arachnids in BC seem to have survived about 10 millennia of

competition with humans. Our development of their habitat has undoubtedly reduced the

numbers of some species, but others, for example, that can develop in ditches, irrigation

runoff and containers, may have benefited. Dams may reduce seasonal flooding of rivers

but often change their characteristics to the advantage of some species of black flies

(Riegert 1999). Many species of ticks feed on several different hosts during their life cycle,

and changes, for example in the population of lizards, mice or deer, might affect the

numbers of ticks that could feed on humans. There is also valid concern that global

warming, possibly related to the increase in human population might increase the

population of disease-bearing species in what are now cooler parts of the planet. It might

also prolong the life of some vectors, giving them more opportunity to transmit diseases.

ACKNOWLEDGEMENTS

We thank Karen Needham for searching Spencer’s photographs and Bar and Jack Gregson

and Paul Riegert for valuable information.

REFERENCES

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British Columbia. Canadian Veterinary Journal 29: 718-723.

Anderson, G.S., P. Belton and E.M. Belton. 1993. A population study of Culicoides obsoletus Meigen

(Diptera: Ceratopogonidae) and other Culicoides species in the Fraser Valley of British Columbia. The

Canadian Entomologist 125: 439-447.

Anon. 1991. Lyme disease in Canada. Journal of the Canadian Medical Association 144 (2): 177.

Belton, P. 1983. The Mosquitoes of British Columbia. British Columbia Provincial Museum, Handbook

41, 189 pp.

Bennett, R.G. 2002. Hyberbole and hysteria along the path to enlightenment OR Show me your genitalia

and I’ll tell you what species you are OR The brown recluse (Loxosceles reclusa) and the hobo spider

(Tegenaria agrestis) in British Columbia (Araneae: Sicariidae, Agelenidae). Boreus (Newsletter of the

Entomological Society of British Columbia) 21 (2). In press

Binford, G. 2001. An analysis of geographic and intersexual chemical variation in venoms of the spider

Tegenaria agrestis (Agelenidae). Toxicon 39: 955-968.

Curtis, L.C. 1941. A preliminary list of the species of Culicoides in Western Canada. Proceedings of the

Entomological Society of British Columbia 37: 18-19.

Curtis, L.C. 1954. Observations on a blackfly pest of cattle in British Columbia, Diptera: Simuliidae.

Proceedings of the Entomological Society of British Columbia 51: 3-6.

Curtis, L.C. 1967. The Mosquitoes of British Columbia. Occasional Papers of the British Columbia

Provincial Museum No. 15, 90 pp.

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Davies, G.R. 1963. A probable case of necrotic arachnidism. Proceedings of 18" Annual Meeting of the

International Conference on Diseases in Nature Communicable to Man. Pp. 13-16.

Goddard, J. 1999. Physicians guide to Arthropods of medical importance. 3" ed. CRC Press, Boca Raton,

USA. 387 pp.

Gregson, J.D. 1956. The Ixodoidea of Canada. Canada Dept. of Agriculture Publication 930, 92 pp.

Holbrook, F.R., W.J. Tabachnik, E.T. Schmidtmann, C.N. McKinnon, R.J. Bobian and W.L. Grogan.

2000. Sympatry in the Culicoides variipennis complex (Diptera: Ceratopogonidae): a taxonomic

reassessment. Journal of Medical Entomology 37: 65-76.

Holland, G.P. 1949. The Siphonaptera of Canada. Canada Dept. of Agriculture Publication 817, 306 pp.

Kleider, N. and M.J. Lees. 1984. Culicoides hypersensitivity in the horse. 15 cases in southwestern British

Columbia. Canadian Veterinary Journal 25: 26-32.

McLean, D.M. 1975. Arboviruses and human health in Canada. Associate Committee on Scientific Criteria

for Environmental Quality, NRC Canada #14106. Pp. 1-35.

McMullen, R.D. 1978. Culicoides (Diptera: Ceratopogonidae) of the Okanagan area of British Columbia.

The Canadian Entomologist 110: 1053-1057.

Neilson, C.L. 1955. Handbook of the main economic insects of British Columbia. II Livestock insects.

British Columbia Ministry of Agriculture. 40 pp.

Palmer, J.H. and J.M. Crawford. 1933. Relapsing fever in North America with report of an outbreak in

British Columbia. Journal of the Canadian Medical Association 28: 643-647.

Riegert, P.W. 1999. A history of outbreaks of blackflies and their control in Western Canada 1886-1980.

Rampeck Publishers, Regina SK, 68 pp.

Riegert, P.W. 2000. Federal Government Research on insects of medical and veterinary importance in

Western Canada 1940-1980. Rampeck Publishers, Regina SK, 110 pp.

Slocombe, J.O.D. 1999. Heartworm in dogs in Western Canada in 1998. Proceedings of the Heartworm

Symposium °98, M. Soll Ed. Pp. 1-8.

Teskey, H.J. 1990. The Insects and Arachnids of Canada. Part 16 The horse flies and deer flies of Canada

and Alaska. Agriculture Canada Publication 1838, 381 pp.

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 127

Forensic entomology in British Columbia: A brief history

G.S. ANDERSON

SCHOOL OF CRIMINOLOGY, SIMON FRASER UNIVERSITY,

8888 UNIVERSITY DRIVE, BURNABY, BC, CANADA VSA 1S6

Forensic, or medicolegal, entomology is the study of the insects associated with a dead

body, primarily to determine time of death. In its broadest sense, forensic entomology

actually refers to any legal activity that involves insects or other arthropods. This includes

urban entomology, involving insects which affect the human environment, such as

structural damage by termites or carpenter ants and even spider bites (Haskell et al. 1997)

and stored products entomology, involving the insects and insect residue in stored products

such as grain and flour. However, it is the medicolegal or medicocriminal aspects that are

most commonly referred to by the general term ‘forensic entomology’.

Determining time of death is paramount in a death investigation. Knowing time of

death focuses the police investigation into the correct time frame, can support or refute a

suspect’s alibi, helps in the identification of an unknown victim, improves efficacy of the

police investigation and most importantly, is vital in determining time line prior to death,

victim’s whereabouts, associates seen with victim, etc. Determining time of death is,

therefore, vital. Pathologists can estimate time of death based on several medical

parameters (Henssge ef al. 1995) but these are only valid for the first few hours after death,

becoming less valuable as time passes and can usually not be used beyond about 72 h.

However, homicide victims are frequently not discovered for days, weeks or months.

Forensic entomology is the most accurate, and frequently the only method of determining

time of death when more than a day or two have elapsed (Kashyap and Pillai 1989). It

continues to be valuable up to a year or more after death. Forensic entomology also can be

used to determine whether the body has been moved from one site to another, whether the

body has been disturbed after death, the position and presence of wounds, efc., but its

primary use is to determine time of death.

A dead body, whether animal or human, is a rich but temporary and ephemeral

resource, exploited by many organisms, primarily insects. Within minutes of death,

assuming conditions are suitable, insects, primarily the Calliphoridae and Sarcophagidae,

colonize a body, developing at predictable rates, based on environmental and

meteorological conditions (Anderson and Cervenka 2001; Anderson and VanLaerhoven

1996). As the body decomposes it goes through rapid biological, chemical and physical

changes, attracting a sequence of colonizing insects until nothing of nutritional value is

left. This sequence of colonization depends on biogeoclimatic zone, habitat, season,

microclimate efc. but is predictable within those parameters (Anderson 2000a). This

predictable and sequential colonization of a body allows an entomologist to determine the

tenure of the insects on the body, and therefore the minimum time since death.

The use of insects in death investigations dates back to 13th Century China (McKnight

1981) and came into some use from the mid 19th Century in Europe (Smith 1986;

Yovanovitch 1888). This early interest led to a study on insect succession on human

corpses in Quebec, Canada, in 1897 (Johnston and Villeneuve 1897). However, little

forensic entomology was used in Canada after this time until the 1970s, when police

interest in forensic entomology began to grow in North America, with interest developing

in three major centres in North America: one of those was Vancouver, British Columbia.

In the 1960s and 1970s Peter Zuk, Vancouver Research Station, Agriculture and Agri-

Foods Canada (then Agriculture Canada) was involved in several cases for the police,

including the infamous Clifford Olsen cases. He was probably the first to present forensic

128 J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

entomological evidence in court in Canada, and certainly in British Columbia. In the early

1970s, police came to Dr. John Borden at Simon Fraser University wondering whether an

insect found on a dead railway worker might be implicated in his death. John identified the

insect as Monochamus species (Coleoptera : Cerambycidae), and stated that it could only

be the cause of death if the decedent suffered from an extreme case of entomophobia!

Forensic entomology at Simon Fraser University was born!

During the 1970s and early 1980s, Professor Thelma Finlayson, of the Centre for Pest

Management at Simon Fraser University, received preserved specimens from several cases

and was able to provide identifications and expertise. Cases were handled when a police

officer had a specific question about insects involved in an investigation, but only on a

sporadic basis, with Professor Finlayson, Dr. Borden and members of his lab providing the

expertise. Other entomologists were also sometimes called in on a case by case basis. In

the early 1980s, the BC Coroners Service approached John Borden about providing

forensic entomology expertise on a more regular basis. John enlisted the aid of Akbar

Syed, the head of the Simon Fraser University Insectary, and Akbar was involved in about

a dozen cases until 1987. He attended crime scenes and autopsies and testified in court as

an expert witness in forensic entomology (Skinner ef al. 1988).

By 1987, police interest in the field was increasing, and case work and the need for

court appearances was growing. I was just starting my Ph.D. in medical and veterinary

entomology, with Dr. Peter Belton and Dr. John Borden at Simon Fraser University in the

Centre for Pest Management. In late 1987, John Borden approached me and asked if I

would like to take over the forensic entomology, as case work was increasing and Akbar

Syed no longer wished to continue taking cases. Intrigued by the idea, I agreed. Although I

immediately delved into the literature, nothing happened until August when I was thrown

into the deep end with two cases arriving on the same weekend! One was a young man and

another a single human thigh. I attended my first two autopsies on the same day! |

determined the young man had been dead for a little over 3 weeks, based on the presence

of Calliphoridae puparia and pupae, and the thigh had been colonized 3-4 days prior to

discovery, based on Calliphoridae larval development. Both cases remain unsolved at time

of writing. Although already an entomologist and continuing in my entomology training

under John and Peter’s guidance, I had no knowledge of forensic science, crime scene

analyses, autopsies, court procedure efc. However I was fortunate to be kindly guided in

this by then Corporal Bob Stair, (now Staff Sergeant, Retired) Royal Canadian Mounted

Police (RCMP), Regional Forensic Identification Support Section (RFISS), Coroners Bart

Bastien and Chico Newell of the BC Coroners Service Forensic Unit, and pathologists Dr.

Sheila Carlyle, Dr. Laurel Grey and Dr. Rex Ferris. Everyone in the forensic field was

extremely supportive and enthusiastic and I was hooked!

Case work was increasing every year and by late 1991 I was about to complete my

graduate work. Therefore John Borden met with the Coroners office and the University

and raised the money between the two to establish a position in forensic entomology at

Simon Fraser University for me, which began in 1992. The Information and Identification

Services Directorate and the Training Directorate, Royal Canadian Mounted Police

(RCMP) later also supported this position.

It had become apparent to me during my case work that there was a desperate need for

more research in this field: there had been no research in Canada since 1897 (Johnston and

Villeneuve 1897), and none in BC. As a newly minted Ph.D. and professor I could now

devote my research to forensic entomology. Analysis of maggot development, used in the

early days and weeks after death, is based primarily on temperature, humidity and species

so literature reports, although at the time few in number, could be applied. Insect

succession over time, however, is much more dependent on local conditions and so

literature reports from other areas, seasons, habitats efc. are not applicable. Therefore, the

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 129

first research project in forensic entomology this century in Canada, and ever in BC was

begun in 1992, funded by a small start up grant from SFU. I hired a keen young second-

year year undergraduate student, Sherah VanLaerhoven and our first pig project began!

This first study looked at insect succession on carrion beginning in one season,

summer, one scenario, direct sunlight, and was conducted in the Lower Mainland of BC,

the Coastal Western Hemlock zone, using freshly killed pig carcasses. Pigs have long been

accepted as the best model for human decomposition studies (Catts and Goff 1992) due to

their similarity in skin type, gut bacteria, size, and relative lack of hair. Insect colonization

on the carcasses was studied over a 10-month period. Results were dramatic. In previous

studies in other temperate countries, with very similar climates to the Lower Mainland of

BC, such as Britain, publications had suggested that key colonizing groups, such as

Dermestidae (Coleoptera) and Piophilidae (Diptera) arrived months after death.

Dermestidae were generally considered to be the last group to feed on the remains

worldwide in temperate zones, with the majority of adults and larvae being collected in the

final stages of decomposition when only skin and bones remain (Early and Goff 1986;

Smith 1986; Rodriguez and Bass 1983; Payne and King 1970; Easton 1966; Reed 1958;

Fuller 1934), although the actual post mortem interval varied with geographic region.

Early colonization was only reported from much more tropical regions, such as Hawaii

(Hewadikaram and Goff 1991; Early and Goff 1986). The only other report of insect

succession on carrion in Canada states that Dermestidae, including Dermestes frischii

Kugelman, were not collected on human remains until 3-6 months after death, despite

compelling evidence of a case in which Dermestes sp. were found less than 5 weeks after

death (Johnston and Villeneuve 1897). However, in the Lower Mainland of BC,

Dermestidae larvae were first collected 21 days after death, and were commonly collected

after 43 days post mortem (Anderson and VanLaerhoven 1996).

Piophilidae or skipper flies were similarly collected earlier than previously reported,

with larvae being present 29 days after death. This confirmed case work in which I had

collected Piophilidae from human remains 26 days after death (Anderson 1995). This

initial study (Anderson and VanLaerhoven 1996) highlighted the need to conduct research

in all biogeoclimatic zones, seasons and habitats in which forensic entomology is used.

Such data are not necessarily transposable to other regions.

When large numbers of maggots congregate on a body, they form masses which

generate high temperatures, affecting development rates. It had been previously assumed

that although diurnal temperatures obviously fluctuated, internal carcass temperature

fluctuated less, remaining at a higher, more constant temperature than ambient (Deonier

1940). Previous studies had measured maggot mass temperature daily but only at a single

time each day (Shean ef al. 1993; Early and Goff 1986; Payne 1965). In our experiments,

we placed datalogger probes inside the carcasses, continuously recording internal carcass

temperature, and found that although internal carcass temperature does increase

considerably during active decay, there is greater fluctuation in internal than ambient

temperature, with diel differences of more than 35°C (Anderson and VanLaerhoven 1996).

This study became the foundation for our future work. Research was needed in

different geographical areas and situations as I was training police officers, in BC and

across North America (Anderson ef a/. 1996; Anderson 1993a, b, 1991), so case work was

increasing rapidly (Anderson 1995). I was also testifying in court as an expert witness, or

my report was admitted as evidence in more and more cases.

Funding from the Canadian Police Research Centre (CPRC) allowed me to take on my

first M.Sc. student, Leigh Dillon, and to expand the research into three biogeoclimatic

zones, the Coastal Western Hemlock zone, the Sub-boreal Spruce Zone and the Interior

Douglas Fir Zone. Although BC is divided into 14 biogeoclimatic zones, these three zones

include the most populous areas, such as the majority of the Lower Mainland, Vancouver

130 J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

Training police officers to collect insect evidence in a mock homicide scene.

Island, Prince George Region and the Okanagan, so consequently are the areas from which

most forensic cases originate. This work again used pig carcasses as human models and

insect succession was studied in the three zones, in spring, summer, and fall, in sun and

shade.

Colonization times varied with season, habitat and biogeoclimatic zone, and occurred

in a predictable sequence with similar species colonizing in each area, although

colonization times varied (Dillon 1997; Dillon and Anderson 1996a, 1995). Some species

were found in all areas, but in some areas certain species predominated. A distinct

difference was noted between pigs in sun and shade, in species composition and

abundance, as well as seasonal differences in both. Level of shade impacted decomposition

rates, actual species attracted and arrival times. Pigs in shade were also scavenged more

heavily by vertebrates, which impacted insect colonization (Dillon 1997; Dillon and

Anderson 1996a, 1995).

This work generated excellent databases of insect succession on carrion for these

biogeoclimatic zones and also confirmed our previous observations of earlier colonization

times for many species in BC in comparison with some other regions, and the diurnal

temperature fluctuations within the maggot mass. However, this work also showed that in

some cases, primarily those in shade, or cooler seasons, the oldest maggots would enter the

prepupal stage and leave the body before the high temperatures were generated by the

maggot mass (Dillon 1997; Dillon and Anderson 1996b). This obviously has a major

impact on determining time of death using maggot development. Maggot mass

temperature is usually taken into account when determining age of the oldest maggots, but

this work indicates that in some cases, the oldest maggots may not be impacted by the

maggot mass.

However, early on in the research, although the pigs in spring and fall, and those in the

shade in summer decomposed in a manner similar to the human cases I was analyzing at

the time, we noticed that Leigh’s summer, sun pigs decomposed much faster than a human

body, with the skin and extremities actually mummifying (Dillon 1997; Dillon and

Anderson 1995). We realized that, while the pigs that Leigh was working with were naked,

the majority of the human cases I was involved with were clothed. We also noticed that the

majority of my previous cases involved either clothed, partially clothed or wrapped bodies.

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 131

We postulated, therefore, that clothing might be having an effect on pig decomposition and

insect colonization, as clothing absorbs body fluids, and provides habitats for insects. |

sent a simple little electronic message to my departmental colleagues requesting donations

of used clothing. The message somehow reached the media; to Associated Press and then

went international! I received large quantities of used clothing from all over and we were

able to clad our pigs! For years, I continued to receive disreputable packages of old

clothing! However, some good did come of all this as although my work had received a

tremendous amount of media attention prior to this, which continues to this day, it was this

particular attention that was noticed by an entomology student in Ontario, Niki Hobischak,

who became my third graduate student.

In 1995, a wildlife enforcement officer noticed this work and contacted me, asking

whether forensic entomology could be applied to poaching cases. Bear poaching is a major

crime in BC with animals being taken illegally as trophies and for body parts, primarily for

the Traditional Chinese Medicine market. Most carrion insects are ubiquitous and do not

discriminate between carrion species, except in the case of particularly small carcasses

(Denno and Cothran 1975), and most species are more likely to colonize animal carcasses

than human bodies, simply due to availability. However, although pigs are accepted as

good models for human decomposition, it was not known whether they would be valid as

models for bear decomposition. Therefore, supported by various wildlife organizations,

including World Wildlife Fund and the Vancouver Foundation, we added several bear

carcasses to Leigh’s project. These were bears that had been killed as nuisance bears,

which are normally incinerated. In 1995, they were donated to forensic entomology

research! Obviously we could not get enough bears at the same time for a full experiment,

but Leigh was able to compare bear decomposition and insect colonization with her

extensive pig experiments (Dillon 1997; Dillon and Anderson 1997, 1996b). Leigh Dillon

graduated in 1997 and is presently a coroner with the BC Coroners service.

In 1995, I was called into my first bear poaching case, in which two cubs had been

killed for their gall bladders. The case involved first instar Calliphoridae eclosion rates and

I testified in the court case in 1996 (Anderson 1999). The insect evidence indicated a time

of death that successfully linked two suspects to the scene. Both defendants were found

guilty of two counts of poaching under the Provincial Wildlife Act of Manitoba. They

were sentenced to 3 months in jail per count. This was the first time in Canada where a jail

term was secured for the actual poachers of the animals in question and has set a

precedent.

By this time, the compilation of an extensive database was under way in BC for bodies

found above ground, but many bodies are buried and there was little research worldwide

on buried carrion, and no research for BC. This point was brought home when the body of

a child was found in a shallow grave nearly two months after the child was last seen alive.

I collected the insects from the grave and autopsy but had no direct database with which to

compare. Sherah VanLaerhoven, my first forensic entomology research assistant, was

completing her undergraduate degree with a small research project in my _ lab

(VanLaerhoven and Anderson 2001) and decided that she would like to do her Master of

Pest Management on insect colonization of buried bodies. Again funded by the CPRC, and

with support from the RCMP, large numbers of clothed pig carcasses were buried in

shallow graves in two biogeoclimatic zones, with above ground carcasses as controls.

Three pigs were exhumed each time and their fauna analyzed at 2 and 6 weeks, 3, 6, 12

and 16 months. This work showed that burial encouraged some species and discouraged

others. In particular, species in the family Calliphoridae, although present, were much rarer

on buried bodies (VanLaerhoven and Anderson 1999; VanLaerhoven 1997; VanLaerhoven

and Anderson 1996), than on above ground carcasses (Dillon 1997) and Muscidae were

much more common, possibly due to the lack of competition with Calliphoridae. In some

132 J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

cases, the same species were present above and below ground, but at very different times.

For instance, Fannia species (Diptera: Fanniidae) were found 6 weeks or more after death

in above ground bodies (Dillon 1997), but were found within 2 weeks of death on buried

bodies (VanLaerhoven and Anderson 1999; VanLaerhoven 1997; VanLaerhoven and

Anderson 1996). Decomposition was also greatly slowed by even such shallow burial. As

Calliphoridae were low in numbers, no maggot masses formed so carcass temperature

remained very close to soil temperatures (VanLaerhoven and Anderson 1999;

VanLaerhoven 1997; VanLaerhoven and Anderson 1996). This is quite different from data

from Tennessee in human bodies (Rodriguez and Bass 1985), and also from our

preliminary research in Alberta in which maggot masses did form, indicating major

geographic differences. This work indicated that it is difficult if not impossible to

extrapolate data from above ground research to buried victims. I used these data when |

testified in both the preliminary and supreme court trials associated with the death that had

prompted this research in the first place. Sherah went on to complete a Ph.D. in

entomology at the University of Arkansas and has now returned to BC as an NSERC

postdoctoral fellow at Agriculture and Agri-foods Research Station, Agassiz and Simon

Fraser University with Dr. Dave Gillespie and Dr. Bernie Roitberg. This is coming full

circle, as Sherah first became intrigued with entomology in Dave’s lab as a volunteer at the

research station when she was just 17!

We had developed databases for terrestrial environments, both above ground and

buried. However, whenever I gave seminars to police officers, one of the first questions

asked always concerned what happens to a body in water? So, in January 1996, Niki

Hobischak began an MPM degree on the effects of freshwater submergence on a body,

funded by CPRC. Pig carcasses were again used to model human decomposition and were

compared with human deaths in similar habitats. Niki looked at decomposition and faunal

colonization on carcasses in freshwater streams and ponds and found that variation in

aquatic organisms occurred over time, based primarily on season but also on

decompositional stage, with Trichoptera being the major scavengers. Calliphoridae also

colonized when parts of the body were exposed, but were rarely successful (Hobischak

and Anderson 2002; Hobischak 1997; MacDonell and Anderson 1997). The pig carcass

research compared well with human death cases with known elapsed time since death in

the same time period (Hobischak and Anderson 1999). Niki was a Coroner after

completing her MPM in 1997, and is now the Research Coordinator for the Forensic

Entomology Lab at Simon Fraser University.

In 1997, I moved to the School of Criminology at Simon Fraser University, and in

1999 received funding from the Ministry of the Attorney General of British Columbia,

Proceeds of Crime Fund and the University, to build the first laboratory in Canada

dedicated to forensic entomology. Through the lab, I am now coordinating research across

Canada to develop databases of insect succession on carrion in several biogeoclimatic

zones. This work involves extensive collaboration with entomologists, forensic scientists

and students in several provinces.

Although by then data existed for the effects of freshwater decomposition on a body,

no data were available for bodies in the marine environment. In talking with Cpl. Bob

Teather, RCMP (retired) a distinguished police diver, I mentioned my desire to look at the

effects of marine submergence on faunal colonization and decomposition rates of a body.

However, this project was limited by a lack of resources such as boats, divers efc. Bob said

“well, we have divers, we have boats!”. So the marine project was born. Funding was

provided again by the CPRC, but massive in kind support, including divers, boats,

hovercrafts, field research facilities, safety equipment efc. was provided by the RCMP,

Canadian Coast Guard, Canadian Amphibious Search Team and the Vancouver Aquarium

Marine Sciences Centre. Pig carcasses were submerged in the waters off Vancouver

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 133

shortly after death in both early summer and fall, and divers observed, photographed and

sampled the carcasses every few days. In general Niki and I found that decomposition was

much slower than in terrestrial environments, although the carcasses went through much

the same stages. The tissue itself remained intact for much longer than on land, and

decomposition was affected by whether the body was in contact with the sediment because

the diversity of animals were limited when the body floated. A sequence of marine fauna

colonized the remains, focusing at first on wounds but very shortly after, on non-wound

areas, unlike terrestrial colonization where wounds and orifices are the focus (Dillon 1997;

Anderson and VanLaerhoven 1996; Dillon and Anderson 1996a, 1995). Fauna included

crustacea, mollusks, annelids and echinoderms (Anderson and Hobischak 2001). This

work is ongoing.

Forensic entomology is now an established part of death investigations in Canada, as

well as providing greatly needed knowledge on carrion ecosystems, an area which has

often been sadly neglected in the past. It has extended into studying ancient human

remains and a future collaboration will involve looking at Calliphoridae myiasis in live

people for maggot debridement therapy for wounds. Present research includes extending

our aquatic work into large bodies of freshwater and white water, the effects of commonly

used human drugs, illicit and therapeutic, on insect development and the effects of several

microclimatic features on insect development. Of course, the lab continues to receive

many cases every year from BC and across Canada as well as from other countries. Each

case is unique and often provokes more questions which will be addressed through the

Forensic Entomology Lab at Simon Fraser University.

One of the most satisfying things about doing research in forensic entomology, for

myself and my students is the knowledge that the data we generate can and will be used in

a death investigation and a court of law, often very soon after it has been generated. In a

case in Manitoba, I was testifying in the second degree murder trial of a man accused of

murdering a teenage girl. Empty pupal cases of Phormia regina (Meigen) (Diptera :

Calliphoridae) indicated that the insects had completed an entire life cycle on the body. and

I based my determination of a minimum elapsed time since death of 30 days (or 480.5

accumulated degree-days) on my lab generated data for this species (Anderson and

Cervenka 2001; Anderson 2000b). During the preliminary trial, defense counsel argued

that the data were lab generated and therefore could not be applied to a field situation.

Although it was true that the data were lab generated, Calliphoridae larval development is

primarily temperature driven, so lab conditions can simulate the main parameters involved.

However, it was true that I did not have field data to back up my lab data. Just weeks

before the supreme court trial, by sheer coincidence, Leigh Dillon’s field work had been

conducted in almost the exact same weather conditions as those in the case. She noted that

it took the first specimens of Phormia regina exactly 30 days to complete development

under those temperature conditions (Dillon 1997; Dillon and Anderson 1996a), confirming

the lab data exactly. The defendant was convicted.

ACKNOWLEDGEMENTS

I would like to thank Professor Thelma Finlayson, Dr. John Borden and Mr. Akbar

Syed for pioneering forensic entomology in BC, and particularly, Dr. John Borden, for

getting me started in this exciting and unusual field, and for his continued encouragement.

I would also like to thank my past, present and future graduate students for expanding and

extending this field. | am also extremely grateful to the many financial supporters of this

field, including the Canadian Police Research Centre, BC Ministry of Attorney General,

the Information and Identification Services Directorate and the Training Directorate, Royal

Canadian Mounted Police, Vancouver Foundation, World Wildlife Fund, BC Coroners

Service and Simon Fraser University.

134 J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

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11.

Anderson, G.S. 1993b. Forensic entomology. Insects and crime scenes. Identification Canada

Oct/Nov/Dec. : 10-11.

Anderson, G.S. 1995. The use of insects in death investigations: an analysis of forensic entomology cases

in British Columbia over a five year period. Canadian Society of Forensic Science Journal 28(4): 277-

292,

Anderson, G.S. 1999. Wildlife forensic entomology: determining time of death in two illegally killed black

bear cubs, a case report. Journal of Forensic Sciences 44(4): 856-859.

Anderson, G.S. 2000a. Insect succession on carrion and its relationship to determining time of death.

Pp.143-175 In: J.H. Byrd and J.L. Castner (Eds.) Forensic Entomology: The utility of arthropods in

legal investigations Boca Raton, CRC Press. 440pp.

Anderson, G.S. 2000b. Minimum and maximum developmental rates of some forensically significant

Calliphoridae (Diptera). Journal of Forensic Sciences 45(4): 824-832.

Anderson, G.S. and V.J. Cervenka. 2001. Insects associated with the body: their use and analyses. Pp. 173-

200 In: W.D. Haglund and M. Sorg (Eds.) Forensic Taphonomy, The Postmortem Fate of Human

Remains. Boca Raton, CRC Press. 544pp.

Anderson, G.S. and N.R. Hobischak. 2001. Effects of marine submergence on pig decomposition and

faunal colonization in British Columbia. Proceedings of the American Academy of Forensic Science

Annual Meeting, Seattle, WA.

Anderson, G.S. and S.L. VanLaerhoven. 1996. Initial studies on insect succession on carrion in

southwestern British Columbia. Journal of Forensic Sciences 41(4): 617-625.

Anderson, G.S., J.J. Gaudet and B.A. Yamashita. 1996. A practical exercise in Forensic Entomology.

RCMP Gazette 58(9): 3-6.

Catts, E.P. and M.L. Goff. 1992. Forensic entomology in criminal investigations. Annual Review of

Entomology 37: 253-272.

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Entomological Society of America 68: 741-754.

Deonier, C.C. 1940. Carcass temperatures and their relation to winter blowfly activity in the Southwest.

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Dillon, L.C. 1997. Insect succession on carrion in three biogeoclimatic zones in British Columbia. M.Sc.

Thesis. Dept. of Biological Sciences, Simon Fraser University, Burnaby. BC: 76 pp.

Dillon, L.C. and G.S. Anderson. 1995. Forensic entomology: The use of insects in death investigations to

determine elapsed time since death. Ottawa, ON. Canadian Police Research Centre. TR-05-95.

Dillon, L.C. and G.S. Anderson. 1996a. Forensic entomology: A database for insect succession on carrion

in Northern and Interior BC. Canadian Police Research Centre. TR-04-96

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Dillon, L.C. and G.S. Anderson. 1997. Forensic Entomology -- Use of Insects towards illegally killed

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VanLaerhoven, S.L. and G.S. Anderson. 1999. Insect succession on buried carrion in two biogeoclimatic

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VanLaerhoven, S.L. and G.S. Anderson. 2001. Implications of using development rates of blow fly

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J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 137

Bees and pollination in British Columbia

PAUL VAN WESTENDORP

BC MINISTRY OF AGRICULTURE, FOOD & FISHERIES,

1767 ANGUS CAMPBELL ROAD, ABBOTSFORD, BC, CANADA V3G 2M3

DOUG M. MCCUTCHEON

BEE BOOKS & THINGS,

2525 PHILLIPS STREET, ARMSTRONG, BC, CANADA VOE I1B1

Bees co-evolved with flowering plants. Their wasp-like ancestors modified their diet

and behavior in order to utilize the floral food resources that became available some 90

million years ago. In turn, plants were able to accelerate species diversification. The inter-

dependency between flowering plants, especiaily fruit-bearing plants and insect pollinators

became obligatory and in some cases developed into extreme specialization. In general, the

abundance and species diversity of insect pollinators play a critical role in the ecology of

many habitats in the world. Economic interests and demand for optimized yields in

modern crop production systems have intensified the interdependency between crops and

insect pollinators.

British Columbia’s temperate environment and habitat diversity has facilitated the

evolution of a rich pollinator fauna. The majority of its indigenous pollinators are solitary

bees, characterized by the female establishing a nest on her own, and provisioning each

cell with pollen, nectar and an egg. Many solitary bee species are gregarious and nest in

the soil.

In comparison, only a small number of social bee species are indigenous to BC. These

include bumblebees (Bombus spp.) characterized by one female (queen) having assumed

sole responsibility of egg laying while all other individuals are sterile workers and males

(drones). The vast majority of the offspring are workers responsible for food gathering,

nest building, brood rearing and protection.

Winston and Graf (1982) identified six families of solitary bees in BC. These included

Halictus, Andrena, Augochlora, Chelastoma, Melissodes and Xylocopa. In 1987,

Scott-Dupree and Winston (1987) examined the diversity of bees in Okanagan Valley

orchards and found Halictus spp. most abundant, followed by Andrena, Megachilidae and

Anthophoridae. Among bumblebees, Bombus terricola was most often recorded followed

by B. bifarius, while at least ten additional species were noted. Mackenzie and Winston

(1984) found B. mixtus the most abundant species in Fraser Valley berry crops followed by

B. occidentalis, B. terricola and B. flavifrons.

Management of Bee Species for Crop Pollination

Although all bees play an important role in the ecology of most habitats, the

development of agriculture in the province magnified the significance of their role

(Matheson ef a/. 1996). John Corner, BC Provincial Apiarist from 1950 to 1983, assessed

the suitability of using wild bees to improve pollination of various crops (Corner 1963).

Corner selected the alkali bee Nomia melanderi for further trials. These bees were

gregarious ground nesters from Oregon and were used in alfalfa pollination on a limited

scale. Soil beds were established adjacent to alfalfa fields near Ashcroft and Kamloops and

soil cores containing larvae were imported and successfully introduced. The bees became

well established but the project was abandoned, as alfalfa seed production remained

limited in the southern interior. Management suitability of Bombus spp. was also assessed

138 J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

in the Peace region for the pollination of red clover Trifolium pratense. The long-tongued

species of B. californicus and B. auricomis were identified as most effective pollinators

while B. bifarius nearcticus was deemed unsuitable.

Other trials involved the alfalfa leafcutter bee Megachile rotundata at sites in Creston,

Vernon, Kamloops, Williams Lake and Peace River. Although leafcutter bee management

proved successful under BC conditions, the alfalfa growers' unfamiliarity with seed

production caused the abandonment of the projects in southern and central BC. Leaf cutter

management proved a successful enterprise for some Peace operations specialized in

alfalfa seed production.

Scott-Dupree and Winston (1987) noted a low count of the indigenous Blue Orchard

Mason Bee Osmia lignaria in Okanagan orchards. This solitary and gregarious species had

been previously identified for its manageability and excellent pollination characteristics

under poor weather conditions. Scott-Dupree recommended this species be considered for

management in BC’s tree fruit and berry crops. In the same study, B. occidentalis and B.

nearcticus were also suggested as candidates for commercial use. Since then, O. lignaria

was never adopted as a significant pollinator in commercial crops, but it has gained

considerable popularity among gardeners in urban settings in recent years.

Researchers have assessed the management and rearing suitability of a wide range of

insect pollinators over many years. Pollinators with unique characteristics may still be

selected for special crop pollination requirements in the future. Yet, of more than 4,000

insect pollinators identified in North America, only a few species have ever been managed

in significant numbers for crop pollination purposes. These include the non-indigenous

honeybee Apis mellifera L., two species of bumblebees (B. occidentalis and B. impatiens),

the alfalfa leafcutter bee M. rotundata, and the Orchard Mason Bee O. lignaria.

Orchard Mason Bees

The Orchard Mason Bee (also called the Blue Orchard bee, Mason Bee or Osmia Bee)

is the ideal ‘urban bee’. During the 1990s, O. lignaria propingua became popular in urban

garden settings because of its non-defensive behavior, low maintenance, and high

pollinating efficiency in early blooming fruit bearing plants. Its popularization was further

enhanced with the introduction of ‘condominiums’ or nest boxes that are now

commercially available at garden centers and selected nurseries. Many initial enthusiasts

were former beekeepers who no longer kept honeybee colonies following the introduction

of the obligate parasitic mite Varroa destructor (formerly V. jacobsoni Oudemans).

The Osmia Bee has nesting behavior similar to M. rotundata where the gregarious

female selects a tubular cavity where she lays up to 10 eggs in succession, each

provisioned with nectar and pollen, and closed off with a plug. Since reproduction rates

remain limited, this pollinator is not considered suitable as a primary crop pollinator in

large-scale settings.

Bumblebees

Over 30 indigenous species of bumblebees (affectionately called ‘Bumbles’) have been

identified in BC. The bumblebee tolerance to poor weather conditions has made this insect

an ideal pollinator of early blooming plants. The characteristic ‘buzzing’ causes sticky,

moist pollen grains to be dislodged, further enhancing the bumblebee’s pollinating

versatility. While honeybees have a complex communication system enabling them to

utilize pollen and nectar sources over great distances, the bumblebee’s solitary, non-

directional food gathering limits its foraging range and makes it an ideal pollinator in the

confined space of the greenhouse. Their smaller nests and comparatively low defensive

behavior also reduces conflict with greenhouse workers. Bumblebees have become the

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 139

principal pollinators in greenhouse tomato production providing higher crop yields,

improved quality and earlier maturation at substantially lower costs than manual

pollination. In BC, growers purchase nest boxes from eastern Canadian suppliers that

include the indigenous Bombus occidentalis and the non-indigenous Bombus impatiens.

Honeybees

Honeybees are not indigenous to the Americas. They were first brought to North

America from Europe in the 1600s. The first introduction of honeybees in BC was in the

1850s. Initially, these colonies were not prolific as most of the natural vegetation of the

province did not offer sufficient nectar and pollen sources. As agricultural activities

expanded in southwestern BC and the Okanagan, honeybees became well established.

The introduction of intensive agricultural practices in modern times has made the

honeybee indispensable. Monocultural practices, pesticides, and the alteration of the soil

and natural vegetation have contributed to the decline in the abundance and species

diversity of wild insect pollinators (Matheson et a/. 1996). Even with a natural abundance

of pollinators, the pollination requirements of monocultural crop management systems can

not be met. It has been estimated that an hectare of mature highbush blueberries produce 9

— 10 million flowers in spring when weather conditions are often not conducive to insect

foraging activity. Only honeybee colonies with their large populations can meet crop

pollination requirements at the time of bloom. The value of annual crop production in BC

attributable to honeybee pollination has been estimated at over $161 million (Anon. 1999).

For Canada, this value is estimated at approximately $700 million (Scott-Dupree 1995). In

the USA with its milder climates, honeybee pollination has been valued at over $14 billion

worth of agricultural production per year (Morse and Calderone 2000).

The first reference to using honeybees in pollinating crops for a fee was in the 1953

Annual Report of the BC Department of Agriculture. A total of 155 colonies were rented

for tree fruit pollination at $2.50 - $6 per colony. By 1975, 5000 hive sets were recorded at

an average pollination fee of $15 per colony. During the 1990s, 29,000 hive sets were

rented each year with fees ranging from $50 in tree fruit orchards to $90 in cranberry

Vaccinium macrocarpon. With the ongoing expansion of berry crops in the Fraser Valley

and high-density plantings of tree fruits in the Okanagan, the demand for honeybee

pollinating units has increased to 47,000 in 2001.

Honeybee Diseases and Pests

The perennial colony nest of honeybees offers ideal conditions to a range of pathogenic

and non-pathogenic organisms. In 1586, Jacob of Germany first described American

Foulbrood disease (AFB) caused by the spore-forming bacterium Paenibacillus larvae

(formerly Bacillus larvae L.) (Otten 1999). The highly resistant spores remain viable for

decades and pose a source of infection to bee brood in any infected hive equipment. Many

countries in the world have enacted legislation to control the disease, which has

traditionally involved the depopulation of colonies followed by burning of all the

equipment. In the 1950s, antibiotics were introduced enabling beekeepers to control AFB

effectively. However, the bacterial spores would not be killed but merely prevented of

germination. The incessant use of antibiotics in beekeeping management has led to the

development of antibiotic-resistant strains of P. /arvae (r-AFB) in recent years.

Other bee brood diseases include European Foulbrood (Bacillus alvei), Chalkbrood

(Ascosphaera apis) and viral diseases such as Sacbrood. Most of these ailments are stress

related and can be managed relatively easily by the beekeeper. Nosema disease caused by

the protozoan Nosema apis affects the midgut and ventriculus of adult bees, causing

140 J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

impairment in nutrient absorption. Effective control is obtained with the application of the

antibiotic fumagillin.

With the arrival of parasitic mites, beekeeping changed radically. In 1986, the first

infestation of the microscopic mite Acarapis woodi was confirmed in BC. Initially, the

impact of this mite, affecting the tracheal tubes of adult bees, was much feared because of

the widespread destruction of colonies that had been reported on the Isle of Wight in 1919.

Since then, the tracheal mite has proven manageable for most beekeepers. Comparative

studies on tracheal mite resistance in honeybees during the 1990s showed that resistant

strains occurred in BC. Through selection, many beekeepers developed beestock with

some level of tracheal mite resistance.

The first infestation of the highly destructive mite Varroa destructor was confirmed in

1990. Despite efforts to isolate the pest through colony movement restrictions, the mite

eventually spread to most beekeeping areas in BC. V. destructor originated in Southeast

Asia where it was a common pest of the eastern honeybee, Apis cerana. With the

introduction of the western or European honeybee, the mite found a perfect host without

defenses parasitizing brood and adult bees.

The high pathogenicity of V. destructor invariably leads to the demise of the colony if

no controls are applied. In Canada, formic acid and the synthetic pyrethroid fluvalinate

marketed under the trade name Apistan, have been registered to control this pest. The

development of Apistan-resistant mites signals the end of the usefulness of this product.

There are currently efforts to obtain an emergency registration of Coumaphos of Bayer,

while other control methodologies are being sought at various research facilities in North

America.

Use of Hive Products

Honey is the end product bees produce from the collection of floral nectar sources.

While nectar is a sugary solution containing approximately 80% water, honey is a solution

of enzymatically converted monosaccharides containing between 14 and 20% water. Its

low water content prevents microbial growth and when kept airtight and in cool

conditions, honey can be stored for many years. Virtually all stored honeys will undergo

the physical process of crystallization over time. The rate or crystallization is determined

by the relative abundance of the different sugars. Reversal to liquid honey can be

accomplished through warming and stirring. In North America, honey is viewed as a fancy

alternative sweetener to table sugar. In most other countries of the world honey is regarded

as a scarce and valuable product to which many medicinal qualities are ascribed. While per

capita consumption of honey in Canada is only 0.86 kg per annum, consumption in the

Middle East exceeds 10 kg per capita per annum (Anon. 2001).

Pollen constitutes the primary protein source needed for brood rearing. Through special

management, some beekeepers collect pollen for feeding to bees at a future date, for sale to

other beekeepers, or for human consumption. Some of the pollen is also collected as a

nutrient supplement for racehorses. The total amount of pollen sold for commercial

purposes is limited.

Propolis (Pro — for; polis — the community) is a resinous material collected from floral

and foliar buds. The material is used for plugging holes in the nest cavity, or encasing and

mummifying foreign materials inside the nest that bees can’t remove. Its strong anti-

microbial and hydrophobic properties have long been recognized, from the ancient

Egyptians to today’s pharmaceutical industry.

Bee venom has long been used for controlling rheumatoid arthritis and various other

ailments. A collection device has been developed where bees release their venom when

exposed to a small electric charge. The pharmaceutical industry is the primary market

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 14]

although in recent years, the alternative medicinal practice of apitherapy has gained

popularity.

Royal jelly is a protein-rich excretion of the hypo-pharyngeal glands in young worker

bees. The material is the principal food source of the queen throughout her life. Worker

brood is only fed small quantities of royal jelly mixed with honey and pollen during early

larval development. High labor costs in colony management and harvesting have

prevented commercial production of royal jelly in North America and Europe. The world’s

largest producers include China and Korea.

Queen and Honeybee Stock Production

Prolonged winter conditions have always been among the most important stress factors

to honeybee colonies. Average winter mortality has been about 16% for the province but in

some northern regions average losses have often been much higher. To replace winter

losses and improve the quality of stock, beekeepers purchase queens or package bees (a

cage containing approximately 8,000 bees and a queen) from breeders. In former times,

large-scale commercial operators in the Peace region purchased thousands of packages

from California each spring to stock their hives. These “package operators” killed off all

their colonies after the honey harvest, as wintering was considered too costly and

expensive. When parasitic mites were first discovered in the US, Canada closed its borders

to the import of bees from the US, forcing Canadian beekeepers to winter their bees and

rely on domestically produced beestock or bees imported from Australia and New Zealand.

Most of the domestic bee breeders became established in coastal BC. Due to high

production costs and late availability in spring, the growth potential of this sector remains

limited.

REFERENCES

Anon. 1999. Factsheet #504. Apiculture Program, BC Ministry of Agriculture and Food.

Anon. 2001. World honey market. American Bee Journal 141(12): 859.

Corner, J. 1963. Annual Report, Apiculture Program, BC Department of Agriculture, Internal Document,

Unpublished.

Matheson, A., S.L. Buchmann, C. O’Toole, P. Westrich and I.H. Williams. 1996. The Conservation of

Bees. Linnean Society of London and the International Bee Research Association. Academic Press,

London.

McKenzie, K.E. and M.L. Winston. 1984. Diversity and abundance of native bee pollinators in berry crops

and natural vegetation in the lower Fraser Valley, British Columbia. The Canadian Entomologist 116:

965-974.

Morse, R.A. and N.W. Calderone. 2000. The value of honey bees as pollinators of US crops in 2000. Bee

Culture, http://bee.airoot.com/beeculture/pollination2000/pg1.html

Otten, C. 1999. Epidemiology of the American Foulbrood in Germany. Proceedings Apimondia °99

Congress, Vancouver, BC. Pp. 52-53.

Scott-Dupree, C. (Ed.). 1995. A Guide to Managing Bees for Crop Pollination. Canadian Association of

Professional Apiculturists.

Scott-Dupree, C.D. and M.L. Winston. 1987. Wild bee pollinator diversity and abundance in orchard and

uncultured habitats in the Okanagan Valley, British Columbia. The Canadian Entomologist 119: 735-

745.

Winston, M.L. and L.H. Graf. 1982. Native pollinators of berry crops in the Fraser Valley of British

Columbia. Journal of the Entomological Society of British Columbia 79: 14-20.

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J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 143

Fifty years of entomological research in orchard and

vegetable crops in British Columbia

R.S. VERNON

AGRICULTURE AND AGRI-FOOD CANADA,

PACIFIC AGRI-FOOD RESEARCH CENTRE, AGASSIZ, BC, CANADA VOM 1A0

British Columbia has seen tremendous advances and accomplishments in the broad field of

entomology over the past fifty years. This is especially true in relation to the orchard and

vegetable industries, where entomologists have played essential and often pivotal roles in the

production, protection and sustainability of these crops. I have attempted to list as many of the

entomologists (and their affiliations) working in orchards and field vegetables as I have been

able to retrieve from memory and various archival sources. Unfortunately, to adequately

summarize the specific endeavors of the many scientists involved would take more space than

allotted for this article. For more in-depth information on the entomologists of BC and their

various research specialties, the reader is referred to the joint Entomological Society of BC

and Entomological Society of Canada publication ‘Entomologists of British Columbia’

compiled by P.W. Riegert in 1991. For this paper, I have chosen in part to present a number of

the more highly publicized research programs and the entomologists involved that have played

pivotal roles in the direction and advancement of entomology in certain orchard or vegetable

crops. Since this year marks the 100" anniversary of the Entomological Society of BC, and the

98" Volume of the Journal of the Entomological Society of BC (JESBC), it also seemed

appropriate in part to link certain aspects of this article towards key papers published in the

Journal since 1951.

General Trends

Reviewing the past fifty issues of the JESBC was a tremendously educational and nostalgic

experience, not only from an entomological point of view, but also as a general interest and

historical exercise. Who could resist reading, for example, ‘An authenticated case of black

widow bite’, by Carl and Perry in 1959, or so many of the other key articles that in retrospect

helped capture our interest and shape our profession in BC. When I had gathered and

summarized all of the papers pertaining to entomology in orchards and vegetables since 1951,

I was greatly impressed by the diversity and efficacy of our Society’s entomologists, both past

and present, and by the ebb and flow of various research themes over time.

Volume 47 of the Proceedings of the Entomological Society of British Columbia, issued

July 15, 1951, began with an ad by the Nichols Chemical Company congratulating the

Entomological Society of British Columbia on its 50" anniversary. I found it interesting to

note that there was a total of 8 pages of advertisements by various pesticide companies in that

issue, most of which (Monsanto being a notable exception) are no longer in existence. One ad

in Volume 49, 1953, had the ominous title ‘Improved Controls for Entomologists’, and on

another page was a photo of two workers (possibly the entomologists referred to in the first ad)

wearing no safety equipment, in short-sleeved shirts, applying pesticides in an apple orchard

using hand-held sprayers (the chemicals advertized in that ad included DDT, parathion, and

lead arsenate). Such photographs of course, are now used to illustrate how ‘not’ to apply

pesticides, and these advertisements went the ultimate way of DDT following the 1954 issue.

144 J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

The prominence of pesticide advertisements in the JESBC at that time reflected the

prominence and influence of the pesticide industry on entomological research. The papers

published in the Journal between 1951 and 1960 were heavily biased towards pesticide trials

(including studies on efficacy, resistance, phytotoxicity, application technology, etc.),

seemingly at the expense of general biological studies (general species descriptions, host range

and damage, life history, insect ecology, behavior, etc.) and studies on alternative pest control

methods (biological, cultural, natural, physical, mechanical, genetic, semiochemical or SIR)

(Fig. 1). It is interesting to note that the number of pesticide-related articles published in the

Journal has declined linearly from a high of 25 papers from 1951-60 to a low of 3 papers in the

last decade. This probably reflected in part the gradual erosion in the number of existing and

pending product registrations, as well as the reviving interest in insect biology and the

development of integrated pest management (IPM) theory, tools and strategies (alternative

controls, sampling and forecasting, economic thresholds, blended control strategies, etc.). The

number of papers in the general biology category overtook pesticide articles in the Journal

between 1961-70, and articles depicting IPM and alternative control approaches have also

been gradually increasing (Fig. 1).

Mm General biology

Pesticide + resistance

[CJ] Alternative controls

Number of papers

Figure 1. The number of papers published between 1951 and 2000 in Journal of the

Entomological Society of BC relating to: general biological studies; pesticide studies; and

alternative control methods of insects and mites in orchard and field vegetable crops.

1961-70 1971-80 1981-90 1991-00

pth

fo)

Although the emphasis and general infrastructure of entomological research has changed

dramatically in orchard and field vegetable crops over the past fifty years, the insects and mites

involved have changed but little. The next sections chronicle the entomologists who have

studied insects and mites in orchards and vegetables in BC, and a few examples of key pests

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 145

that have demanded a relay team of entomologists and effort spanning the past fifty years are

given. Since most entomological research in orchards and vegetables can be classed under

biological, chemical or alternative IPM research, these will also serve as broad themes for the

discussions to follow.

Entomology in Orchard Crops

The majority of orchard crops in BC, (primarily apple, pear, peach, apricot, plum or prune,

and cherry) are grown in the Okanagan and Similkameen Valleys with some production in the

Kootenays and more recently in the lower Fraser Valley. Most of the research in orchards over

the past fifty years, however, has focused on the Okanagan and Similkameen areas, primarily

by Agriculture and Agri-Food Canada scientists at the Pacific Agri-Food Research Centre

(PARC) at Summerland (formerly the Summerland Research Station). An interesting and

historic paper was published in the JESBC in 1953 entitled “A decade of pest control in BC

orchards” by J. Marshall, senior entomologist at the Summerland Research Station. A buildup

of federal and provincial staff at Summerland was occurring at that time and Marshall spoke of

the facility as having struck a good balance between fundamental long-term biological studies

and the chemical investigations that were considered a season-to-season necessity by the

industry. In the 1950s, scientists at Summerland included: J. Marshall (specialty: pesticides);

D.P. Pielou (specialty: aphids and resistance to insecticides); C.V.G. Morgan (specialty:

Eriophyid mites and scale insects); M.D. Proverbs (specialty: codling moth irradiation) and;

R.S. Downing (specialty: insecticides and mites). In the 1960s, W.H.A. Wilde transferred to

Summerland from the Creston substation in 1961, retired in 1963 and was replaced by R.D.

McMullen (their specialty: pear psylla bionomics) who transferred from the Harrow Research

Station in Ontario in 1964. H.F. Madsen (specialty: integrated control) joined Summerland in

1964 and replaced Marshall (retired in 1963) as head of entomology. In the 1970s, F.L.

Banham, formerly studying vegetable insects began orchard research in 1971 (specialty: stone

fruit insects), and following the retirements of Morgan in 1974, Downing in 1979, and

Proverbs in 1980, the entomology team at Summerland by 1982 consisted of McMullen,

Banham, Madsen and newcomers N. Angerilli (Specialty: orchard mite control, San Jose scale)

and V.A. Dyck (specialty: management of codling moth). J.E. Cossentine (specialty: biological

control) replaced the retired Madsen in 1985, and G.J.R. Judd was appointed in 1989

(specialty: insect chemical ecology and behavior). M. Smirle (specialty: resistance

management) joined the station in 1990 and D. Thielmann (specialty: insect baculoviruses)

transferred from the Vancouver Research Station upon its closure in 1996. H. Thistlewood

transferred to Summerland from the Vineland Ontario Research Station in 1998, was

temporarily seconded to the SIR program as general manager between 1998 and 2001 and is

now at PARC, Summerland (specialty: insect ecology). Surrounding this core of research

scientists have been many technical staff, including, C.J. Campbell, W.W. Davis, M. Gardiner,

L.B. Jensen, C. Jong, C. Krupke, D.M. Logan, T.K. Moilliet, J.R. Newton, and J.M. Vakenti

(only individuals, whose names have appeared in the literature as author or co-author are

listed).

In addition to the federal government researchers mentioned above, significant

contributions have also been made by entomologists from the Provincial Government (C.L.

Nielson; J.C. Arrand; J. Corner; P.J. Procter; and H. Philip), private consultants (S. Haley;

J.M. Vakenti; L. Edwards; F. Peters; D. Thomson) and the Okanagan-Kootenay Sterile Insect

Release Program (K.A. Bloem and S. Bloem). Contributions to orchard entomology have also

been made by the various BC universities, including UBC (D.A. Chant) and Simon Fraser

University (J.H. Borden; B.P. Bierne; B.D. Roitberg; M. Mackauer; G. Gries;) which has had

Close ties to PARC, Summerland through the Centre for Pest Management (hosting the Master

146 J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

of Pest Management program (MPM)). The impact of SFU and the Centre for Pest

Management on current research personnel at PARC Summerland is evidenced by the fact that

three out of five current research scientists (Judd, Smirle and Thistlewood) as well as

numerous technical staff (M. Gardiner, M. Claudius and C. Krupke) are former SFU graduate

students. Numerous collaborations between SFU and PARC, Summerland scientists involving

graduate students have also resulted in significant contributions to orchard pest management,

and PARC, Summerland scientists have hosted the orchard pest management summer course in

the MPM program since 1973.

General Biology

The list of insect and mite pests as well as the associated suite of beneficial organisms in

orchard crops in BC is very long, and has provided entomologists with an abundance of

challenging material for study over the years. Downing ef al. (1956) compiled a list of 63

species of insects and 14 species of mites known to be economically (E) or sporadically (S)

injurious to apples (28 E; 22 S); apricot (10 E; 12 S); peach (11 E; 14S); pear (12 E; 15 S);

plum or prune (15 E; 23 S); and cherry (12 E; 26 S). This list has grown since then, and I’m

sure the next fifty years will see a number of major new introductions and challenges to the

industry, the apple maggot, Rhagoletis pomonella (Walsh), for example, being an imminent

threat.

Much of the attention of orchard entomologists over the past fifty years has focussed on:

scale insects (e.g. San Jose scale, Quadraspidiotus perniciosus Comstock), mites, both pest

(e.g. the McDaniel mite, Tetranychus mcdanieli McGregor, and European red mite,

Panonychus ulmi (Koch)) and predators (e.g. phytoseiid mites including 7yphlodromus

occidentalis Nesbitt); lepidopterans (e.g. codling moth, Cydia pomonella (L.), obliquebanded

leafroller, Choristoneura rosaceana Harris); pear psylla, Cacopsylla pyricoli (Foerster) and

the western cherry fruit fly, Rhagoletis indifferens Curran. In addition to their popularity as

research organisms, some of these species are also the focus of anumber of success stories that

will be highlighted below.

Insect and Mite Management

The introduction of organochlorine insecticides to the BC orchard industry in the 1940s

had an immediate impact on the population dynamics of insect and mite populations, as well as

on the job descriptions of many entomologists. The initial efficacy of these new products was

so impressive, and the influence of the pesticide companies so great, that much of the research

efforts in the 1940s, 50s and 60s were directed at evaluating various products against the key

economic orchard pests present at that time. A number of the Summerland scientists mentioned

earlier in the 1950s and 60s were very prolific in evaluating insecticides (Marshall, Pielou,

Proverbs) and acaracides (Downing, Morgan). It was fortunate for the industry, however, that

these entomologists were also aware of the drawbacks to indiscriminate pesticide use, and their

concurrent biological and ecological observations and work with less toxic alternatives such as

dormant oil sprays (Downing, Madsen) gradually gave rise to more discriminate pesticide use

and ultimately to widely adopted IPM programs.

As early as 1953, the need for judicious use of chemicals in orchards in order to preserve

beneficial organisms had been recognized by Marshall and others (Marshall 1953). Marshall

published another important paper in JESBC a decade later (1963), entitled, “Background for

integrated spraying in the orchards of British Columbia”, which made reference to the recently

published and “woefully biased” ‘Silent Spring’ by Rachel Carson in 1962 (also reviewed in

JESBC by Marshall in 1962), and which essentially described the evolving concept of IPM in

Summerland and other fruit growing areas of the world. Among other advances, Summerland

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 147

scientists had recently shown that less spraying for insects and mites was possible without crop

losses, and the use of more selective pesticides used only when necessary was being advocated

for various regions of the Okanagan Valley. In apples, a number of breakthroughs in mite and

codling moth control helped shape and direct the course of pest management related research

in this major crop.

Much of our understanding of the life histories and distribution of the pest and predatory

mites in BC orchards can be attributed to the early efforts of N.H. Anderson (e.g. Anderson ef

al. 1958), Morgan (e.g. Morgan ef al. 1955), and Downing (e.g. Downing and Moilliet 1971)

whose orchard survey and life history work formed the foundation for the integrated mite

controls now standard throughout the industry. An important tool in the understanding and

management of mites was the mite brushing apparatus, which was adapted by Morgan for use

in Okanagan orchards (Morgan et a/. 1955) and has been in standard use for about 50 years.

This apparatus allowed researchers to study both phytophagous and predaceous mites, and

with the development of accompanying action thresholds it has become a cornerstone of apple

and pear IPM programs delivered by private consultants and packing houses. Along with the

findings that dormant oil sprays could control the European red mite (and certain scale insects)

without impacting predator mites, and that the predatory mite, 7yphlodromus occidentalis had

developed natural resistance to organophosphates such as azinphos methy] (Guthion), growers

have been able to rely heavily on naturally occurring biological control backed up by

surveillance-based miticide applications since the early 1970s.

The codling moth, Cydia pomonella, is the key insect pest of apples and pears world-wide

and much of the general biology of this pest has been determined elsewhere, or pre-dates the

current review. However, it is in the development of sophisticated IPM tools and strategies for

codling moth management that a number of BC entomologists have distinguished themselves.

In the Okanagan and elsewhere, codling moth could only be managed in the 1950s and 60s by

repeated use of a variety of insecticides, however three major developments, including:

autocidal control; pheromone trapping; and mating disruption, have irrevocably changed this

tradition over the past 30 years.

Autocidal control, later to become known as the Sterile Insect Release (SIR) program, was

initiated in Summerland in 1956 by M.D. Proverbs with the ultimate aim of eradicating codling

moths from geographically isolated areas. Probably the most challenging aspect of this

program was the development of an artificial diet and mass rearing facility for codling moth,

the efficiency of which was evolved by Proverbs’ team to the point that about 17 million sterile

moths were eventually being reared and released annually. The rearing facility itself at the

Summerland Research Station was an amazing example of what can be accomplished with

ingenuity and a shoestring budget, and the larger present day facility near Osoyoos was

modeled very closely after the original. The efficacy of SIR was demonstrated in a number of

isolated orchards between 1964 and 1976, and from 1976 to 1978 was expanded to include

520 ha of apples and pears in the geographically isolated Similkameen Valley. By the end of

the project in 1978, codling moth populations and associated apple damage had been virtually

eradicated from the Valley (Proverbs ef a/. 1982), and no additional measures for codling moth

control were required in any of the orchards until 1981.

Proverbs’ SIR program had gained worldwide attention and recognition by his retirement

in 1980, and in 1988 plans for the resurrection and expansion of the SIR program to cover the

entire Okanagan and Similkameen valleys were established in a cooperative effort between the

Summerland Research Station (V.A. Dyck) and the BC Fruit Growers’ Association. In 1990,

the Okanagan Similkameen SIR program was formally launched with the goal of eradicating

codling moth from key growing regions of the Okanagan and Similkameen Valleys of BC by

1999. The hiring of staff, including K.A. Bloem as program manager (succeeded in 1998 by H.

148 J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

Thistlewood), began in 1992, and releases of sterile moths began in 1994 in the south

Okanagan (Summerland and Naramata south to the U.S. border), Similkameen (Cawston,

Keremeos) and Creston regions (known as Zone | of the SIR). The program has since

expanded to the central (Zone 2) and north (Zone 3) Okanagan regions. After 7 years of moth

releases, populations of codling moth had declined dramatically throughout Zone 1, but

functional eradication had only been achieved in those regions of the more segregated

Similkameen valley where Proverbs did his initial work. Although the goals of the SIR

program were shifted from eradication to area-wide suppression in 1999, the autocidal control

approach has reduced codling moth populations over large areas to unprecedented low levels.

An interesting paper was published in the JESBC by Madsen and Davis in 1971 which

described the use of female-baited traps as indicators of codling moth populations and apple

damage. This paper was followed by another JESBC paper by Madsen and Vakenti in 1973,

which recorded the initial use of traps baited with synthetic pheromones for codling moth and

fruit-tree leafroller, Archips argyrospilus (Walker) in orchards in BC. Pheromone traps soon

after became a widely used and indispensable tool in orchard IPM in BC, and provided an

important cornerstone to the codling moth SIR program still under development at that time as

well as in the current SIR program.

Possibly the most exciting development in apple IPM in the past decade has been the

development of pheromone-based mating disruption technology for area-wide management of

codling moth and other lepidopterous pests of apples and pears (i.e. Choristoneura rosaceana

and Pandemis limitata (Kearfott)). This work, collectively, has been led by G.J.R. Judd at

PARC, Summerland in association with industry (Pacific Biocontrol, 3-M Canada, and

PheroTech Inc.) and colleagues at SFU, including professors J.H. Borden, B.D. Roitberg, and

G. Gries and graduate students M.L. Evenden, N. Delury, H. McBrian and J.P. Deland. In

1992, Isomate-C was registered in Canada by Pacific Biocontrol (Vancouver, WA) as a mating

disruption product for codling moth control, and has since been used in Canada and the USA

for area-wide management of codling moth (e.g. Judd ef a/. 1996). Mating disruption has

become the preferred method for reducing codling moth populations in the clean-up stage of

both conventional and organic orchards in Zones 2 and 3 of the Okanagan Similkameen SIR

program, and is now being rationally integrated with the sterile moth release program for non-

chemical management of codling moth in Zone |. Mating disruption has also been

demonstrated for a number of leafroller pests in BC orchards, and eventual registrations for

these additional mating-disruption products will further reduce the reliance of the industry on

insecticides.

Entomology in Vegetable Crops

Commercial field vegetable crops are grown in many areas of BC, with the largest

concentration of acreage historically being in the lower Fraser Valley and south central interior

including the Okanagan Valley, Kamloops and Kootenay areas. As was observed in orchard

crops, most of the published entomological research in vegetable crops up until the last decade

has involved Agriculture and Agri-Food Canada scientists working out of various research

stations or substations in the above regions. In the 1950s, entomological research in vegetables

was strong in Kamloops (at that time the Dominion Field Crop Insect Laboratory), Victoria

(Saanichton Station), Agassiz (formerly the Agassiz Research Station) and Vancouver

(formerly the Dominion Laboratory of Plant Pathology at UBC). In Kamloops in the 1950s,

scientists included: R.H. Handford (officer in charge); D.A. Arnott; H.R. MacCarthy; D.G.

Finlayson; and F.L. Banham. At Saanichton, scientists in the early 1950s included K.M. King,

A.T.S. Wilkinson and A.R. Forbes, however, vegetable research was phased out in the mid-

1950s with the retirement of King in 1956, and the transfer of Wilkinson (specialty: soil

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 149

insects and biological control of weeds) and Forbes (specialty: aphids and aphid morphology)

to Vancouver. At that time, federal entomologists in Vancouver were located on the UBC

campus in a building since converted into a cafeteria known as ‘the Barn’, and also included

MacCarthy (specialty: virus vectors) who transferred from Kamloops in 1955. They relocated

to the newly built Vancouver Research Station (VRS) at UBC in 1959, and were joined by

Finlayson (specialty: root maggots; toxicology) who transferred from Kamloops that year.

B.D. Frazer (specialty: aphid ecology) joined the VRS in 1967 and the group remained intact

until the retirement of MacCarthy in 1976. In 1981, R.S. Vernon (specialty: vegetable insect

IPM) replaced the retired Finlayson (1980), and D.A. Raworth (specialty: biological control)

was appointed in 1984. Vegetable research at Agassiz was conducted by R. Glendenning until

his retirement in 1953, as well as H.G. Fulton (specialty: vegetable insects), who was stationed

at the Entomology Laboratory substation of Agassiz in Chilliwack BC (which later became a

VRS substation) until his retirement in 1967. With the closure of the VRS in 1996, Vernon and

Raworth (as well as small fruit specialist S. Fitzpatrick, specialty: semiochemicals) were

transferred to Agassiz (now the Pacific Agri-Food Research Centre (PARC)) to join D.

Gillespie (specialty: greenhouse vegetables), V. Brookes (specialty: minor use registration)

and J.T. Kabaluk (specialty: geographic information systems and microbial control). In 1965,

Banham transferred from Kamloops to Summerland and was their sole vegetable insect

specialist until his specialty was changed to tree fruits in 1971. Excellent technical staff have

complimented this core of research scientists, including, C.J. Campbell, C.K. Chan, T.

Kabaluk, M. Knott, J.R. Mackenzie, R.R. McGregor and M.D. Noble (again, only individuals,

whose names have appeared in the literature as author or co-author are listed).

As was observed above in orchards, additional professionals have contributed in various

ways to entomology in field vegetables, including extension officers from the Provincial

Government (H. Gerber, H. Philip, B. Costello, J.C. Arrand, T. Kluge, L. Gilkinson) and

private consultants (R.S. Vernon, G.J.R. Judd, W.B. Strong, B.D. Henderson, S.Y. Li).

Contributions to vegetable entomology have also been made by the various BC universities,

including UBC (M.B. Isman) and Simon Fraser University (e.g. J.H. Borden; B.P. Bierne;

B.D. Roitberg; M. Mackauer; G. Gries;), the latter again having close ties to the Vancouver

and Agassiz Research Stations through the Centre for Pest Management. In fact, the

development and implementation of many of the vegetable IPM programs in current use in BC

has been through collaborative efforts between the Vancouver and Agassiz Research Stations

and SFU staff and graduate students since the mid 1970s.

General Biology

Vegetable crops in BC include a multitude of plant families, species and cultivars, and as

such the list of insects associated with these crops Is also very long. It is interesting to note that

many of the key pests of our more important vegetable crops are not native to Canada, and it is

these pests that has demanded much of our attention over the past fifty years. Among the more

important introduced pests to BC are: root feeding maggots (e.g. the onion maggot, Delia

antiqua (Meigen), the cabbage maggot, Delia radicum (Bouche) and the carrot rust fly, Psila

rosae (F.)); wireworms (e.g. the dusky wireworm, Agriotes obscurus (L.) and the lined click

beetle A. /ineatus (L.)); lepidopterans (e.g. imported cabbageworm, Pieris rapae (L.)); and

aphids (e.g. the lettuce aphid, Nasonovia ribis-nigri (Mosley)). Other important pests are

endemic to the USA and may have entered BC by natural avenues of dispersion or through

man’s activities (e.g. the tuber flea beetle, Epitrix tuberis Gent. and the Colorado potato beetle,

Leptinotarsa decemlineata (Say)). Much of the research effort in BC vegetables has involved

root maggots, tuber flea beetles and various aphids, and brief summaries of the research

activities devoted to these pests are given below.

150 J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

Root maggots. Root maggots are pests of many vegetable crops (i.e. all cruciferous crops,

onions and carrots) in BC, and entire fields can be destroyed without adequate protection.

Most of what is known about the general biology of root maggots attacking cruciferous crops

in BC (particularly D. radicum) is through the independent or combined work of Forbes and

Finlayson (e.g. Forbes and Finlayson 1957). From the 1950s to the 1980s the majority of

research on D. radicum as well as on the onion maggot, D. antiqua, and carrot rust fly, P.

rosae, involved the screening of a wide variety of insecticides (Forbes, Fulton, Wilkinson,

Finlayson, Mackenzie, Vernon). The amount of spraying for these pests, however, had become

excessive, and in the late 1970s to early 1980s a number of monitoring devices and threshold-

based IPM programs were developed for onions and carrots through graduate studies at SFU

by Vernon and Judd (under the guidance of J.H. Borden). These programs dramatically

reduced insecticide use in these crops in the Cloverdale Valley, and formed the basis of BC’s

first grower-funded private IPM consulting company, Monagro Consultants, in 1979.

Flea beetles. The tuber flea beetle, E. tuberis, at one time was one of the most important

pests of potatoes in BC, and all published literature on the biology of this insect has been

through the efforts of MacCarthy, Banham, Finlayson, Fulton, Vernon and Thomson. As with

root maggots, considerable research on this pest since 1950 has been devoted to establishing

chemical controls, and growers had become accustomed to spraying their potato fields on a

routine basis with broad-spectrum insecticides. This practice also established the need for

routine sprays for aphids, largely due to the coincident elimination of aphid parasitoids and

predators. In the early 1980s, a monitoring program was established through the efforts of

Vernon and SFU graduate students K. Giles and M. Cusson, and this program has been

provided to the majority of potato growers in BC since 1980 by private consultants (in

particular by Monagro Consultants, and ES Cropconsult Ltd.). Since flea beetles enter potato

fields from the field margins, it has been shown that early-season monitoring can detect when

and where they first occur, and populations can generally be controlled for the entire season

with one (or zero) carefully timed and placed edge spray. By not spraying the entire field ona

routine basis, the control of aphid populations through the natural buildup of biological control

agents is now common practice.

Aphids. Aphids are found as pests of many vegetable crops, and the efforts of

entomologists at the Vancouver Research Station (until its closure in 1996) have been pivotal

in determining their biology and control. The early work of H.R. MacCarthy, for example,

helped build our understanding of the relationship between aphids and their ability to transmit

various virus diseases to potatoes. There is no doubt, however, that the research of A.R. Forbes

and his technician C.K. Chan has been most instrumental in building our base of knowledge on

the biology and distribution of aphids in BC, many of them pests of vegetable crops. Their

work, along with contributions by MacCarthy and Frazer, was published almost annually in a

series of 21 papers in the JESBC between 1973 and 1993 (e.g. Forbes and Chan 1991). With

the last paper published (Chan and Frazer 1993), this team had described 412 aphid species

collected from 1243 different host plants and had identified 2434 aphid-host associations. The

collective efforts of these entomologists helped set the stage for the development of

management programs for several key aphid species (e.g. lettuce aphid, NV. ribis-nigri and the

European asparagus aphid, Brachycolus asparagi Mordvilko).

It is hoped that this paper has adequately portrayed the personalities, scope and at least

some of the more public aspects of entomological endeavor in orchards and field vegetable

crops in BC. The work of entomologists not profiled in the limited space available is of no less

importance, and like a jigsaw puzzle, every piece is required to make the image complete.

Assuming some of us will still be around in 2051, it will be interesting to see how much more

of the orchard and vegetable entomology puzzle will have been completed when the next fifty

years of entomology is reviewed.

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 15]

REFERENCES

Anderson, N.H., C.V.G Morgan, and D.A. Chant. 1958. Notes on occurrence of 7yphlodromus and Phytoseius

spp. in southern British Columbia (Acarina: Phytoseiinae). The Canadian Entomologist 90: 275-279.

Carl, G.C. and A.W. Perry. 1959. An authentic case of black widow bite. Journal of the Entomological Society

of British Columbia 56: 21-22.

Carson R. 1962. Silent Spring. Houghton Mifflin, Boston.

Chan, C.K. and B.D. Frazer. 1993. The aphids (Homoptera: Aphididae) of British Columbia 21: Further

additions. Journal of the Entomological Society of British Columbia 90: 77-82.

Downing, R.S. and T.K. Moilliet .1971. Occurrence of phytoseiid mites (Acarina: Phytosetidae) in apple

orchards in south central British Columbia. Journal of the Entomological Society of British Columbia 68:

33-36.

Downing, R.S., C.V.G. Morgan, and M.D. Proverbs. 1956. List of insects and mites attacking tree fruits in the

interior of British Columbia. Journal of the Entomological Society of British Columbia 52: 34-35.

Forbes, A.R. and C.K. and Chan. 1991. The aphids (Homoptera: Aphididae) of British Columbia 20: Further

additions. Journal of the Entomological Society of British Columbia 88: 7-14.

Forbes, A.R. and D.G. Finlayson. 1957. Species of root maggots (Diptera: Anthomyiidae) of cruciferous crops in

British Columbia. Journal of the Entomological Society of British Columbia 54: 25-28.

Judd, G.J.R., M.G.T. Gardiner, and D.R. Thomson. 1996. Commercial trials of pheromone-mediated mating

disruption with Isomate-C to control codling moth in Brisith Columbia apple and pear orchards. Journal of

the Entomological Society of British Columbia 93: 23-34.

Madsen, H.F. and W.W. Davis. 1971. A progress report on the use of female-baited traps as indicators of codling

moth populations. Journal of the Entomological Society of British Columbia 68: 11-14.

Madsen, H.F. and J.M. Vakenti. 1973. The influence of trap design on the response of codling moth

(Lepidoptera: Olethreutidae) and fruittree leafroller (Lepidoptera: Tortricidae) to synthetic sex attractants.

Journal of the Entomological Society of British Columbia 70: 5-8.

Marshall, J. 1953. A decade of pest control in British Columbia orchards. Journal of the Entomological Society

of British Columbia 49: 7-11.

Marshall, J. 1962. Book review (on Silent Spring). Journal of the Entomological Society of British Columbia 59:

53-55.

Marshall, J. 1963. Background for integrated spraying in the orchards of British Columbia. Journal of the

Entomological Society of British Columbia 60: 26-29.

Morgan, C.V.G., D.A. Chant, N.H. Anderson and G.L. Ayre. 1955. Methods for estimating orchard mite

populations, especially with the mite brushing machine. The Canadian Entomologist 87: 189-200.

Proverbs, M.D., J.R. Newton and C.J. Campbell. 1982. Codling moth: A pilot program of control by sterile

insect release in British Columbia. The Canadian Entomologist 114: 363-376.

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 153

History of forest insect investigations in British Columbia

This three-part paper describes the history of forest entomology in British Columbia

during the past century. The first part discusses programmes and personnel in the federal

and provincial governments, private enterprise, and academic institutions. This section was

co-authored by current or former employees of these agencies as a reflection of the close

co-operation between them over the years ranging from insect management programs,

research, and training of forest health personnel. The second part describes a unique

federal entomology program, the history of the former Forest Insect and Disease Survey

(FIDS) unit of the Canadian Forest Service. The third part is a brief history of the

establishment and programmes of the federal Vernon Laboratory, the first permanent

facility in British Columbia dedicated to forest insect investigations.

I. Forest entomology education, research, and insect management

a. Forest entomology in the British Columbia Ministry of Forests and

private sector

b. Teaching and research at academic institutions

c. Research in the federal government

II. Forest Insect and Disease Survey in the Pacific region

III. The Vernon Laboratory and federal entomology in British Columbia

Throughout the history of forest entomology in BC, there has been substantial and

productive interaction between provincial, federal and academic institutions. In addition,

industry has supported major initiatives in research, e.g., on ambrosia beetles and bark

beetles. Thus, there is unavoidable overlap when attempting to synthesize the contributions

of each.

I. Forest entomology education, research,

and insect management

P. M. HALL

BC MINISTRY OF FORESTS,

1450 GOVERNMENT ST., VICTORIA, BC, CANADA V8W 9C2

J. M. KINGHORN

1253 PALMER RD., VICTORIA, BC, CANADA V8P 2H8

B. S. LINDGREN

FACULTY OF NATURAL RESOURCES AND ENVIRONMENTAL STUDIES,

UNIVERSITY OF NORTHERN BRITISH COLUMBIA,

3333 UNIVERSITY WAY, PRINCE GEORGE, BC, CANADA V2N 4Z9

J. A. MCLEAN

FOREST SCIENCES, UNIVERSITY OF BRITISH COLUMBIA,

2357 MAIN MALL, VANCOUVER BC, CANADA V6T 1Z4

L. SAFRANYIK

NATURAL RESOURCES CANADA, CANADIAN FOREST SERVICE,

PACIFIC FORESTRY CENTRE, 506 WEST BURNSIDE ROAD,

VICTORIA, BC, CANADA V8Z 1IM5

154 J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

a. Forest entomology in the British Columbia Ministry of Forests and private sector

Background

“For several years reports have been received from various points in British Columbia

indicating considerable loss from Bark-beetle attack to standing timber and logs. The

lumber industry of the province is of such importance, and the destruction by forest insects

in the States to the south has been reported as so serious in recent years, that it was

thought advisable to make a survey of the actual conditions in regard to injurious forest

insects in British Columbia forests. The Forestry Branch of the Department of Lands of

British Columbia had in the meantime requested the Division of Entomology to undertake

such an investigation. Accordingly, with the assistance and co-operation of the Provincial

Forestry Branch, a survey was made during the summer of 1913 the object of which survey

was to determine the location and extent of the chief forest insect injuries, and to decide

upon proper control measures for the more serious outbreaks.” (Swaine 1914)

And so began the history of forest entomology in British Columbia. Until relatively

recent times, the British Columbia Forest Service (BCFS) did not employ entomologists or

pest management personnel; it relied on the federal government and universities for this

expertise. However, there were numerous instances where the BCFS continuing interest in

forest insect impacts and management was shown.

The Chief Forester of British Columbia, Mr. H.R. MacMillan, made the initial request

to the Division of Entomology and provided funding support to J.M. Swaine to carry out

the survey. Subsequently, the BCFS utilized the expertise of federal entomologists such as

R. Hopping in supervising a number of bark beetle control efforts utilizing crews for

falling and burning infested trees and directed harvesting of infested stands and trees.

These control programs were funded by the provincial Forest Service (Hopping 1921). As

bark beetle infestations continued to arise, so did further control programs, still relying on

federal entomologists for professional expertise (Hopping and Mathers 1945).

The vast and diverse forests in British Columbia result in a similar diversity of forest

insects, many of which can cause damage that affects different forest resource values.

However, timber losses have long been of greatest interest to management agencies and

industry. Through the years, each decade presented the province with one or more major

insect-related issues, most often in the form of extensive outbreaks of bark beetles,

Dendroctonus spp., or defoliators.

Up until about the 1960s, much of British Columbia’s forest industry was concentrated

in coastal areas. However, the extensive stands of spruce, Picea spp., and lodgepole pine,

Pinus contorta, in the interior of the province beckoned, and, as markets increased forest

industry operations in the interior areas of the province expanded.

The modern era

A major step towards a provincially coordinated forest pest management program came

in 1974 when the Forest Pest Review Committee (FPRC) was formed (Pearse 1976). This

committee was chaired by the Chief Forester for British Columbia and was comprised of

representatives from the Forest Service, other provincial and federal agencies, and industry

associations. The purpose was to discuss specific insect and/or disease outbreaks and make

recommendations as to management directions. Several issues in the 1970s made the

committee relevant and very active:

e the beginnings of the Chilcotin outbreak of mountain pine beetle;

e mountain pine beetle outbreaks in the Prince Rupert Region and in the Flathead

Valley in Nelson Region;

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 155

e the beginnings of the Bowron Lakes spruce beetle outbreak;

ea large scale trial of treatments against Douglas-fir tussock moth in the

Kamloops Region carried out in co-operation with the US Forest Service;

e proposed spraying of a western spruce budworm outbreak in the Fraser Canyon;

and,

e the first detection of gypsy moth in British Columbia found in Kitsilano.

In response to the major forest insect and pesticide related issues, Mr. J.M. (Mike) Finnis

was hired by the Protection Section of the BC Forest Service to act as an in-house advisor

and specialist in pest management issues. Mr. Finnis served primarily as a co-ordinator of

activities and as liaison between provincial and federal agencies (most notably the

Canadian Forestry Service). Paul Wood was hired by the Cariboo Forest Region in the

mid-1970s to address the increasing mountain pine beetle outbreak. Additionally, the

Forest Service routinely provided summer students to assist Canadian Forestry Service

entomology researchers.

The report from the Royal Commission on Forest Resources was published in 1976

(Pearse 1976). This report made a series of recommendations to change how the forest

resource was managed and administered in British Columbia. It set the stage for the

creation of the Ministry of Forests Act (Anon. 1978a) and the Forest Act (Anon. 1978b).

These legislative acts gave a mandate to the Ministry of Forests to “manage, protect and

conserve the forest and range resources of the government, having regard to the immediate

and long term economic and social benefits they may confer on British Columbia”

(Section 4b, BC Ministry of Forests Act, 1978). The new legislation led to a large-scale

reorganization of the British Columbia Forest Service in 1980.

Staffing up

By 1979 it became clear that the Forest Service required some in-house expertise, who

were capable of training field staff, advising the Ministry Executive on forest health issues,

and implementing large and small management programs. In early 1980, Mr. R.S. (Bob)

Hodgkinson was hired as a forest entomologist by the Forest Service Protection Branch

and based in, Prince George, to evaluate the use of trap trees for reducing losses caused by

spruce beetle, and Mr. P.M. (Peter) Hall was hired as the regional forest entomologist for

the Cariboo Forest Region, Williams Lake, BC.

The reorganization of the Forest Service progressed through the summer of 1980. For

the first time, the provincial government staffed professionals in pest management

positions. In Victoria headquarters, the positions of a Manager, Pest Management (R. F.

(Bob) Deboo), Forest Entomologist (P.M. Hall), Forester (Mike Finnis) were filled. Later,

a Forest Pathologist (J.A. (John) Muir) and a Pesticide Specialist (JF. (John)

Henigman) were added to the Pest Management Section at Protection Branch. Branch

personnel are responsible for developing policy and procedures, advising the Ministry

Executive and Government, providing co-ordination with other provincial, federal and

private agencies, monitoring management programs at the regional and district level, and

providing expert technical advice. The name of the Pest Management Section was changed

to Forest Health in 1990 to reflect a shift in philosophy away from traditional pest control

and towards more integrated forest management.

156 J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

At the regional level, Pest Management Co-ordinators were staffed at each of the six

regional offices:

Region Location Initial Pest Management Subsequent Forest

Co-ordinator Positions Health Forester

Positions

Cariboo Williams Lake D. Doidge M. Hamm*

Kamloops Kamloops R. Edwards D. Calder*

Nelson Nelson A. Renwick J. Monts

E. Morris

R. Stewart*

Prince Prince George R, Cozens S. Taylor*

George

Prince Smithers V. Barge M. Geisler

Rupert B. Young*

Vancouver Vancouver S. Raine P. Wood

R. Heath*

* the last to hold the position

The position of Pest Management Co-ordinator was eliminated in 1987 and was

replaced in various regions with the position of Forest Health Forester. The latter position

was not necessarily supervisory to the regional forest entomologist.

The regional entomologist position evolved from necessity in those regions

experiencing a wide variety of issues. Staffing of this position occurred over a period of

years, with the formal creation of the position in 1984:

Forest Entomologist

Region Location Initial* Subsequent*

Cariboo Williams Lake P. Hall R. Heath

L. Rankin*

Kamloops Kamloops R. Chorney T. Maher

L. Maclauchlan*

Nelson Nelson D. Gray E. Morris

A. Stock*

Prince George Prince George R. Hodgkinson*

Prince Rupert Smithers A. Stock T. Ebata

K. White*

Vancouver Vancouver/ E. Jeklin D. Heppner*

Nanaimo

*the last (current) to hold the position

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 157

Regional entomologists are responsible for providing expert technical advice to

regional and district staff, co-ordinating and monitoring management programs, training,

and liaison with other agencies, industry and the public. Regional forest pathologists were

also hired in most regions.

Additionally, entomologists or pest management specialists were hired in other areas of

Ministry of Forests operations:

Position Specialist/Entomologist*

Nursery Pest Management Specialist G. Shrimpton

D. Trotter*

Cone and Seed Pest Management Specialist D. Summers

R. Bennett* (Victoria)

W. Strong* (Vernon)

*the last (current) to hold the position

No specific pest management staff were placed in district offices at that time; the

District Resource Officer — Protection, carried out the function of pest management. Since

then, several districts with ongoing forest health issues have established a Forest Health

officer position. Many of these staff have provided long-term continuity to insect

management programs at the district (operational) level. District staff has contributed

substantially to the development of management programs, and has been instrumental in

ensuring that the BC Forest Service remains one of the best Forest Health organizations in

North America.

Contractors and industry

The forest industry has usually depended upon the Canadian Forestry Service, the

universities, and the BC Forest Service for professional entomological advice. They also

regularly employ consultants and contractors to carry out specific studies or projects

relating to forest entomology. There have, however, been instances where the industry

directly employed entomologists, particularly to deal with issues such as ambrosia beetles

which were beyond the mandate of the BC Forest Service. After his retirement from the

Canadian Forest Service, H.A. Richmond consulted to MacMillan-Bloedel to deal with

ambrosia beetle damage, and he was later employed as an in-house consultant to the

Council of Forest Industries. He provided advice on a variety of forest insect issues

(Richmond 1983). Further, Pacific Forest Products hired R. (Dick) Heath as a pest

management specialist, Northwood Pulp and Timber hired 7. (Tom) Maher, and Finlay

Forest Industries of Mackenzie, BC, hired D. (Darrell; Devlin to deal with pest

management issues. At the current time, though, no BC forest company has an

entomologist on staff.

Examples of companies that have provided a wide variety of products and services in

forest entomology, are Phero Tech Inc., Delta, BC, and Bugbusters Pest Management,

Prince George, BC. Phero Tech is one of the major North American companies providing

commercial quantities of forest insect pheromones. Both of these companies, as well as

other consulting companies, have employed numerous forest entomologists over the past

15 years.

The number of consulting entomologists and pest management specialists has increased

dramatically in British Columbia since 1980. The increasing interest in and management of

forest insects has created varied opportunities for individuals and companies with

entomological expertise. Downsizing of federal and provincial agencies, as well as

158 J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

increased interest in the field by students entering graduate programs at universities, have

contributed to a growing pool of talented individuals who have contributed to the

development and refinement of management of forest insects in the province. A listing of

individual contractors and companies is impractical in this publication and it would be

impossible to ensure that all deserving individuals were included.

Milestones and accomplishments

The Ministry of Forests has accomplished a great deal in the area of forest entomology

and management of the forest resource to reduce damage caused by insects. The Forest

Service has evolved from an organisation primarily reacting to crisis to having ongoing

integrated programs and staffing dedicated to dealing with all aspects of forest insect

biology and management. Some of the accomplishments have been specific to a particular

insect problem, while others have been in the realm of legislation and program integration

and acceptance. All of the accomplishments have been achieved through the close working

relationship between the forest entomology “team” of the Forest Service.

Some of what has been done is briefly noted in the following table:

Issue Insect Accomplishment

Legislation All Incorporation of forest health principles into all aspects of the

Forest Practices Code of BC Act and Regulations (Anon.,

2001).

Staffing N/a Forest Entomologists are in place in all 6 Forest Regions,

Victoria headquarters, Cone and Seed Program, and Nursery

Program.

Several Forest Districts have dedicated Forest Health Officers.

Manuals and N/a Management guidebooks for a wide variety of forest insects

Publications have been produced under the authority of the Forest Practices

Code of BC Act. These include guidebooks for bark beetles,

defoliators, terminal weevils and others.

Extension literature and publications are now available.

The Ministry of Forests website contains a great deal of

information relating to forest entomology:

http://www.gov.bc.ca/for/

Surveys All The Forest Service is now responsible for conducting the

annual aerial overview survey formally conducted by the

FIDS Unit of the CFS. The responsibility for this survey was

assumed by the province in 1996 and has been fully

implemented in 2001.

Support All The Forest Service has consistently supported research efforts

by the Canadian Forestry Service and universities through

direct funding or provision of logistical support.

Specific Bark The Forest Service was responsible for operationally

Management Beetles implementing the use of aggregation pheromones for bark

beetle management.

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 159

Large-scale operational management programs for mountain

pine beetle, Douglas-fir beetle and spruce beetle have been

conducted periodically. Expenditures on these issues have

exceeded $10 million annually.

Province-level and _ landscape-level strategic planning

frameworks have been developed and accepted by all levels of

the Forest Service and industry.

Ongoing support and encouragement for the development of

predictive modelling techniques to improve understanding of

population dynamics and the impacts of management efforts

(e.g., SELES/MPBSIM)

The items noted in the table below show a small subset of the accomplishments in

forest entomology achieved by the forest entomology professionals within the Ministry of

Forests. This group has consistently worked co-operatively together and with others to

ensure that management objectives are met and that impacts of forest insects are

minimized.

Insect Accomplishment

Douglas-fir The Forest Service was the first jurisdiction to use nuclear

tussock moth polyhedrosis virus operationally to terminate DFTM outbreaks.

And the Forest Service has implemented an integrated detection,

evaluation and treatment system for this insect. A true Integrated

Pest Management System.

Spruce weevil The Forest Service has actively supported the development of

assessment models and identification and propagation of resistant

genotypes of spruce.

Gypsy moth The Forest Service has actively supported and participated in efforts

to maintain British Columbia free of gypsy moth.

Completed a detailed Risk Assessment for gypsy moth in British

Columbia

Major issues

Currently, forest insect issues in British Columbia relate to the same insects as in the

past: bark beetles, defoliators, ambrosia beetles, terminal weevils and regeneration

problems, and others. However, new issues arise that deal with responsibility-sharing

between government and industry, changing management objectives over time, new

legislative initiatives, introduction of exotic, potentially damaging insects, encouraging

and implementing new survey and management technologies, and, as always, trying to

grapple with a class of organisms that often seem to be better at what they do than we are

at doing what we do. Perhaps they just have more experience.

b. Teaching and research at academic institutions

Research on forest insects has been conducted at all major institutions in British

Columbia. Some of the major contributions are listed in the next Table.

160 J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

The earliest collections of insect damaged wood in the collections at the University of

British Columbia bear the name of Dr. G. (George) Spencer. Materials he collected more

than half a century ago are still in use at the Faculty of Forestry today. Forest Entomology

has always been a core subject in the academic programs for students wishing to become

professional foresters. The first full time professor of forest entomology was Dr. K. (Ken)

Graham whose interests were in ambrosia beetles and bark beetles. He also wrote one of

the first texts in Forest Entomology (Graham 1963). His successor in 1977 was Dr. J.A.

(John) McLean who also was interested in the general biology of ambrosia beetles,

especially the population surveys and mass trapping in sawmills with pheromone-baited

traps as well as economic impacts of attacks on lumber values. His graduate students have

contributed to our knowledge on ambrosia beetles, defoliators and leader weevils. Several

of McLean’s graduate students hold prominent positions with the BC Forest Service and

the Canadian Forest Service, Victoria, BC. Several other UBC professors have also

contributed to forest entomology studies, especially in the areas of population modelling,

Dr. C.S. (Buzz) Holling; in tent caterpillar and biological control studies, Dr. J. (Judy)

Myers; as well as in insects and weather, Dr. W.G. (Bill) Wellington. In recent years

concerns for sustainable forest management have centered on biodiversity studies of

carabids, sucking insects and bark beetles by Dr. G. (Geoff) Scudder and riparian insects

by Dr. J. John) Richardson. Dr. M.B. (Murray) Isman studied the effects of botanical

insecticides, e.g., neem, on many insects, including forest seedling nursery pests and the

mountain pine beetle.

A strong chemical ecology research group was established by Dr. J.H. (John) Borden,

along with chemists K.N. (Keith) Slessor and A.C. (Cam) Oehlschlager, at Simon Fraser

University (SFU) in the 1970s. Many of Dr. Borden’s graduate students, who over the last

35 years have completed the Master of Pest Management program, as well as more

traditional Masters and Ph.D. programs, now contribute professionally to forest

entomology in BC and are mentioned elsewhere. The SFU team, led by Dr. Borden, and

graduate students have made milestone contributions to the chemical ecology of forest

insects. Over the years, they have identified numerous forest insect pheromones, many of

which have been put to use in pest management applications.

Issue Significant contributions

Bark beetle ecology and Significant contributions in the field of chemical ecology

management and basic biology of numerous important bark beetle

species. Development of applications. Ongoing

investigations in the role of anti-aggregation pheromones

and non-host odours.

Ambrosia beetle ecology Identification and synthesis of semiochemicals for the

and management most important species, and development of applications,

many of which are commercially available. Important

findings regarding economic impact to coastal industry.

White pine weevil ecology Important discoveries on host selection and host resistance

and management mechanisms in collaboration with CFS.

Seed and cone insect Development of semiochemicals and investigations on the

ecology and management ecology and impact of significant seed and cone insects in

collaboration with BCFS and CFS.

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 161

Dr. G. (Gerhard) Gries is currently very active in the area of identification of

semiochemicals in forest insects. Drs. Borden and Gries teach graduate courses in forest

pest management and forest entomology at SFU. In the early program at Simon Fraser, Dr.

B. (Bernie) Roitberg supervised several graduate students studying mate selection

behavior in bark beetles. Dr. A. (Bert) Turnbull wrote on biological control (Turnbull and

Chant 1961), 7. (Thelma) Finlayson described many hymenopterous parasites, and Dr. J.

(John) Webster studied interactions between nematodes and insects, especially the

pinewood nematode.

Forest entomology is taught as part of a Forest Health course at the University of

Northern British Columbia by Dr. B.S. (Staffan) Lindgren, whose major interests are in

forest insect ecology and management. At the University of Victoria, extensive studies on

the biodiversity of insects in the tree crowns of mature coastal rain forest have been

conducted by Drs. R. (Richard) Ring and N. (Neville) Winchester.

Technicians trained at the Technical Institutes and colleges in British Columbia have

received their forest entomology education from a number of instructors over the years,

and it would be impractical to name all of them here. Suffice it to say that forest

entomology is taught at every major college and University College in BC. As well,

teachers and researchers at these institutions have made important contributions to forest

insect biology and management.

c. Research in the federal government

The following is a brief overview of entomology research conducted by the federal

Government in British Columbia. We have listed the scientists grouped approximately by

the decade in which they commenced permanent employment. For each scientist, we give

a brief description of the main subject area(s) of work and/or the highlight of

accomplishments. We do not give a complete list of laboratory and research directors as

this list is available elsewhere. Instead, we listed only those managers who were trained

entomologists and did some forest entomology research during their careers. We made an

effort to provide a complete list of all scientists who, at least some time during their

careers, were permanently employed by the federal government and worked at one or more

of the three entomology labs (Vernon, Vancouver, Victoria). We sincerely apologize for

inadvertent omission of any names. In compiling this information we drew upon our

personal knowledge and the following references: Richmond (1983); Riegert (1991); and

Swaine (1918).

The early years

Significantly, organized forest insect investigations in British Columbia by the

dominion government were started in response to extensive outbreaks by bark beetles

during the first three decades of this century. The following quote from C. Gordon Hewitt,

Dominion Entomologist in Ottawa described the bark beetle problems that existed up to

1917 as follows.

“ The bark-beetles constitute the chief insect enemies of our coniferous forests, and it is

impossible to give even an approximate estimate of the enormous annual loss caused by

their depredations throughout Canada. Much of the dead timber whose destruction is

attributed to fire is the result of outbreaks by bark beetles; this is particularly true in

British Columbia.”

To this day, bark beetles affecting mature pines, spruces, Douglas-fir and sub-alpine fir

collectively remain the most important cause of tree mortality from forest insects in the

Province.

162 J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

Although not residing in British Columbia, J.M. Swaine was the first dominion

government entomologist to carry out major investigations of forest insect biology and

management in the Province. During these early years, there was a lack of information on

the taxonomy, bionomics, and methods of coping with, destructive bark beetles. In

response to the urgent need for this basic information, and the constant demand by

lumbermen, foresters and others for practical methods of control, J.M. Swaine, Assistant

Entomologist in charge of Forest Insect Investigations, undertook a study of the Canadian

bark beetle fauna. This work culminated in the publication in 1917-18 by the Dominion of

Canada, Department of Agriculture of the two-part Bulletin No. 14 entitled Canadian bark-

beetles. It describes several new species from British Columbia. In 1918 The Commission

of Conservation, Canada Committee on Forests published Forests of British Columbia

which included a chapter by Swaine on injurious forest insects. He described outbreaks of

bark beetles affecting ponderosa, western white and lodgepole pine, Engelmann and sitka

spruce, western hemlock, lowland and alpine fir. Most of these outbreaks were occurring

in the southern interior and the southern Rockies.

The beginning of organized research

R. (Ralph) Hopping was hired from California by the Federal Government in the late

1910s to organize suppression programs against outbreaks of the mountain pine and the

western pine beetle in ponderosa and lodgepole pines in the Nicola-Aspen Grove-Merritt

areas. The Provincial Government financed the control operations. In 1923 J.M. Swaine

established the first dominion government forest insect laboratory in Vernon and R.

Hopping was appointed to undertake a program of forest insect investigations in British

Columbia and Alberta. Initially, the total staff of the Vernon laboratory consisted of four

persons: R. Hopping in charge; his son G. (George) R. Hopping, assistant: H. (Hec) A.

Richmond and W. (Bill) G. Mathers, research assistants. For the next two decades, these

entomologists made significant contributions to our knowledge of bark and ambrosia

beetle biology and control, and the biology and control of some important wood borers

(e.g., the red cedar borer) and defoliators (e.g., the hemlock looper, blackheaded

budworm). H. (Hugh) B. Leech collaborated with W.G. Mathers on aspects of bark beetle

research. In his Masters thesis, H.A. Richmond produced the first morphologic description

of adult mountain pine beetles. G.R. Hopping became an authority on the North American

bark beetle genus /ps.

Expansion of entomology investigations

The early 1940s saw a great proliferation of defoliator outbreaks in the coastal regions

of the Province. Dr. M. (Malcolm) L. Prebble was transferred from Fredericton to British

Columbia and established a forest insect laboratory in Victoria. He was joined by Dr. K.

(Ken) Graham and together they carried out detailed investigations of the nature and

causes of outbreaks in the coast hemlock forests by a number of defoliating insects such as

the blackheaded budworm, the hemlock looper, the hemlock sawfly, and the rusty tussock

moth. These investigations included the nature of natural control and considerations for

chemical treatment. D. (Don) N. Smith and G.R. Wyatt assisted with various phases of this

work as well as with subsequent studies by M.L. Prebble and K. Graham on the biology

and damage caused by ambrosia beetles in softwood lumber on Vancouver Island. D.N.

Smith’s main research areas involved nursery and regeneration insects and insect pests of

wood in service. In 1948 K. Graham became professor of forest entomology at UBC and

M.L. Prebble transferred to Sault Ste. Marie, Ontario. Dr. M. (Margaret) R. MacKay

worked as a taxonomist at Vernon until 1949 when she was transferred to Ottawa.

E.D. (Dave) A. Dyer and J. (Jim) M. Kighorn both joined the entomology laboratory

in Victoria in 1946 as student assistants and later (1950) as permanent staff. Over a long,

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 163

productive career that spanned three decades, J.M. Kinghorn made major contributions to

our knowledge of hemlock looper biology, especially the effect of stand conditions on its

epidemiology; ambrosia beetle management, and chemical control of the mountain pine

beetle. During the last decade prior to retirement, he switched careers and pioneered

development of containerized production of seedlings for reforestation. E.D.A. Dyer and

Dr. J. (John) A. Chapman, who joined the entomology lab in Victoria in 1952, were

among the first scientists to demonstrate pheromone production in ambrosia beetle and

spruce beetles. E.D.A. Dyer also made significant contributions to the biology and

management of the spruce beetle. Other scientists that joined the entomology lab in

Victoria during the late 1940s were S. (Stu) Brown, M.G. Thomson, and J. (Jack)

Walters. S. Brown worked on population change in the mountain pine beetle and, in

collaboration with J.A. Chapman and J.M. Kinghorn, ambrosia beetle biology and

management. M.G. Thomson and J. Walters conducted one of the first investigations in the

Province on spruce beetle biology; the former also worked on hemlock looper and

ambrosia beetle problems.

Dr. W. (Bill) G. Wellington joined the Victoria lab in 1953. He made fundamental

contributions to the field of bio-meteorology and insect ecology, especially studies of

insects and climate, and individual differences among insects as factors in population

dynamics. R (Ray) R. Lejune replaced H.A. Richmond as officer in charge of the Victoria

lab in 1955 as the latter started a highly successful career as an entomology consultant; he

was the first entomology consultant in the Province. A. (Al) F. Hedlin moved to Victoria

from the Indian Head station in 1954. He worked on seed and cone insects, became an

international authority on the subject and authored and co-authored a number of definitive

publications on seed and cone insects and their management in Canada and North

America. Dr. L. (Les) H. McMullen joined the forest insect lab in Vernon in the early

1950s and moved to the Victoria lab in 1954. In collaboration with Dr. M. (Mike) D.

Atkins, who joined the Victoria lab in the mid 1950s, Dr. McMullen carried out the first

comprehensive investigations of the ecology of the Douglas-fir beetle in British Columbia.

These studies included flight capacity and dispersal, brood mortality in relation to natural

factors, and the effects of stand conditions and harvesting practices on beetle populations.

This work has lead to the development of the highly effective “Douglas-fir beetle clauses”

in the early 1960s which were written into timber sale licenses in high beetle hazard areas

in the BC interior to ensure sanitary logging practices and treatment of logging residue. Dr.

McMullen has also contributed to studies of mountain pine beetle dispersal and the sitka

spruce weevil biology and management, being the major author of the first computerized

model of spread, intensification of damage and direct control. M. Seger and S. (Sergei) F.

Condrashoff joined the Victoria lab during the early 1950s. Mr. Seger, an insect

pathologist, investigated disease associations of damaging forest lepidoptera. Condrashoff

worked on the biology of needle and leaf-mining insects and regeneration pests such as the

weevil Steremnius carinatus. Dr. D. (Doug) A. Ross joined the Forest Insect and Disease

Survey (FIDS) unit at Vernon in 1950. The main areas of his research were wood borer

biology and control. D. (Dave) Evans joined the FIDS as a taxonomist in the early 1950s.

The main areas of his research concerned the life histories of forest insect pests and

introductions of exotic natural enemies for controlling forest defoliators. Dr. G.T. (Tom)

Silver replaced E.D.A. Dyer as FIDS Survey Head in 1953. Dr. Silver made important

contributions to several fields including the biology of several species of forest defoliators,

sitka spruce weevil biology, and sample survey methods. Dr. D. (Don) K. Edwards, insect

physiologist, joined the insect lab in Victoria in the late 1950s. A productive scientist, Dr.

Edward’s work was centered on techniques for measuring activity rates and rhythms in

insects such as the effects of electrical fields or acclimatization, and development of

sampling techniques to measure defoliator density.

164 J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

The new forest research laboratory was opened on Burnside Road, Victoria, during the

early 1960s and all subsequent entomology work was done out of this facility with the

exception of a few entomologists working with the FIDS who remained in Vernon until

1970 when that laboratory was closed. Dr. C.S. (Buzz) Holling worked at the Victoria lab

during the 1960s. His main interest and work concerned basic principles of insect

predation and process modelling in insect ecology. He was recognized internationally as an

authority on these subjects. In 1960, Dr. O. (Ozzie) N. Morris succeeded M. Sager as

insect pathologist. He investigated, and produced a catalogue of insect pathogens

associated with forest lepidoptera in British Columbia. Drs. J.R. (Rod) Carrow

(entomologist), D.B. (Bir) Mullick and G. (George) S. Puritch (plant physiologists) joined

the entomology research group in the mid-1960s. Their research focussed on the nature

and effects of resistance in true firs to the balsam wooly aphid, and induced physiological

and biochemical changes in the tree as it affected aphid establishment and survival. Dr. J.

(John) W.E. Harris who joined the FIDS unit in 1964, collaborated with assessment of

natural enemy complexes associated with the balsam wooly aphid. Other accomplishments

included the biology of the poplar and willow borer, development of survey methodology,

including remote sensing, and assessment of the effectiveness of introduced parasites and

predators in controlling the larch sawfly, the larch casebearer and other forest insects. Dr.

T. (Tara) S. Sahota, insect physiologist, joined the entomology research group in 1967.

He made important contributions to several fields including hormonal regulation of

metamorphosis and reproduction in bark beetles, and quality of individual insects as

affecting numerical changes in populations. Currently semi-retired, his work during the

past 5 years concerns the reproductive physiology and behaviour of the spruce weevil as

affected by host factors. D.R. (Ross) Macdonald moved to PFC from the Fredericton lab

in 1968 to become Section Head, Forest Entomology. In 1969 he became Program

Director for Forest Protection. After serving as Director of the Forest Protection Branch in

Ottawa, Mr. Macdonald became Director General, Pacific and Yukon Region (1980-87),

Canadian Forest Service (CFS), a post from which he retired. Mr. Macdonald’s research

experience and expertise included spruce budworm ecology and the effects of spray

operations on budworm natural enemies. Dr. R. (Roy)F. Shepherd moved to Victoria

from the Edmonton lab of CFS in 1969 to undertake researches on the population biology

and management of defoliating insects. This research has lead to development of sampling

methods for several defoliators, development of pest management methods for the

Douglas-fir tussock moth and the black army cutworm, monitoring systems for endemic

populations with pheromone traps, and prevention of outbreaks through early treatment

with nuclear polyhedrosis virus.

Dr. H. (Henry) A. Moeck joined the bark beetle research group in Victoria from the

Forest Products Laboratory in Vancouver in 1970. His field of research involved primary

attraction in bark and ambrosia beetles and assessment of native and exotic natural

enemies of bark beetles as potential biocontrol agents. Dr. S. (Steve) IInytzky joined the

entomology group at PFC in the early 1970s. His main research concerned the use of

pesticides in insect control. He developed bioassays of insecticide residues in soils and

investigated phytotoxicity of pesticides to forest seeds. C. (Cliff) E. Brown moved to PFC

from Ottawa in 1972 to become a Program Manager and Deputy Director, positions from

which he retired. The focus of Mr. Brown’s earlier work concerned the development of

automated recording of FIDS data at regional and national levels. Drs. M. (Malcolm) D.

Shrimpton (tree physiologist), H.S. (Stu) Whitney (plant pathologist) and L. (Les)

Safranyik (insect ecologist) were transferred to Victoria from the Edmonton lab of CFS in

1972 in order to concentrate bark beetle research in CFS at the Victoria lab as these insects

continued to be the most destructive in the forests of British Columbia. Based largely on

previous research in the East Kootenays by this team and colleagues from Alberta, they

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 165

produced two milestone publications on the mountain pine beetle in 1974: “An

interpretation of the interaction between lodgepole pine and the mountain pine beetle with

its associated blue stain fungi in Western Canada” and “Management of lodgepole pine to

reduce losses from the mountain pine beetle” The first paper describes the nature and

affects of the three-way interactions among the host, beetles and fungi and their role

determining the course of infestations. The second paper, intended for practicing foresters

and forest health specialists, emphasizes that in order to effectively cope with the beetle

the focus of management should be logepole pine and not the beetle. This publication was

an all time “best seller” in Canada and the United States; two printings of a couple

thousand copies each are sold out. Other major publications followed on the population

dynamics of the spruce beetle. Dr. Safranyik’s current research involves a population

dynamics model of the mountain pine beetle, competitive exclusion of the mountain pine

beetle, and spruce beetle ecology. Dr. A. (Al) J. Thomson joined PFC in the late 1970s.

An insect ecologist but not formally part of the entomology research group, Dr. Thomson

does much work of entomological nature such as a landscape-level model for mountain

pine beetle, collaborative work on bark beetle population quality with Dr. Sahota and a

mountain pine beetle model with Dr. Safranyik. H. (Howard) A. Tripp succeeded Dr.

Silver as FIDS Survey Head in 1973. His interests were biological control and seed and

cone insects. Dr. C.D. (Doug) F. Miller moved to PFC from Ottawa to become Research

Program Manager in 1978.His entomological work was done in agriculture concerning the

taxonomy of ants, wasps and parasitic insects. Dr. Miller has published several

monographs on the classification of these insects.

Dr. G. (Gordon) E. Miller joined PFC to take over seed and cone insect research

following the retirement of A.F. Hedlin in 1980. Dr. Miller’s research work was centered

on the population ecology and management of cone and seed insects and produced over 30

scientific and technical papers on these subjects. Since 1986 he pursued a career in

management and is currently Director of Research for CFS in Ottawa. Dr. L (Imre) S.

Otvos moved to PFC from the Newfoundland lab of CFS inl1981. His main work at PFC

centers on the use of natural enemies (predators, parasitoids, fungi, bacteria and viruses)

for controlling defoliating insects. Some major accomplishments include assessment of the

potential of introduced parasitoids and fungal pathogens for the control of the hemlock

looper; pioneering the concept of creating epizootics of naturally occurring pathogens to

control defoliators, and co-developing with Dr. R.F. Shepherd a management system for

Douglas-fir tussock moth. Dr. R. (Rene) I. Alfaro joined the FIDS program in 1980 and

was assigned to entomology research full time a few years later. The main foci of his

research are quantification of damage caused by forest pests, management and ecology of

the spruce weevil with emphasis on the nature of host resistance and_host-insect

interaction. His comprehensive research on insect impacts has enabled private and

provincial pest management agencies to measure potential losses and to justify control

programs. Continuing work on weevil-host interactions resulted in numerous scientific and

technical papers concerning the nature and effects of host resistance, and the dynamics of

intensification and spread of infestations at the tree, stand and landscape levels. Dr. M.

(Mike) A. Hulme joined PFC from the Eastern Forest Product Laboratory in 1982. The

main area of his work involves biological control. Dr. Hulme co-edited the book entitled

“Biological control programs against insects and weeds in Canada 1969-1980”. Other

accomplishments include demonstration of pheromone-based mating disruption § in

Douglas-fir tussock moths, a first in this field, and showing that host phenology has a

major affect on resistance to the sitka spruce weevil. Dr. 7. (Terry) L. Shore transferred

from the FIDS to bark beetle research in 1985 following the retirement of Dr. L.H.

McMullen. His main areas of research are development of hazard-risk rating systems and

decision support systems to enable forest managers to develop effective and efficient

166 J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

programs of managing destructive bark beetles. A system for rating lodgepole pine

susceptibility and risk of loss from mountain pine beetle developed by Drs. Shore and

Safranyik is included in the bark beetle management guidelines section of the Forest

Practices Code for the province, and is being widely used by forest protection staff in

industry and government in BC as well as in neighbouring states in the USA. Dr. A.

(Hugh) J. Barclay, an insect ecologist, joined the growth and yield program at PFC in

1982. Although his research assignments were, and continue to be, in a field other than

entomology, he has authored and coauthored a number of scientific papers on

entomological subjects. Significant contributions include the mathematical treatments of

sterile males, inundative releases of natural enemies, for controlling insects, and

contribution to the development of interactive models of spruce beetle and mountain pine

beetle population dynamics. Dr. L. (Lee) M. Humble joined FIDS in 1985 as an insect

taxonomist. In addition to the taxonomic work, his main areas of research include insect

biodiversity, specifically the diversity of canopy-dwelling insects in the boreal forest, and

the establishment and biology of introduced insects.

Following program reorganization in CFS during the mid-1990s two entomologists

transferred to PFC from other CFS establishments: Dr. V. (Vince) G. Nealis from Sault

Ste. Marie and Dr. A. (Allan) L.C. Carroll from St. John’s, Newfoundland. Dr. Nealis is

developing a program on the population ecology of spruce budworms in British Columbia.

The focus of Dr. Carroll’s current work involves investigations of the ecology of mountain

pine beetle populations in the endemic state.

Technical support

The contributions of technical support staff to all phases of research and development

are highly important and impressive. Some technicians, owing to special skills and or

experience gained on the job, had their own research projects which they conducted semi-

independently from the research officer in charge. For example, W. (Bill) W. Niyholt has

made substantial contributions to ambrosia beetle management and pioneered the use of

pine oil for controlling bark and ambrosia beetles and, with H.A. Richmond, water-misting

of log decks in dry-land sorts to reduce ambrosia beetle damage. Some others who worked

in similar manner are T. (Tom) G. Gray on pheromone-based monitoring of defoliators and

R. (Bob) Duncan in insect taxonomy. Still others co-authored an impressive list of

scientific and technical papers.

Trends in research

Of necessity, much of the research conducted during the first 25 years or so involved

identification, taxonomic description, geographic distribution, and exploratory work on the

biology and control of injurious forest insects. Due to shortage of manpower and expertise,

often a single scientist was assigned to work on several problem areas involving several

insect species in different taxa. With an increase in the number of scientists assigned to

forest entomology, research became specialised. Scientists normally worked in a single

field such as physiology, population ecology and often with one or a few related species

such as the budworms. There was an emphasis on studying insect populations under

outbreak conditions. Research projects, however, were still mainly headed up by single

scientists. This general trend continued to the beginning of the 1970s. From this time on,

gradually there was a change from studies of epidemics to studies of factors affecting

populations in the endemic state. As well, there was more emphasis on insect management

through cultural practices and natural enemies. Multidiscipline team approaches to tackling

difficult problems became more prevalent. These trends continue today but with greater

emphasis on development of practical decision support systems that are based on a solid

ecological foundation. There is increased need for understanding the interactions of insects

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 167

with their host trees and the host environment at the landscape level and the consequences

of natural and man-made disturbances, including insect management actions, on

biodiversity and sustainability of forest management.

REFERENCES

Anon. 1978a. Ministry of Forests Act of British Columbia

Anon. 1978b. Forest Act of British Columbia

Anon. 2001. Forest Practices Code of BC Act.

Graham. K. 1963. Concepts of Forest Entomology. Reinhold Pub. Corp., New York.

Hopping, R. 1921. The Control of Bark-Beetle Outbreaks in British Columbia. Dominion of Canada, Dept.

of Agriculture, Entomological Branch. Circular No. 15. 15 pp.

Hopping, G.R. and W.G. Mathers. 1945. Observations on outbreaks and control of the mountain pine

beetle in the lodgepole pine stands of western Canada. The Forestry Chronicle, June, 1945: 9-11.

Pearse, P.H. 1976. Timber Rights and Forest Policy in British Columbia. Report of the Royal Commission

on Forest Resources. 395 pp.tappendices.

Richmond, H.A. 1983. Forever Green: the Story of One of Canada’s Foremost Foresters. Oolichan Books,

Lantzville, BC. 203 pp.

Riegert, W.P. 1991. Entomologists of British Columbia. Friesen Printers, Altona, Manitoba. 90 pp.

Swaine, J.M. 1914. Forest Insect conditions in British Columbia. A Preliminary Survey. Dominion of

Canada, Dept. of Agriculture, Division of Entomology. Entomological Bulletin No. 7. 41 pp.

Swaine, J.M. 1918. Canadian bark beetles. Part 2. A preliminary classification, with an account of habits

and means of control. Dominion of Canada, Dept. of Agriculture, Division of Entomology. Technical

Bulletin No. 14. 143 pp.

Turnbull, A.L. and D.A. Chant. 1961.The practice and theory of biological control in Canada. Canadian

Journal of Zoology 39: 697-653.

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 169

II. The Forest Insect and Disease Survey

in the Pacific Region

A. VAN SICKLE, R.L. FIDDICK and C.S. WOOD

NATURAL RESOURCES CANADA, CANADIAN FOREST SERVICE, PACIFIC

FORESTRY CENTRE, 506 WEST BURNSIDE ROAD, VICTORIA, BC, CANADA V8Z IM5

The combined Forest Insect and Disease Survey (FIDS) originated with the 1962

unification of the pre-existing Forest Insect Survey and the Forest Disease Survey. FIDS

was an early national network within the Canadian Forestry Service, and its predecessors,

to detect and monitor insect and disease conditions in Canada's forests. Responsibilities

among the six regional units, such as the British Columbia and Yukon unit, included

monitoring forest pest conditions, keeping records to support quarantines and make

predictions, supporting forestry research with records, and disease (herbaria) and insect

collections, testing and developing survey techniques, and providing advice on insect and

disease conditions.

FIDS and pest surveys were an early and integral part of the Canadian Forest Service in

its various forms and departmental associations. Dr. J.M. Swaine was appointed as the first

full time forest entomologist in Canadian government service in 1912. He conducted

numerous inspections (surveys) and collecting trips in western Canada and published

"Canadian Bark Beetles" in 1918.

In 1922 the Canadian government established an insect laboratory at Vernon to

investigate forest insect problems in the BC interior, the Rocky Mountain National Parks

and the east slopes of the Rockies. The four staff sharing an office in the Vernon

courthouse included Ralph and George Hopping, Bill Mathers and Hec Richmond. Initially

efforts focused on extensive bark beetle infestations in pine stands in south central BC and

in Banff National Park, where from 1941-1943 more than 27,000 infested trees were cut

and burned to prevent an outbreak of the mountain pine beetle from developing to

epidemic proportions. Although the Vernon establishment was made a_ sub-lab

administered from the Victoria centre in 1945, by 1951 the 18 staff were accommodated in

a new office building on British Columbia Forest Service land on Pleasant Valley Road.

An insectary and large garage-shop was added by 1954. When the Vernon lab was closed

and staff transferred to Victoria in 1970, the facilities were sold to the British Columbia

Forest Service for $1.00

A sub-laboratory for insect and disease studies in the coastal forests was opened in

Vancouver in 1925 in conjunction with the Forest Products lab.

In 1928, Dr. J.M. Swaine strongly urged the establishment of an "insect intelligence

service" for a survey of forest insects because of the severe outbreaks of budworm, sawfly

and bark beetles. This was supported by representatives of forest industries through the

Canadian Pulp and Paper Association and the Canadian Society of Forest Engineers. In

1931, The Division of Forest Insects of the Entomological Branch of the Department of

Agriculture organized the "Forest Insect Intelligence Service" A number of circulars giving

popular accounts of the principal forest pests were prepared. These circulars, together with

questionnaires were sent to industrial organizations and forest services, with the request

that they be filled out by forest workers. In 1936 the Forest Insect Survey was established

as a permanent, independent unit and 528 insect samples and records from eastern Canada

reached Ottawa for identification and recording. In 1937 forest insect surveys from Vernon

commenced as part of a survey which had national scope by 1939. To process the

collections being sent to Vernon a Forest Insect Survey insectary was built nearby at

170 J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

Trinity Valley in 1937. Mind you, this was only after obtaining direct approval from Dr.

J.J. de Gryse, Chief, Forest Insect Investigations in Ottawa (and workers today think they

are tightly controlled). Labor was to be at 70 cents an hour with total costs not to exceed

$300. With closure of the Vernon lab, and following some vandalism and a fire in 1968,

the Trinity Valley site was turned over to the Department of Education about 1971 for

outdoor studies.

The nation wide survey project was administered on a regional basis, coordinated

through a headquarters at Ottawa, initially with the following objectives:

1. Conduct a year-to-year survey of the status of potentially destructive insect species.

2. Accumulate information relating to the many thousands of species affecting forest trees:

the characteristics of their attacks on the trees;

their distribution throughout Canada;

the tree species attacked in different parts of their range;

the parasites, predators and diseases which attack the insects;

the relation of the development of destructive insect populations to forest

composition, age and other environmental factors.

These objectives were updated periodically, to include forest disease surveys in 1952,

and damage appraisal, survey methodology and remote sensing in the Pacific Region, in

1966.

A permanent forest insect centre was established in 1940 in Victoria firstly in the "old,

old" Post Office building (now the Customs and Immigration Building on Government St.

and Wharf St.). About this time two wood-walled tent-covered frames at Lake

Cowichan/Mesachie Lake Experimental Station were refurbished by Drs. Prebble and

Graham (entomology) and Dr. Buckland (pathology). These were upgraded, although still

far from being luxurious, by Survey ranger staff from 1947-1949. They along with a 1960s

laboratory (now a meeting centre) served as a field station for research by Federal staff

until 1983 when they were turned over to the British Columbia Forest Service.

After World War II permanent ranger staffs were assigned to the Forest Insect Survey

with their first offices in Victoria in the Central Building on View Street before moving to

the old post office. R.L. (Lew) Fiddick was taken on staff in the spring of 1945 as the first

federal insect ranger in the Pacific Region. The first assignment under the direction of Dr.

M.L. Prebble was an assessment of a western black-headed budworm outbreak on north

Vancouver Island in the Holberg Inlet-Port Alice area. A spray program had been

organized, but a very labor intensive egg survey on numerous, large, hand-felled trees

showed the infestation had collapsed due to natural causes. Over the next 30 years FIDS

was to be involved in forecasting and assessing population levels, forest damage and

control efficacy for more than 25 infestations of numerous different defoliators (see

Appendix). It contributed to the development of aerial spray technology applicable to the

unique and challenging conditions of BC. Effective dosage levels were progressively

decreased and the more acceptable biological insecticides were introduced and tested.

By 1949 the Department of Agriculture was advertising for additional Forest Insect

Rangers, Grade | at the annual salary of $2160-$2460. Expense account allowances were:

breakfast 70 cents, lunch 85 cents, supper $1.15 = $2.70/diem. The midnight ferry from

Victoria to Vancouver was fare $3.50, truck $6.00 berth $4.50. At its peak a total of 16

Forest Insect Survey Rangers were assigned to individual districts throughout BC.

In 1948, to undertake surveys of the more than 11,000 km of shore line at a time when

there were virtually no roads, an 18 m (60 foot) vessel was purchased from War Assets and

converted to a floating laboratory, the JM. Swaine. Cramped quarters existed for the

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 171

skipper, a cook, engineer and two or three survey rangers or assistants. The vessel served

the coastal areas until 1953 when it was sold and replaced by the smaller two-man vessel,

the Forest Biologist. With somewhat improved road access and increased usage of aircraft

the Forest Biologist was last used in 1968, although smaller one-person trailered boats

continued to be used for sampling on smaller islands and along the lower coast. In 1968, a

new ranger hired from the second graduating class of British Columbia Institute of

Technology forest technicians, Peter Koot, had his first "sink or swim" summer surveying

from Jervis Inlet to Smith Inlet using "The Okalla Queen". This was a V6 powered

plywood, 6.4 m (21 foot) cabin cruiser/tub built by the "very experienced" boat builders

from Okalla Prison. It would only get up and plane with a nearly empty gas tank, very little

food on board, no water in the bilges (a rarity because it always leaked) and preferably

without the student assistant.

Also in 1948 a Forest Insect Survey insectary and rearing facility in Langford was

completed by the versatile Survey staff. In 1965 with the new Pacific Forest Research

Centre research facilities in Victoria the old insectary building was turned over to the

municipality and is now the offices of the Capital Regional District and part of Mill Hill

Regional Park.

Between 1950 and 1963 a network of seasonal Forest Insect Survey field stations was

built including Kye Bay (1950), Cultus Lake, Lakelse Lake (1951), New Denver, Christina

Lake (1953), Wasa Lake (1954), Williams Lake (1952), Prince George, and Babine Lake.

In following years additional stations were moved to or acquired at Terrace (1959),

Kamloops, Agassiz, Powell River (1963), Smithers and Summerland. With the long-term

and conspicuous involvement in many communities throughout the province the Survey

became one of the most visible parts of the Canadian Forest Service and maintained a

reputation for consistent service to the forest community. The cabins greatly reduced

accommodation costs during the 4 decades of service and greatly appreciated in value

compared to the average cost of construction (Cultus Lake $3418; Kye Bay $2937; Lakelse

Lake $3855; Williams Lake $4045; Wasa Lake $4359; Babine Lake $5804 - costs were

said to be higher here because of local shortages due to the ongoing construction of the

Kitimat aluminum plant). Minimizing operating costs is a regular government mantra, but

consider the extent in the 1950s: in July 1957, B.M. McGugan, Co-ordinator, FIDS in

Ottawa wrote Ray Lejeune, Officer-in-charge, Forest Biology Laboratory, Victoria, "policy

regarding ranger cabin furnishings will not allow the purchase of refrigerators". And in

1959, $4.40 per month for a phone was considered too high to consider in the budget.

Remember at this time the paved roads ended at Clinton, the Hope-Princeton Highway

was just being opened and the Rogers Pass and Kaslo-Salmo highways didn't exist. Initially

the Survey organization "inherited" army surplus 4-wheel-drive vehicles called Heavy

Utility Personnel Carriers (HUP's). Only slightly suitable for rough overland travel at slow

speeds they were clumsy, noisy, dusty, right-hand drive and subject to break-down. When

eventually new 1950 Ford vans were delivered to Victoria they didn't have heaters, Ottawa

considered them unnecessary.

A Forest Disease Survey was formally organized in 1952 to include a survey of tree

diseases. For both the Forest Insect and the Forest Disease Survey, a standard punch-card

system of recording and sorting insect and disease records was established across Canada

("as many as 12 columns simultaneously and 400 cards per minute" were processed using

Remington Rand sorting machines).

Annual forest insect and disease surveys in the Yukon were started in 1952, although a

special survey for mosquitoes had been conducted in 1948 with an Ottawa-based scientist

and support staff from Victoria (they certainly didn't collect them all!). The annual Yukon

surveys by FIDS from the Pacific Forest Research Centre continued until 1965, and then

172 J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

again from 1974 to 1996. From 1966 to 1972 the Yukon surveys were conducted from the

Northern Forestry Centre in Edmonton. Initially an older house trailer in Whitehorse was

used during the Yukon surveys before being replaced by a roll-up, sectional camper and

then a modern hard sided truck mounted camper. One year, the FIDS ranger sleeping in the

camper awoke suddenly to find a mother and two cub bear faces pressed against the

window only inches away from his own. Being a sectional camper, one good shove would

have separated the sections; fortunately the bears climbed down from the hood of the truck

and departed, probably followed quite quickly by the ranger. Another time a bear climbed

into the back of the truck while the ranger was away collecting, and then... but there are so

many bear stories.

Although the first sketching of infestation boundaries from an aircraft was published as

early as 1920, and periodic mapping was done for major outbreaks, it wasn't until the early

1950s that aircraft availability and cost was reasonable enough that organized annual

overview flights and mapping of the major defoliators and bark beetles was initiated on an

annual basis. Federal Government austerity always seemed to exist and there was seldom

much money for aerial surveys. Acknowledgement is made of the considerable provision of

flying time and cooperation by the British Columbia Forest Service and forest industry.

The considerable amount of flying time provided to FIDS from other agencies annually for

many years speaks well for the negotiating skills of the ranger staff and for the value of the

information they provided. Without the many excellent contacts and cooperation

established and maintained by the rangers the several decades of aerial detection and

damage records would not have been as extensive.

Cooperation was not limited to just flying time, but included information, lodgings,

assistance and loan of equipment. One courtesy provided in 1958 to ranger, Norm

Alexander in the remote West Prince Rupert District was the loan of a large box "portable"

radio which required that the antenna be thrown up over a branch then back away the 30.5

m (100 foot) length to make your call (hardly today's cellular phone).

Aircraft and pilot reliability was a somewhat different matter at times. e.g. One pilot

refused to fly about 300-500 m above the ground necessary to distinguish and map damage.

He was dubbed "sky blue Lou" and probably fortunately, was always on days-off when

other FIDS flights were scheduled. On the other hand there was the pilot who, after

mapping mountain pine beetle damage, bypassed the refueling facilities at McBride and

headed for Prince George. The FIDS ranger made a few enquiries about running out of fuel

and sure enough over Purden Lake the engine sputtered. Abrupt rocking of the plane kept

the engine alive for a few moments, but not for long and elevation was being lost. Gliding

silently over Tabor Lake, boaters were seen making hasty retreats to the shore and even

into the lake. Fortunately the glide carried far enough for a relatively smooth landing in a

hay field just beyond the lake. A different plane and pilot, but the same FIDS ranger

finished mapping later that week and continued doing so for years.

Other enjoyable(?), memorable flights: En route to Sandspit, dark oily spots started

appearing on the window and increased until visibility was nearly nil. Upon landing, smoke

curled up from the engine cowling and the engine bust into flame. Luckily the pilot

anticipated this (had it happened before?) and doused the flames with an extinguisher.

Another incident occurred on an early 1970s flight made from Campbell River in a Beaver

on floats. After dodging the many anglers in the harbor and just about to lift off, the engine

died, causing a near nose dive into the chuck. These were the joys of a new pilot on a plane

just brought back to Canada from the Peruvian Air force with an instrument panel still in

Spanish and the gas tank switches opposite to those of other Beavers (the pilot had

switched to an empty instead of full tank). Most rangers had hair-raising experiences of

flying in marginal weather using fish boats and lighthouses for navigation, float-plane

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 173

landings hitting rough seas, logs or even basking sharks, and very tight turns in narrow or

dead end (not a nice term) valleys. Invariably, they would comment "If it had been any

other type of plane but a noisy Beaver, it would not have continued flying."

In 1960 the Forest Biology Division, Science Service, Department of Agriculture was

joined with the Forestry Branch, Department of Northern Affairs and Natural Resources to

form the Department of Forestry.

The Forest Insect Survey and the Forest Disease Survey were unified in 1962 as the

Forest Insect and Disease Survey (FIDS).

While the Honourable J.R. Nicholson, the Minister of Forestry may have laid the

cornerstone in August 1963 for the new Forest Research Laboratory on Burnside, it was

FIDS rangers in 1957 who laid the more important first culvert and driveway for access to

the site. This included hauling gravel from Goldstream in the back of the panel trucks

because of government spending constraints. Guess who got the most press!

On February 15, 1965 the new Pacific Forest Research Centre on Burnside Road was

opened, and for a short while the entire coastal unit of FIDS were together along with the

herbarium and insectary. Then, after the 1969 austerity review, the Vernon lab was closed

and all projects and personnel were transferred to Victoria (10 transferred, 2 quit and 2

positions were terminated). Along with staff transferred from the Calgary/Edmonton lab the

Burnside lab became crowded and offices and labs were created for FIDS in the old

headerhouse which had been constructed in 1957-1958 along with a separate greenhouse.

(The herbarium cabinets stood just about where the Christmas luncheon is tabled).

Although slightly separated from the main lab it was only a wet run away during coffee

breaks and FIDS staff maintained a strong team unity (only the screams from the serious

table tennis players in the silviculture group's annex next door shattered the noon-hour

peace). With the addition of the "new wing" in 1985, the herbarium and all FIDS staff

returned to the third floor again in association with the insectary. The new greenhouses

were then built attached to the headerhouse.

Forest insect and disease punch card records were computerized in 1967 and new

collections were recorded on a revised enclosure slip. Data processing was centralized in

Ottawa.

In 1975 the FIDS unit at PFC led a national task force on Damage Appraisal and Pest-

Caused Losses. The approaches developed were the basis for loss estimates contributed to

Statistics Canada for the next couple of decades and used widely by the British Columbia

Forest Service and others.

In 1979 there was another A-Base Review and a task force was struck by Canadian

Forest Service Headquarters to review the FIDS program. Based on interviews and

correspondence with over 150 individuals in federal departments, provincial agencies and

the forest industry, the task force concluded: "there are sufficient and compelling reasons

why the FIDS should be maintained as a national program". It also recognized that the

transfer of FIDS from Agriculture to Forestry without provision for a mandate to replace

the Plant Quarantine Act, and the de-emphasis of the national aspects of the program in the

1969 reorganization, required action to readopt and reaffirm the original strong rationale

for a FIDS to provide essential data input to federal forestry programs and to federal policy

and decision making. It was recognized that the demand for pest information was certain to

grow at an increasing rate over the next decade, but periodic funding reviews would

continue to target programs as large as the national Forest Insect and Disease Survey.

In 1984 FIDS at the Pacific Forest Research Centre began developing a computerized

mapping and analysis geographic information system (GIS). It was up-graded in 1992

when the system was standardized nationally. GIS is now commonly and nationally utilized

and the historical distribution and infestation records are available on the internet.

174 J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

The Forest Insect and Disease Survey celebrated 50 years of service in 1986. Over this

period of time and beyond, the motivation and self-reliant nature of FIDS staff was key to

many accomplishments. They undertook annual in-service training programs most winters

and tested and embraced new tools. Funds were found for 2-way radios in 1983 after Leo

Unger walked more than 33 kms from a mud hole. Pheromone trap monitoring for gypsy

moth detection was used since 1978 and pheromone monitoring systems for numerous

other forest insects were tested and calibrated. Satellite tracking geographic positional

systems and electronic field data recorders were added early in the 1990s.

From 1985-1996, FIDS provided information, undertook special surveys, developed

and tested protocols and served on Science/Policy panels concerning the pinewood

nematode and its effect on the export of Canadian lumber.

In 1995, after (yet another) program review and reorganization of the Canadian Forest

Service, which included a 30% cut in resources, the decision to discontinue annual,

operational surveys of important forest pests on a national basis was announced. Several

provinces, including BC, felt that they could not politically accept the “off-loading” of

what had been a Federal activity, or did not have the resources to continue the surveys.

Subsequently the FIDS units were disbanded through early retirements, other packaged

departures, and transfers to the various research networks that were established within the

Canadian Forest Service.

PERSONNEL

FIDS has been an early and integral part of the history of the Canadian Forest Service

in Canada and in BC/Yukon. Over a period spanning almost 6 decades many staff gained

and shared an expertise and knowledge of the forests and its diseases, insects and natural

control factors. More than 70 men and one woman served as Forest Insect and Disease

Survey Rangers. Some remained only for a year or two but for many it was a long and

dedicated career. Ranger staff in the Pacific Region with more than 25 years service

included: Stan Allen, Dick Andrews, Cliff Cottrell, Lew Fiddick (the most senior with

more than 37 years service, first as an insect ranger and then Chief Ranger), Jim Grant,

Colin Wood, Roly Wood, and Bob Erickson, Peter Koot and Leo Unger who are still

contributing as Forest Health network technicians. More than 50 other scientists and

technicians have worked with FIDS in the herbarium, insectary, damage appraisal and

methodology studies. Those with more than 25 years service included: George Brown, Don

Collis, Al Dawson, Dave Evans, Ab Foster, John Harris, Daphyne Lowe, Alec Molnar,

Erika Pass, Doug Ross, Dave Ruppel, Allan Van Sickle, Emil Wegwitz and Wolf Ziller.

Numerous others contributed and received a good foundation in FIDS before

transferring to other projects in the Canadian Forest Service or to other aspects of forest

pest management including (but not limited to): Norm Alexander, Rene Alfaro, Dennis

Beddows, Clare Farris, Brenda Callan, Dennis Clarke, Don Doidge, Bob Duncan, Lee

Humble, Rich Hunt, Ernie Morris, Doug Ruth, Terry Shore, Tom Silver, and Tad Woods.

In addition, many casual employees and students, while working with FIDS to earn funds

for further education, also gained an increased awareness of forest insects, diseases or

geographic information systems, and the experience of federal government employment.

FIDS is indebted to the secretarial staff (Evelyn Andrews, Nancy Mason, B. Sugden

and T. Gabriel at Vernon; Pat McLean, Heather Murchison, Francis Douglas, Susan

Ticknor, and Joan Strobbe at Victoria), who patiently deciphered near incomprehensible

telephone messages and hand writing; and to the many other technicians and scientists,

regionally, nationally and internationally who worked closely with, and contributed greatly

to the many accomplishments of FIDS.

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 175

MAJOR ACCOMPLISHMENTS OF FIDS

For more than 5 decades Forest Insect and Disease Survey staff provided expertise and

a strong liaison for the Canadian Forest Service with the British Columbia Forest Service,

the Yukon Territory, as well as with forest industry, universities and technical schools,

provincial and federal parks, the public and other agencies. This was a time of growing

awareness and concern about forest pests, their life cycles and control factors, both natural

and chemical.

From the 1940s to the 1980s, FIDS staff undertook, often in cooperation with the

Council of Forest Industries, or participated with research staff in the assessment and

monitoring of more than 25 forest insect control projects. In numerous additional cases,

careful assessments of population trends and natural control factors indicated that direct

control was not required.

FIDS developed and implemented practical sampling and survey methods and

frequently provided collections, information and assistance to other scientists across

Canada and internationally.

The FIDS collections and associated data established the extensive data base of more

than half a million records of forest diseases, insects and beneficial natural control agents

from the forests of the Pacific Region of Canada. The curated permanent reference

collection of more than 66,000 insect and 35,300 disease specimens is critical for accurate

identification of native and exotic pests. An extensive photo and color slide collection was

also assembled. Some of this information has recently been made available on the internet.

It has also provided the basis for some biodiversity studies.

Many assessments of the natural biological control factors of forest insects were also

conducted by FIDS. A data base of more than 20,000 records was compiled from

collections and insectary rearings to better understand and predict the effects of natural

occurring parasites and disease on insect infestations. Biological control experiments

started as early as 1931 when parasites from Japan were caged with hemlock sawfly from

the Queen Charlotte Islands. As well as monitoring the larch casebearer from its first

detection in BC in 1966, FIDS initiated and participated in parasite introductions and

assessments from 1969 to 1984. Other successful parasite releases and introductions were

made for balsam woolly adelgid, the winter moth, larch sawfly and apple ermine moth.

In 1975 FIDS led a national task force on damage appraisal and pest-caused losses,

developing the approach and providing estimates of forest insect and disease losses in the

nation's forests for reporting by Statistics Canada for the following decades. These

quantifications increased awareness of the major impacts of forest insects and diseases as

well as identified areas needing further study and support. Damage appraisal, remote

sensing of pest-caused damage, and statistically based sampling studies started first in

FIDS before becoming separate studies.

Extensive biological surveys (insects, diseases, parasites and predators) were conducted

up to 1969, after which emphasis was more on commercially important pests with

increasing attention to introduced insects and diseases of potential risk to Canada's forests.

An increased level of awareness and concern for the latter continues. FIDS provided both

an annual and an historical perspective of forest pest conditions throughout BC and the

Yukon. This was well documented in a great many timely reports and publications (even

videos) and through participation in regional, national and international meetings.

FIDS first implemented a geographic information system (GIS) for the maintenance and

reporting of insect infestations and disease distributions in 1984. Portions of the historical

aerial survey outbreak maps and disease distributions are available on the world wide web.

176 J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

An informative and popular Forest Pest Leaflet series describing the life cycles, damage

and activities of more than 60 common forest insect and diseases was created and

maintained by FIDS. These were also presented in PFC's first CD release (HFOREST -

Hypermedia Forest Insect and Disease Knowledge Base and Diagnosis).

APPENDIX

Early aerial control trials of forest defoliators in BC.

Date Insect Location/area Insecticide

1929 western hemlock looper Burrard Inlet/45ac calcium arsenate

1930 — western hemlock looper + Stanley Park/800ac calcium arsenate

black-headed budworm

1930 western hemlock looper Seymour Creek/800ac calcium arsenate

1946 — western hemlock looper Nitinat DDT

1948 — western false hemlock looper Windermere Valley DDT

/11,200ac

(First time in Canada to use non-fixed wing aircraft, Bell 47 helicopter, in an

operational forest pest control program)

1956 — black-headed budworm Pt. McNeill/360ac DDT

1956 phantom hemlock looper Burnaby, Central Pk DDT

/200ac

1957 phantom hemlock looper Burnaby, Central Pk DDT

/200ac

1957 phantom hemlock looper New Westminster DDT

Queens Pk /75ac

1957 black-headed budworm Pt. Alice-Pt Hardy DDT

/156,000ac

1959 western hemlock looper + Stanley Pk/550 ac DDT

greenstriped forest looper

1960 _ black-headed budworm QCI Moresby Is DDT, Bt

/160,000ac

1960 — saddleback looper Kitimat/1,800ac DDT

1961 saddleback looper Kitimat /9,800ac DDT, Dibrom,

Phosphamidon, Bt

1961 pine butterfly Cameron Lake/1,500ac DDT

1962 Douglas-fir tussock moth Okanagan Valley/160 ac DDT, malathion,

NPV

1964 — western hemlock looper Enderby/50ac Phosphamidon

1964 _ greenstriped forest looper QCI Pt Clements/1,600ac Phosphamidon

1965 hemlock needleminer Holberg/1,000ac Phosphamidon,

Dimethoate

1973 western false hemlock looper Salmon Arm/400 ac Dipel

1973 black-headed budworm Pt. Alice/28,000 ac Fenitrothion

1974 Douglas-fir tussock moth Kamloops/25 ac NPV

1974 ~ western false hemlock looper Chase/120 ac Dipel, juvenile

hormone, Zoecon

1975 Douglas-fir tussock moth + Kamloops/3 1,000 ac Dipel, Orthene, Bt,

western false hemlock looper NPV

1976 Douglas-fir tussock moth Kamloops/20,000 ac Orthene, Bt

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 177

III. The Vernon Laboratory and federal entomology

in British Columbia

RICHARD A. RAJALA

DEPARTMENT OF HISTORY, UNIVERSITY OF VICTORIA,

P.O. BOX 3045, VICTORIA, BC, CANADA V8W 3P4

Over its fifty years of existence the personnel of the Vernon forest entomology

laboratory conducted scientific research on B.C.’s insect species, battled epidemics that

threatened commercial timber, and monitored their populations. Between 1920 and 1970,

first as part of the Department of Agriculture and more recently within the Canadian

Forestry Service, the facility provided tangible evidence of the federal government’s

interest in the provincial forest economy. Prior to the 1930 transfer of natural resources to

the provinces Ottawa’s stake in the Railway Belt provided sufficient reason for such a

presence, perhaps, but the laboratory survived after B.C. claimed complete jurisdiction

over timberlands in the province. Transcending provincial boundaries, forest insect

problems were an appropriate area of Dominion scientific inquiry.

An early twentieth century bark beetle epidemic in the southern interior created the

motivation for Ottawa and Victoria to cooperate in control projects that operated each

spring and summer during the 1920s. Ralph Hopping directed that effort, and stayed on to

head the laboratory’s small staff until passing the reins to his son George in 1938. The

facility remained B.C.’s major centre of forest entomology until losing its research

orientation to a new federal installation in Victoria in the 1950s, then closed entirely in

1970. Ironically, soaring interior bark beetle populations have lowered the value of vast

stretches of timber in subsequent decades.

Forest entomology in Canada began to emerge from its amateur origins in 1909, with

the hiring of C. Gordon Hewitt as Dominion Entomologist. Hewitt, in a refrain

entomologists would repeat often in coming decades, observed that the depradations of

insects received much less attention than fire in Canadian conservation circles. Some

research had been conducted on larch sawfly and spruce budworm outbreaks, but their

significance paled in comparison to the damage caused by several species of bark beetles

which had yet to receive any scientific attention. The first step in correcting this situation

came with the 1912 appointment of J.M. Swaine, a graduate of Cornell and recognized

North American bark beetle authority, as Assistant Entomologist for the study of forest

insects in the Department of Agriculture’s Entomology Branch. Swain took charge of the

new Division of Forest Insects, and immediately undertook a study of bark beetle

outbreaks in Canadian forests. '

A cooperative agreement between the Dominion Department of Agriculture and the

B.C. Forest Branch took Swaine to the west coast in the summer of 1913 for a survey of

forest insect damage. Swaine covered the Kootenay, Okanagan, Similkameen, lower coast,

and Vancouver Island regions. His preliminary investigation found the coastal forests

relatively free of extensive outbreaks, but in the southern interior pine beetles were

'MLL. Prebble, “Forest Entomology,” The Canadian Entomologist 88 (July 1950), p. 351;

D.R. Wallace, “Forest Entomology or Entomology in the Forest? Canadian Research and

Development,” Forestry Chronicle 66 (Apr. 1990), p. 120 [hereafter FC]; “Will Study

Forest Insects,” Canadian Forestry Journal 8 (Jan. — Feb. 1912), p. 26 [hereafter CFU]; C.

Gordon Hewitt, “Investigations on Forest Insects, and Forest Protection,” CFU 8 (Mar. —

Apr. 1912), p. 36.

178 J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

inflicting serious losses to yellow pine stands. Conditions were particularly acute in the

Princeton area, where an outbreak had killed much valuable timber. “The clumps of “red

tops” may be distinguishing upon the mountain-side and in the valleys for many miles,” he

noted in a report on the infestation.’

Swaine returned in the summer of 1914 to discover that the Okanagan infestation had

spread as far west as Princeton. On Okanagan Lake the hillsides appeared “as though

swept by a great fire,” with only isolated patches of Douglas fir surviving. In other areas

beetle attacks were decimating the white pine, and authorities noted the relationship

between the slash left by logging operations and beetle outbreaks. Hewitt advocated the

adoption of “vigorous control methods” to prevent the spread of beetles from fresh slash to

standing timber. “It should be a settled policy in British Columbia to burn all pine slash

each season between October and May, as an aid to bark-beetle control,” he wrote.>

The infestation continued running its course over the next few years, with Swaine

investigating outbreaks and pressing the provincial government for funds to control the

devastation which threatened the province’s white and yellow pine. Infested trees should

be logged and handled in a manner that destroyed broods contained in the bark, reducing

beetle populations to an extent that they would find sufficient breeding material in slash

rather than standing timber. U.S. Bureau of Entomology control projects had demonstrated

that the removal of 75 percent of infested trees would check an outbreak. Floating logs in

water killed some of the broods, barking trees and burning the debris was another option,

but if profitable utilization was not possible simply burning infested timber provided the

cheapest means of control. Finally, in 1919 Swaine and B.C. Forest Branch officials

entered into serious negotiations aimed at developing a cooperative effort to control the

bark beetle menace. By this time the epidemic around Princeton had died out after killing

about 130 million feet of yellow pine, but a new outbreak had emerged around Merritt.

Swaine’s discussions with B.C. authorities produced an agreement to establish a

laboratory at Vernon, and he asked Ralph Hopping, then employed by the United States

Bureau of Entomology in California, to take charge. Regarded as one of the “leading

students of forest-insect life on the west coast,” Hopping accepted the position as

Dominion Forest Entomologist for B.C., settled in at the Vernon courthouse in December

1919, and prepared to launch what Minister of Lands T.D. Pattullo called the province’s

“war on the pine beetle.” It would prove to be a significant campaign, occupying

Hopping’s energies for much of the next decade.

7 H.R. MacMillan, “In British Columbia,” CFJ 9 (July 1913), p. 106; H.R. MacMillan,

“British Columbia Forest Work,” CFU 9 (Oct. 1913), p. 156; J.M. Swaine, “A Forest Insect

Survey in British Columbia,” CFJ 9 (Nov. 1913), pp. 166-67; J.M. Swaine, “The

Economic Importance of Canadian Ipidae,” Proceedings of the Entomological Society of

British Columbia (1913), pp. 41-43.

°C. Gordon Hewitt, “Forest Insect Investigations in British Columbia,” CFJ 10 (Oct. —

Nov. 1914), pp. 102-3.

* “Fighting Forest Insects,” CFJ 11 (Mar. 1915), p. 42; J.M. Swaine, “Problem of the Bark

Beetle,” CFJ 11 (June 1915), pp. 89-92; “Beetles are Killing the Yellow Fir in B.C.,”

Western Lumberman 16 (Oct. 1919), p. 49 [hereafter WL]; R.H. Hopping, “Annual Report

for 1923, Forest Insect Control, Pacific Forest Centre Library [hereafter PFCL].

> Kenneth Johnstone, Timber and Trauma: 75 Years with the Federal Forestry Service,

1899-1974 (Ottawa: Minister of Supply and Services, 1991), pp. 106-7; “Fighting Pine

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 179

A provincial allotment of $10,000 permitted direct control operations to begin during

the spring of 1920 in the Midday Creek valley, a tributary of the Coldwater River. In the

Railway Belt, a Dominion Forest Service crew under D. Roy Cameron worked in the Spius

Creek valley about fifteen miles south of Canford. Designated men first inspected infested

areas, marking trees for cutting. A working party followed, cutting down the trees, peeling

the bark, and burning both bark and limbs. By starting projects early in the spring Hopping

hoped to destroy the beetles before they matured and took flight. “The epidemic can be

practically eradicated,” he declared, given the allotment of sufficient funds to undertake a

comprehensive control program. The province had made admirable progress in its fire

suppression program, but fiscal support for handling the insect menace remained

insufficient to the task. The control effort was "successful as far as it has gone with the

limited funds,” but Hopping had no illusions about his capacity to cope with the ecological

and economic consequences of unregulated cutting. Winter logging operations left an

abundant supply of cull logs and limbs to become infested the following June and July.

Lacking fresh slash that summertime logging would have provided, the beetles attacked

standing trees after emerging. Another winter's cut simply added impetus to the outbreak,

fuelling an epidemic capable of spreading even in standing timber by virtue of the

immense populations. “The removal of slash and infested standing timber is absolutely

necessary if we are to reduce the loss to the minimum, and prevent a state of chronic

menace to our forests,” he warned.°

The cooperative project had three distinct components: direct control, involving cutting

and burning of infested trees; timber sales in areas accessible to mills; and a preventative

emphasis on the burning of slash and cull logs. Hopping supervised all three phases of the

operation, overseeing the work of control crews, inspecting timber sales worked by Nicola

Pine Mills of Merritt, and instructing operators about proper logging methods. Swaine

described the first season’s work as “remarkably successful,” but estimated that some $20-

$40 million worth of yellow pine remained at risk.’

Four control projects involving forty-eight men worked between April | and June 30

the following year in the Vernon and Kamloops Forest District. Crews at Kingsdale, Spius

Creek, Midday Creek and the north end of Adams Lake cut and burned thousands of trees,

but funds were inadequate to keep pace with the infestation. Hopping requested a doubling

of the provincial allotment to $20,000 but held out little hope that the increase would be

granted. The Pacific Coast Lumberman supported his argument that the province should

make a small immediate investment rather than suffer heavier future costs.°

Provincial expenditures rose to $15,500 in 1922, enabling Hopping to place three crews

in the field around Merritt. Together with a federal project on Spius Creek, they treated

over 8,000 trees. ‘““We have found that direct control work on infestations reduces the

infestations 80 percent,” Hopping reported. An untreated epidemic, on the other hand,

Bug in British Columbia,” WL 17 (Mar. 1920), p. 40; R.C. Traherne, “A Further Review of

Applied Entomology in British Columbia,” Proceedings of the Entomological Society of

British Columbia (1921), p. 144.

° “Epidemic of the Bush Beetle,” WL 17 (May 1920), p. 49; “Fighting Bark Beetle in Pine

Forests,’ WL 17 (June 1920), pp. 35-36; Ralph Hopping, “Control of Bark Beetle

Infestations,” Pacific Coast Lumberman 5 (Nov. 1921), p. 77 [hereafter PCL].

’J.M. Swaine, “Bark Beetle Successfully Combatted,” WL 18 (Mar. 1921), p. 28.

* RH. Hopping, “Annual Report, 1921,” PFCL; “The Bark Beetle Menace,” PCL 5 (Nov.

ODN) p22.

180 J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

would increase by 100 to 150 percent, or even more in certain conditions. Thus, work

conducted during the 1922 season saved over five million feet of timber, worth perhaps

$8,000. Encouraged, he suggested that another year or two of direct control would reduce

the infestation sufficiently to permit a much lower annual expenditure.’

Five projects — four in the Merritt area and one at Adams Lake — involved eighty-five

men during the 1923 season at a cost of over $20,000. Hopping again expressed regret that

insect control, which had destroyed about 200 million feet of timber around Princeton and

Merritt at a loss to the province of perhaps $6 million, did not receive the same support

accorded fire prevention and suppression. He also stressed repeatedly that the balance of

nature had been disrupted by lumbering. “Promiscuous cuttings, unless regulated by the

government, upset the natural balance and cause such outbreaks as we are having at the

present time,’ the entomologist observed in 1923. The provincial and Dominion

governments then doubled their allotment to about $45,000 for direct control during 1924.

The provincial share amounted to $35,000 in support of five large fifteen to thirty-six man

projects and several smaller “flying crews,” that treated 19,000 trees. Federal crews

continued to work small infestations at Spius Creek and Adams Lake. With a temporary

staff of four, including H.H. Thomas, Kenneth Auden, Norman Cutler and son George,

Hopping found himself short-handed during the hectic summer season. Although epidemic

infestations in controlled areas had been reduced by 90 percent as a rule, the expansion of

lumbering around the province contributed to the rise of new outbreaks, “tremendously

increasing the work of the personnel.” The Vernon laboratory, Hopping declared in a blunt

plea for additional help, “finds that it has been impossible to properly carry on.”””

The appointment of Hector Richmond and William Mathers as Pest Inspectors in 1925

permitted an expanded research effort. Hopping had initiated cage experiments earlier in

the decade in an effort to determine when the emergence of beetles began, peaked, and

ended, and to identify parasites and predaceous insects. Richmond and Mathers were

assigned to this work, felling, bucking, and limbing infested trees and constructing cages

over the trunk and stump. Richmond inspected the cages daily, collecting and preserving in

alcohol the insect material that emerged. Study of the cedar borer also began during this

period at the Vancouver Forest Products Laboratory under George Hopping, and Mathers

commenced an investigation of the life history of the spruce budworm."'

Ralph Hopping declared victory in the war against the bark beetle in the yellow pine

stands around Merritt early in 1928. “The control measures have been entirely successful,”

he reported, “no epidemic now existing in the yellow pine of British Columbia for the first

time in over fifteen years.” He estimated the saving to the province at $5 million,

attributable to the “100 percent method” of cutting and burning every infested tree and

periodically recleaning infested areas. Over 50,000 trees had been treated in this manner

> R.H. Hopping, “Annual Report for 1922,” PFCL; “Saving Yellow Pine,” PCL 6 (Aug.

1922), p. 64.

'° R.H. Hopping, “Annual Report for 1923,” PFCL; Ralph Hopping, “Forest Entomology,”

Proceedings of the Entomological Society of British Columbia (1923), p. 187; R.H.

Hopping, “Annual Report, Entomological Laboratory, Vernon, B.C., 1924”; Ralph

Hopping, ‘Forest Insect Problems of British Columbia and Their Importance,” WL 22

(Sept. 1925), p. 60.

"RH. Hopping, “Annual Reports, Vernon Forest Insect Laboratory, 1925,”; Hector Allan

Richmond, Forever Green: The Story of One of Canada’s Foremost Foresters (Lantzville:

Oolichan Books, 1983), p. 20.

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 18]

since the start of the campaign, at a cost of approximately $100,000. The situation in

lodgepole pine, however, was less bright. Mountain pine beetle infestations had devastated

vast areas of this commercially unimportant species, moving so rapidly that control

operations had little hope of success even if funding had been available. The laboratory

would also continue to monitor the steady growth of Douglas fir beetle infestations, the

spruce budworm, and other forest insects.”

By this time Hopping had achieved some success in educating National Parks Service

and Dominion Forest Branch rangers to the importance of identifying and reporting

incipient outbreaks, but had made no such progress with B.C. Forest Branch personnel.

"The unconcern of forest rangers and others of the Forest Branch with regard to forest

insect infestations throughout the Province, has been strikingly apparent to us for some

time,” he informed Chief Forester P.Z. Caverhill. Many outbreaks had not been reported

for two or three years after their onset, a situation he hoped to correct by having an

information package sent to each Ranger and making lectures by Division of Forest Insect

officers a feature of annual meetings. An educational campaign of visits to Ranger

headquarters around the province went into effect in 1928, permitting joint inspections of

forest districts. Officials hoped that this sort of cooperation would foster prompt reporting

of outbreaks and eliminate the need for costly control projects.”

The Vernon laboratory conducted only one small control project in lodgepole pine

during 1928, reflecting the gradual shift to a focus on education and research. The

following year Hopping requested funds for a crew to take action against a lodgepole pine

infestation in the Railway Belt, but the species had too little commercial value to justify

the expenditure. George Hopping broke new ground on the coast, however, overseeing

B.C.’s first airplane dusting project on a small area around the Wigwam Inn resort on

Burrard Inlet. Undertaken in part to demonstrate the feasibility of the technique to

Dominion and provincial forestry officials, the experiment against the hemlock looper saw

a Western Canada Airlines Boeing flying boat dust forty-five acres with calcium arsenate.

Mathers studied a spruce budworm infestation at Barkerville that summer, work that would

contribute to his discovery of the budworm’s two-year life cycle in the area. The younger

Hopping took charge of a sub-laboratory at the University of British Columbia in 1930,

and oversaw larger dusting projects on Stanley Park and Seymour Inlet that June. These

endeavours involved three Boeing flying boats at a total cost of over $10,000. '*

The onset of the Great Depression had an immediate impact on the activities of the

Vernon laboratory. Unable to secure funds for control of growing bark beetle infestations

in Douglas fir and lodgepole pine, Ralph Hopping did spend $5,000 that spring in response

'* R.H. Hopping, “Annual Report, Forest Insect Laboratory, Vernon, B.C., Fiscal Year

1927-1928;” J.M. Swaine, “Progress in Forest Insect Control in Canada,” FC 4 (Feb.

1928), np.

'? R.H. Hopping to the Chief Forester, 14 Nov. 1927, copy included in 1927-28 Annual

Report; J.M. Swaine, Forest Entomology and its Development in Canada (Ottawa: King’s

Printer, 1928), p. 12.

MRH. Hopping, “Annual Report, 1928,” PFCL; J.M. Swaine, “Forest Insect

Investigations in Canada, 1928,” FC 5 (June 1929), pp. 37-38; R.H. Hopping, “Annual

Report, 1929,” PFCL; G.H. Hopping, “An Account of the Western Hemlock Looper,

Ellopia Sominaria Hulst, on Conifers in British Columbia,” Scientific Agriculture 15

(Sept. 1934), pp. 24-28; W.G. Mathers, “The Spruce Budworm in British Columbia,” FC 8

(Sept. 1932), pp. 154-57.

tg) J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

to a massive beetle attack at Aspen Grove, one of the original project sites. But a cash-

strapped B.C. government curtailed allocations for fire protection in the early thirties, and

in this fiscal context Hopping had no hope of beetle control funding. Federal budget

cutting in April 1932 led to layoffs of all temporary staff at Vernon. Richmond and the

others continued working in the hopes that their positions would be renewed, receiving a

six-month extension in May accompanied by a 10 percent pay cut that reduced

Richmond’s monthly salary to $112.50."

Despite the need for direct control of further beetle outbreaks in the yellow pine stands

around Merritt during the early 1930s, the laboratory was confined almost entirely to

research. Richmond conducted bark beetle studies at Aspen Grove during the summer

months, devoting the autumn to surveys of damage inflicted to interior forests. Hopping’s

confident assertions of victory in the bark beetle campaign appear to have been misplaced,

then. According to Richmond beetle populations subsided during the mid-1930s due to

several factors, primary among them the exhaustion of mature pine stands. The laboratory

enjoyed greater success, apparently, in combatting an infestation of the European larch

sawfly around Fernie during this period. A 1933 inspection by Mathers discovered an

outbreak of some seventy miles in extent, and a parasite obtained from the Dominion

Parasite Laboratory at Belleville was released over the next two years. This early instance

of biological control in B.C. forest entomology had a beneficial impact on sawfly

populations, saving extensive larch stands from the defoliator.'°

In 1934 J.J. De Gryse took over as Director of the Department of Agriculture’s

Division of Forest Insects from Swaine, who became the agency’s Director of Research.

De Gryse would go on to initiate many important organizational developments, beginning

with the nation-wide Forest Insect Survey. Swaine had taken a step in this direction in

1931 with the establishment of the Forest Insect Intelligence Service. In addition to

publishing a number of circulars dealing with Canada’s principal forest insects, Swaine

had these distributed to industry organizations and forest services along with a request that

field men submit reports on outbreaks in their districts. But after an initial favourable

response Intelligence Service activities waned under the impact of the Depression, leaving

the program moribund."’

De Gryse’s effort to revive the initiative began in December 1934, with an

Entomological Branch conference at Ottawa. Industry and Dominion Forest Service

attendance at sessions dealing with forest insect problems prompted discussions of the

need for cooperation to gather information on the threat posed by the European spruce

sawfly to spruce stands in eastern Canada. A committee organized under the auspices of

the Canadian Society of Forest Engineers and headed by De Gryse rounded up support

among heads of protection organizations, corporations, and wildlife departments in

Ontario and Quebec, and developed procedures for the collection and submission of insect

'° R.H. Hopping “Annual Report, 1930,” PFCL; Hector Allan Richmond, “A History of

Forest Entomology in British Columbia, 1920-1984,” Pt. I, B.C. Forest History Newsletter

9 (Nov. 1984), p. 5.

'° R.H. Hopping, “Annual Report of the Vernon Forest Insect Laboratory, 1932,” PFCL;

R.H. Hopping, “Annual Report, 1933,” PFCL; R.H. Hopping, “Annual Report of the

Vernon Forest Insect Laboratory, 1934,” PFCL; Richmond, Forever Green, pp. 102-3;

George R. Hopping, “A Forest Insect Problem in British Columbia,” FC 11 (Dec. 1935),

pp. 258-61].

'’ Prebble, “Forest Entomology,” p. 352; Wallace, “Forest Entomology,” p. 121.

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 183

samples to the Division of Forest Insects. In 1936 about eight hundred cooperators

participated, examining trees in their areas to determine the presence of the sawfly and

other destructive pests. They were provided with collapsible boxes for mailing samples,

instructions, report forms, relevant circulars, and asked to make monthly reports between

June and September. The Ottawa laboratory received 512 samples in 1936, the first stage

in fulfilling De Gryse’s vision of a national forest insect intelligence system.’

The Forest Insect Survey expanded to the Maritimes and B.C. in 1937, with the Vernon

laboratory serving as headquarters for activities in the west. The B.C. Forest Branch and

National Parks Branch participated the first year, field men making collections by placing

a canvas ground sheet beneath a tree and “shaking it vigorously or hitting it with an axe” to

dislodge larvae. Although the Vernon laboratory received no sawfly specimens in 1937,

three hemlock looper outbreaks were brought to light in this fashion. The number of

cooperators increased steadily over the following years, along with the volume of material

sent in for analysis and rearing. De Gryse’s establishment of a corps of trained forest insect

rangers added to the survey’s effectiveness. By 1947 seven rangers operated in B.C.,

divided evenly between the coast and interior districts. Surveying of the coastal forests

reached a new level of efficiency that year with the commissioning of a sixty-foot boat, the

JM. Swaine."”

After supervising the survey through its initial stage, Ralph Hopping retired in

December, 1938. He died at his Vernon home only two years later, acknowledged as one

of the foremost forest entomologists in North America. His son George took over as

director at the Vernon laboratory, now part of the Entomology Division of the Department

of Agriculture’s Science Service after a 1937 reorganization. The younger Hopping’s

leadership coincided with the establishment of a summer field station and insectary at

Trinity Valley. Plans were also in the works that resulted in a similar facility for coastal

studies at the B.C. Forest Branch’s Cowichan Lake Experiment Station. By this time the

most pressing problem facing the Vernon staff involved a massive beetle outbreak in

Kootenay National Park. Infestation of the park’s lodgepole pine stands began in the early

1930s, and covered seventy-two square miles by 1937. Ralph Hopping had recommended

control measures to contain the outbreak, a proposal that fell victim to Depression-era

budget constraints.””

A major restructuring of the Science Service’s western forest entomology organization

in 1940 brought a new laboratory at Victoria into existence. M.L. Prebble was transferred

from Fredericton to head that facility, which took over the equipment from the UBC sub-

'’ Canada, Annual Reports of the Forest Insect Survey, 1936-1937-1938 (Ottawa: King’s

Printer, 1939), pp. 1-2; J.J. De Gryse, “Cooperation in Forest Insect Studies Relating to

Conservation,” Journal of Forestry 36 (1938), pp. 983-86.

Canada, Annual Reports of the Forest Insect Survey, 1936-1937-1938, pp. 26-33; R.H.

Hopping, “Annual Report of the Vernon Forest Insect Laboratory for the Fiscal Year

Ending March 31, 1938,” p. 2; Canada, Annual Report of the Forest Insect Survey, 1947

(Ottawa: King’s Printer, 1947), p. 91.

*’ R.H. Hopping, “Annual Report of the Vernon Forest Insect Laboratory for the Fiscal

Year Ending March 31, 1937,” PFCL; R.H. Hopping, “Annual Report of the Vernon

Forest Insect Laboratory for the Fiscal Year Ending March 31, 1938,” PFCL; “Ralph

Hopping (1868-1941),” Proceedings of the Entomological Society of British Columbia 38

(1942), pp. 3-4.

184 J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

unit. Kenneth Graham moved west from Vernon, while William Mathers returned to

Vernon from the now defunct Vancouver branch. The new Victoria research centre took

responsibility for forest and shade tree insect problems in the coastal area. The shuffle

confined Vernon’s jurisdiction to B.C. forests east of the Cascade range and all of Alberta

with the exception of the province’s southeast corner, to be administered from the Science

Service’s laboratory at Indian Head, Saskatchewan.”’

Canada’s entry into World War II forced funding restrictions at Vernon, where the

insect survey and research on the larch sawfly dominated activities. George Hopping

discontinued work on some of the less pressing research projects, but by 1941 the bark

beetle problem had become acute on the Kootenay and Banff National Parks. By the end

of that year an estimated 400 million board feet of lodgepole pine had been destroyed on

the former, and Banff had become the scene of a large, unrelated outbreak in 1940.

Hopping and Mathers attributed the mountain pine beetle infestations to several years of

below-average precipitation that diminished the vigour of the mature lodgepole pine stands

covering extensive areas in both parks.”

Since the situation on Kootenay National Park was beyond repair, officials decided to

concentrate control efforts at Banff, where the outbreak was in its early stages.

Fortunately, given the severity of wartime labour shortages, the Parks Branch succeeded in

securing a supply of Alternative Service Workers for the project. Three forty-man camps

for the conscientious objectors were established during the fall and winter of 1941, the

crews working until late spring. Three or four men from each crew served as a cruising

party, the remainder cutting, decking, and burning the infested trees. They covered 5,725

acres during the 1941-42 season, treating 9,192 trees. Crews resumed after the 1942 fire

season, completing control work in the Bow valley. This effort, in conjunction with low

temperatures in January 1943 that killed a high percentage of broods above the snow line,

left only a small area to work the following season. Hopping and Mathers praised the

conscientious objectors who, with few exceptions, “proved to be reliable workers rapidly

acquiring proficiency in control procedure.” Thanks to prompt action and the availablility

of the Alternative Service Workers, the Banff project stands out as a rare success in the

direct control of a bark beetle infestation, saving the park from a “major catastrophe.””*

Federal government support of forest entomology increased in the immediate postwar

period as wood products assumed greater importance in the economic boom. In late 1945

Reconstruction Minister C.D. Howe announced the creation of a Forest Insects Control

Board, comprised of federal, provincial, and industry representatives, to coordinate

entomological research and control measures across the country. In B.C., growing

recognition of the threat insects posed to commercial forestlands was reflected in the

establishment of an industry-endowed chair in forest entomology at UBC. George

Hopping received a leave-of-absence from Vernon to initiate the course of instruction

71 M.L. Preeble, “Forest Insect Investigations, Victoria Unit, Report for 1940,” PFCL; G.

Hopping, “Annual Report, Vernon Forest Insect Laboratory, 1940,” PFCL; Hector Allan

Richmond, “A History of Forest Entomology in British Columbia, 1920-1984,” Pt. II, B.C.

Forest History Newsletter 10 (Mar. 1985), p. 3.

*? G. Hopping, “Annual Report of the Vernon Forest Insect Laboratory, 1940,” PFCL; G.

Hopping, “Annual Report of the Vernon Forest Insect Laboratory, 1941,” PFCL.

*> George R. Hopping and W.G. Mathers “Observations on Outbreaks and Control of the

Mountain Pine Beetle in the Lodgepole Pine Stands of Western Canada,” FC 11 (June

1945), pp. 98-108; G. Hopping, “Annual Report, Vernon Forest Insect Laboratory, 1942,”

PFCL; G. Hopping, “Annual Report of the Vernon Forest Insect Laboratory, 1943,” PFCL.

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 185

during the 1947-48 academic year. Industry figures also met with De Gryse in 1948 to

discuss the development of insect control strategies on the west coast. Prebble left Victoria

to take charge of a new research centre at Sault Ste. Marie that same year, bringing

Richmond to Victoria as his replacement. That laboratory became the headquarters for

forest entomological work in B.C. in 1948, relegating Vernon to sub-laboratory status.”*

The Vernon laboratory, with a staff of fifteen during the summer months, remained

responsible for the forest insect survey in the interior. By 1950 there were eleven forest

insect rangers in the interior and nine on the coast. The establishment of permanent sample

plots around the province made for more systematic collections, and the introduction of a

punch card system of data entry in 1952 facilitated rapid analysis at the laboratories. By

this time 171 permanent sample plots had been established in the vast interior of B.C.

Insects sent to Vernon were sorted, and immature forms reared at the Trinity Valley field

station near Lumby for the determination of life histories and host-parasite relationships.”

Airborne chemical attack on defoliators involving the application of D.D.T. captured

the enthusiasm of forest entomologists during the postwar decades. Scientists at the

Victoria laboratory studied an alarming incidence of attacks by the black-headed budworm

and hemlock looper in coastal forests during the 1940s. The Division of Entomology

cooperated with the B.C. Forest Service and industry in the first aerial spraying with

D.D.T. on 12,000 acres in the Nitinat valley in 1946 to control a looper infestation. No

effort to examine the impact on aquatic life accompanied this project, but the conflict

between D.D.T. spraying in defence of timber values and the consequences for the

fisheries resource would come to play a major part in limiting use of the insecticide.

Vernon entomologists participated in the first aerial spraying of D.D.T. in the interior in

1948, when a false hemlock looper infestation threatened the Christmas tree crop in the

Windermere valley.”°

The most extensive application of D.D.T. in B.C. occurred in 1957 in response to a

black-headed budworm infestation on northern Vancouver Island. By 1956 the outbreak

covered some 3,000 square miles. Representatives of firms with holdings in the area

collaborated with Forest Service and Forest Biology Division officials in developing a

control project designed to save timber valued at $300 million. An experimental spraying

that summer proved effective, and plans went ahead to spray 156,000 acres the following

** Johnstone, Timber and Trauma, p. 138; “Destruction of Canadian Timber By Forest

Insects Attains “National Disaster” Proportions, Says Minister Howe,” British Columbia

Lumberman 29 (Dec. 1945), p. 30 [hereafter BCL]; Richmond, Forever Green, p. 131;

“Forest Entomology Vital Study, Says Expert,” BCL 31 (Dec. 1947), p. 114; “Forest

Entomology,” FC 24 (Dec. 1948), pp. 293-94.

> “Forest Insect Survey in Interior B.C. Extended,” BCL 34 (Feb. 1950), p. 103; B.C.,

Report of the Forest Service, 1948 (Victoria: King’s Printer, 1949), p. 55; B.C. Report of

the Forest Service, 1949 (King’s Printer, 1950), p. 72; H.A. Richmond, “Forest Insect

Surveys,” Proceedings of the Entomological Society of British Columbia (1953), pp. 28-

30; George Hopping, “Forest Insect Laboratory at Vernon Responsible for Control in Huge

Area,” BCL 31 (July 1947), pp. 112, 114.

’° MLL. Prebble and K. Graham, “The Current Outbreak of Defoliating Insects in Coast

Hemlock Forests of British Columbia,” BCL 29 (Feb. 1945), pp. 25-27, 42-48; “Forest

Insect Conditions and Research in 1946,” FC 23 (Mar. 1946), p. 89; H.A. Richmond,

‘Forest Insect Problems in British Columbia,” BCL 32 (May 1948), p. 76; “Helicopter

Fighting False Hemlock Looper Attack,” BCL 32 (July 1948), p. 141.

186 J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

summer with costs shared equally by the B.C. Loggers Association, federal, and provincial

governments.~’

By this time some evidence of fish mortality from D.D.T. spraying in New Brunswick

had been accumulated, and officials of the B.C. Game Commission and _ federal

Department of Fisheries participated in the planning process. Industry’s Pest Control

Committee turned down requests to have some fish- producing areas eliminated from the

project. The fisheries biologists were also denied when they asked that the D.D.T. dosage

be cut by half, to one-half pound per acre. Forest Biology Division representatives ruled

that the reduced dosage might not be effective, in rejecting the proposal. The Committee

did agree, however, to make some adjustments in flight patterns to minimize damage to

fish populations. The project was declared a success from an entomological standpoint,

although Richmond observes that budworm populations declined simultaneously on

untreated areas in the following years. More significantly, the project resulted in

considerable mortality in cohoe salmon populations. One industrial forester involved

warned that if D.D.T. spraying was to be continued, it must be on the basis of non-lethal

concentrations or public opinion would forbid its use altogether.”

The 1957 disaster influenced procedures followed in 1960 on Moresby Island, when a

black-headed budworm infestation promped another D.D.T. control program. This time

forestry officials agreed to reduce the dosage to one-quarter pound per acre, resulting in

negligible salmon mortality. But studies carried out in conjunction with the project

indicated that even this concentration was lethal to salmon fry, supporting other research

that prompted a general outcry against use of the insecticide in North America. Awakening

to the need for acceptable alternatives, the Forest Biology Division and B.C. Loggers

acc auon initiated tests of a bacterial insecticide on the Queen Charlotte Islands in

1960.

Bark beetles remained the major problem confronting entomologists in the interior

during the early 1950s, although increasing spruce budworm populations also caused

concern. Large outbreaks of the mountain pine beetle developed in the Columbia valley,

and the Douglas fir bark beetle caused significant losses in the Quesnel region. Scientists

studied the efficacy of ground-spraying insecticides such as D.D.T. in an effort to develop

a faster and less costly method of controlling pine beetle epidemics, but such operations

were themselves considered too expensive for widespread use in the province.”

?7 B.C. Report of the Forest Service for 1956 (Victoria: Queen’s Printer, 1957), p. 79; B.C.

Report of the Forest Service for 1957 (Victoria: Queen’s Printer, 1958, p. 70; G.S. Brown,

A.P. Randall, R.R. Le Jeune, and G.T. Silver, “Black-Headed Budworm Spraying

Experiments on Vancouver Island, British Columbia,” FC 34 (Sept. 1958), pp. 299-306.

°° E.G. Marples, “Significance of Fish Mortality in Forest Spraying Operations,”

Proceedings of the Western Forestry and Conservation Association (1957), pp. 44-55;

R.A. Coulter and E.H. Vernon, “Effects of Black-Headed Budworm Control on Salmon

and Trout in British Columbia,” Canadian Fish Culturalist 24 (1959), pp. 23-40.

*’ 1.S. Todd and K.J. Jackson, “The Effects on Salmon of a Program of Forest Insect

Control with D.D.T. on Northern Moresby Island,” Canadian Fish Culturalist 30 (Dec.

1961), pp. 15-27; Hector A. Richmond, “A New Look at Aerial Spraying in Forest

Resource Protection,” Proceedings of the Western Forestry and Conservation Association

(1960), pp. 30-32.

°° B.C., Report of the Forest Service for 1951 (Victoria: King’s Printer, 1952), p. 85.

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 187

Jack Walters of the Victoria Biological Laboratory and UBC forest entomologist Ken

Graham worked on the Douglas fir beetle problem, investigating the factors contributing to

recent outbreaks. One rewarding study at Douglas Lake involved the felling of “trap trees”

and release of beetles to determine their flight patterns and the types of trees most

vulnerable to attack. By the late 1950s it had been demonstrated that the presence of felled

logs in a stand induced attacks on surrounding timber, insight that informed a silvicultural

control project involving Western Plywoods, the Forest Service, and Forest Biology

Division. This early effort to merge forestry and entomological principles into forest

management planning sought to draw Douglas fir beetles into pre-selected areas with

attractive logs, then remove these along with nearby infested trees in an attempt to lower

populations.”’

Vernon’s status as a centre of entomological research declined during the 1950s, as the

Victoria laboratory became the hub of federal government biological science. By 1955,

when D.A. Ross replaced William Mathers as officer-in-charge at Vernon, the staff

numbered twenty-one permanent employees concerned primarily with carrying out the

insect and disease survey in the interior. Five years later the Forest Biology Division

became the Forest Entomology and Pathology Branch of the new Canada Department of

Forestry under director M.L. Prebble, and by 1963 the Vernon staff had dwindled to

fifteen. Opening of the Department of Forestry’s new $2.5 million forest research

laboratory at Victoria in 1965 further undermined the Vernon station’s prospects.”

During the same period forest insect problems in the interior became, if anything even

more serious. During the early 1960s the central interior experienced the most serious

spruce bark beetle outbreak ever recorded, and damage from the Douglas fir and mountain

pine beetles increased in the southeastern part of the province. Between 1961 and 1965 the

spruce beetle destroyed over 500 million cubic feet of timber in central B.C., an outbreak

presumed to be caused by abundant slash, warm summers, and mild winters. Research on

these infestations was carried out from Victoria, while the B.C. Forest Service supervised

large-scale salvage operations in damaged timber.*”

The end for the Vernon sub-laboratory came in 1970, when an economizing federal

government closed the facility. Most of the staff members transferred to Victoria

laboratory, now the centre of federal forestry science and base of the insect and disease

survey for the entire province. Federal authorities insisted that the closure would not

impair entomological work in any way, claiming improved efficiency for its entire B.C.

operation. Nevertheless, Ottawa's decision to terminate activity at Vernon has been

questioned frequently. Indeed, expansion of the pulp and paper industry in the central and

southern interior since the 1960s has given these forests greater economic importance than

the coastal stands that provided the raw material for the province’s early lumber industry.

>! kK. Graham, “The Bark-Beetle Problem in Douglas Fir of the Interior,” BCL 36 (July

1952), p. 42; “Bark Beetle Spreads,” BCL 36 (Aug. 1952), p. 100; H.A. Richmond,

“Douglas Fir Bark Beetle in B.C.,” BCL 37 (May 1953), pp. 43, 90-92; B.C., Report of the

Forest Service For 1958 (Victoria: Queen’s Printer, 1959), p. 68.

*? Miles Overend, “Intelligence Service Battles Bugs,”BCL 47 (May 1963), p. 38; Dr. D.A.

Ross, “Forest Entomologist, Retires,” Truck Logger (Jan. 1967), p. 23.

*° Canada, Department of Forestry Annual Report, 1963-1964 (Ottawa: Queen’s Printer,

1964), pp. 33-35; “Spruce Beetle Damage Studied,” BCL 51 (Jan. 1967), p. 66; Canada,

Department of Forestry Annual Report, 1964-1965 (Ottawa: Queen’s Printer, 1965), pp.

33-35.

188 J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

Moreover, bark beetles remain a significant obstacle to rational forest management in these

areas. In recent years Ralph Hopping’s old enemy the mountain pine beetle has re-emerged

with great intensity in the Okanagan and Cariboo regions, thanks to favourable climatic

conditions and the advance in postwar fire protection techniques that brought extensive

lodgepole pine stands to maturity. Hopping would also be familiar with government’s

modern response to these epidemics, if not the precise techniques. Although entomologists

now rely on sophisticated computer mapping systems to trace the progress of epidemics,

salvage harvesting is the primary control technique. Much damaged timber now finds its

way to the sawmill or pulp mill, and the use of trap trees and pheromones show promise in

slowing the spread of outbreaks. But the magnitude of recent spruce and pine beetle

infestations leave forestry officials no alternative but to concentrate logging operations on

the most severely affected areas, an approach which in principle differs little from

Hopping’s early twentieth century control projects.*

Journal Abbreviations in footnotes

BCL British Columbia Lumberman

CEI Canadian Forestry Journal

FC Forestry Chronicle

LSJ Logging and Sawmilling Journal

BEL Pacific Coast Lumberman

PFCL Pacific Forestry Centre Library

WL Western Lumberman

ACKNOWLEDGEMENTS

This work was supported in part by a contract from Natural Resources Canada, Canadian

Forest Service.

“Dr. D.A. Ross,” pp. 36-37; Hector Allan Richmond, “A History of Forest Entomology

in British Columbia, 1920-1954,” Pt. II, B.C. Forest History Newsletter (Mar. 1985), p. 3;

“Northwood Confronts Beetle,” Logging and Sawmilling Journal 14 (Nov. 1983), pp. 16-

18 [hereafter LSJ]; L. Ward Johnson, “Chasing the Pine Beetle,” LSJ 21 (July 1990), pp.

14-16; “Beetle Attack Prompts AAC Increases,” LSJ 23 (Oct. 1992), p. 7; Jim Stirling,

“Going to War,” LSJ 31 (Feb. 2000), pp. 5-7.

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 189

Implications of using development rates of blow fly (Diptera:

Calliphoridae) eggs to determine postmortem interval

SHERAH L. VANLAERHOVEN,' and GAIL S. ANDERSON,”

CENTRE FOR PEST MANAGEMENT

DEPARTMENT OF BIOLOGICAL SCIENCES, SIMON FRASER UNIVERSITY,

8888 UNIVERSITY DRIVE, BURNABY, BC, CANADA VSA 186

ABSTRACT

This research examined the eclosion times of blow fly eggs to determine whether

eggs begin to develop at the time of oviposition, or in vivo. Eggs were obtained

from laboratory colonies of Calliphora vicina Robineau-Desvoidy, Phaenicia

sericata (Meigen) and Eucalliphora latifrons (Hough) and observed at 2-h

intervals. All three species had eggs eclose earlier than the expected minimum of

22 h at 21°C. Precocious egg development occurred for 75% of the C. vicina egg

mass, while 100% of the FE. /atifrons and P. sericata egg masses developed early.

Subsequently, we denied an oviposition medium to fresh C. vicina and P. sericata

colonies for 7 and 14 d and compared the eclosion times with that of eggs from

colonies with a continual access to beef liver. In both species, no precocious egg

development was observed as the eggs eclosed 3-4 h after the expected minimum

time of eclosion in both treatments and control. Finally, we examined eclosion

times of eggs laid by blow flies in the wild. Eggs laid in the wild by P. sericata and

C. vicina also took 1-3 h longer to eclose than the expected minimum time of

eclosion. Our first experiment demonstrated that eggs laid by a single female at one

time, can eclose at a wide variety of times, ranging from 2 h to the expected 22 h

after oviposition at 21°C. Our inability to repeat the early eclosion in the laboratory

with new colonies, despite the denial of oviposition media, or in the wild under

natural conditions, is reassuring to those using egg development and eclosion to

determine elapsed time since death. Clearly this phenomenon is not common, and

may be explained as an artifact of laboratory colonies that do not have a regular

influx of wild blow flies.

Key words: forensic entomology, medico-legal entomology, elapsed time since

death

INTRODUCTION

Forensic entomology, or the use of insects to determine the elapsed time since death of

a homicide victim, is a technique that has been employed in many homicide investigations

worldwide (Goff 1992; Leclercq and Vaillant 1992; Lord et al. 1994; Anderson 1995). It is

the most accurate and often the only method available to determine elapsed time since

death after 72 h. However, it also is used during the first 72 h after death, particularly in

high profile crimes, to confirm pathological parameters, or when only a portion of the body

has been recovered. Traditionally, medical parameters are used to determine time since

death in the first hours after death, but these involve many variables (Henssge et al. 1995)

and pathologists are often reluctant to offer an opinion on time since death when more than

' Pacific Agri-Food Research Centre, Agassiz, BC

* School of Criminology, Simon Fraser University, Burnaby, BC

190 J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

a few hours have passed. Thirty percent of forensic entomology cases in Canada in 1995

involved blow fly (Diptera: Calliphoridae) egg evidence alone and this trend has continued

(Anderson and Cervenka 2001). Although the cases were mainly homicide, they also

included one poaching case where blow fly egg development evidence was vital in

connecting time of death of bear cubs with the perpetrators at the scene (Anderson 1999).

Since blow flies usually arrive and begin laying eggs within minutes of death (Anderson

and VanLaerhoven 1996), an analysis using the eclosion times of blow fly eggs will

provide an estimate of the minimum time since death in the early postmortem interval. This

method requires accurate research on the developmental rates of eggs. Previous research

indicates that the time necessary for blow fly eggs to eclose depends on the species and the

temperature (Kamal 1958; Nuorteva 1977; Greenberg 1993; Anderson 2000). However,

these developmental rates and times of eclosion assume that egg development begins after

oviposition and that the eggs do not begin to develop within the adult fly. Our laboratory

research has indicated that this may not always be the case. If in vivo development does

occur, this would change the estimate of elapsed time since death by as much as 24 h. We

hypothesized that female flies which have a suitable oviposition medium available, will

Oviposit eggs which eclose after the normal length of time, after oviposition; whereas flies

which are denied a suitable oviposition medium may have eggs developing in vivo, thereby

decreasing the length of time between oviposition and eclosion.

The objectives of this research were to: determine whether insect eggs laid on a

homicide victim begin to develop at the time of oviposition, or in vivo, as we have

observed occasionally in the lab; and to determine whether early eclosion occurs in the

wild or is an artifact of laboratory conditions.

MATERIALS AND METHODS

We examined egg eclosion under laboratory conditions at 21°C for three species of

blow fly: Calliphora vicina Robineau-Desvoidy, Phaenicia sericata (Meigen) and

Eucalliphora latifrons (Hough). All three species were reared in laboratory colonies

descended from wild specimens collected locally in the Lower Mainland of British

Columbia. They had been under laboratory conditions for approximately a year. On 5

March 1994, beef liver was presented to gravid females and after several hundred eggs

were laid by ~10 females over a 30 min period, the liver was removed from the cages. Each

egg mass was examined for eclosion immediately after oviposition and at 2 h intervals until

eclosion.

The experiment was repeated at 21°C with new colonies of blow flies, this time varying

the availability of an ovipositional medium. Fresh wild caught C. vicina and P. sericata

colonies were established. On | May 1995, newly emerged adults of each species were

exposed to fresh beef liver for 24 h to ensure that all received a protein meal for the

development of ovaries and testes (Erzinclioglu 1996). The adults were then divided into

three groups. The first group was given immediate and continuous access to beef liver as an

oviposition medium. The second group was given only water and sugar for 7 d after the

females were gravid, and was then given fresh beef liver as an oviposition medium. The

final group was given water and sugar but was denied an oviposition medium until 14 d

after the females were gravid. A minimum of 75 males and 75 females of each species were

used, with one cage per species per treatment. Five females were dissected each day to

determine the time taken until eggs were mature. The delayed access to an oviposition

medium was timed after the females were gravid. After each group was presented with beef

liver, and eggs were laid over a 30-min period, the egg mass was removed from the cage

and observed every 2 h until all the eggs had eclosed.

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 19]

We also tested eggs laid by blow fly females in the wild. Petri dishes with

approximately 250 g of fresh beef liver were exposed in partially sunny locations in

Coquitlam, BC. The experiments were conducted between 17-25 September 1996 and 2-10

June 1997. At all times, blow flies were abundant in this mild region. The experiment was

replicated 15 times. After oviposition of at least 100 eggs in a 30-min period, the petri

dishes were covered to prevent further oviposition and moved indoors. Each egg mass was

examined for eclosion at 2-h intervals until eclosion.

Ambient temperature was recorded at 30-min intervals throughout each experiment

using a double channel datalogger (SmartReader 1°, Young Environmental Systems,

Richmond, BC). Temperatures cited are means of records from the time eggs were laid

until eclosion was complete.

RESULTS

All three species had eggs eclose earlier than expected at 21°C (Table 1). Precocious

egg development occurred for 75% of the C. vicina egg mass, while 100% of the E.

latifrons and P. sericata egg masses developed early (Fig. 1).

100

—a— C. vicina

—-«--P. sericata

80 sage oe es (QUITONS

% of Eggs Eclosed

0 2 4 6 8 10 12 14 16 18 20 22 24

Hours After Oviposition

Figure 1. Percent of eggs eclosed from egg masses of three laboratory colonies of blow

flies.

When new colonies of P. sericata and C. vicina were established, no precocious egg

development was observed (Table 2), despite the lack of ovipositional media. Phaenicia

sericata and C. vicina females took 3 d at 21°C to develop mature eggs in their ovaries.

In the field experiments, no precocious egg development was observed for eggs laid by

P. sericata and C. vicina (Table 3). The mean temperature was 20°C for the September

experiment and 23°C for the June experiment.

192 J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

Table 1

Egg eclosion from egg masses of laboratory colonies of blow flies compared to expected

minimum times of eclosion at 21°C (Anderson 2000).

Minimum Time of Eclosion (h)

Species Expected Observed

C. vicina 22 2

P. sericata 21 14

E. latifrons 22 2

Table 2

Egg eclosion from egg masses of laboratory colonies of blow flies: held continuously with

Oviposition media available; held 7 d without oviposition media; and held 14 d without

Oviposition media, compared to expected minimum times of eclosion at 21°C (Anderson

2000).

Minimum Time of Eclosion (h)

Oviposition media Species Expected Observed

Available C. vicina 22 26

Available P. sericata | 24

7d C. vicina D2 26

7d P. sericata 21 24

14d C. vicina 22 26

14d P. sericata 21 24

Table 3

Egg eclosion from egg masses of wild blow flies compared to expected minimum times of

eclosion (Anderson 2000).

Minimum Time of Eclosion (h)

Mean Temperature Species Expected Observed

20°C C. vicina 26 28

20°C P. sericata 25 26

23€ C. vicina 21 24

23.6 P. sericata 21 24

DISCUSSION

It is currently accepted that blow fly eggs do not generally develop in the female fly,

but only begin to develop after oviposition. Therefore, a measure of the developmental

stage can be used to predict the age of the egg, and the time of eclosion can be used to

count backwards to determine the time of oviposition. However, our first laboratory

experiment demonstrated that eggs laid at the same time can eclose at a wide variety of

times, ranging from 2 h to the expected 22 h after oviposition.

Early eclosion of blow fly eggs has been described in the literature, although it is rare

(Auten 1934; Reiter 1984; Erzinclioglu 1990). It is possible that female flies may delay

oviposition until a suitable site is found (Auten 1934). One recent study examined internal

egg development of Phormia regina (Meigen) and stated that only one developing egg can

be withheld by females, as this one egg enters the oviduct and is fertilized, whereas, the rest

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 193

do not enter the oviduct until oviposition (Erzinclioglu 1990). Another study examined

Calliphora terraenovae Macquart, C. vomitoria (L.), C. vicina and P. sericata (Meigen)

and found precocious egg development of at least one egg within all four species (Wells

and King 2001).

The trigger for development within the female remains unknown. Our inability to repeat

the early eclosion in the laboratory with new, wild-captured colonies, despite the denial of

Oviposition media, or in the wild under natural conditions, is reassuring to those using egg

development and eclosion to determine elapsed time since death. Clearly this phenomenon

is not common, and may be explained as an artifact of lab colonies that do not have a

regular influx of wild blow flies; it may even have been an artifact of those specific

colonies, although this seems unlikely. In fact, in a large number of other experiments

conducted over several years, in which eggs were observed every 1-2 h until eclosion, not

once was this phenomenon observed (Anderson 2000). As well, many other researchers

who have performed similar experiments have not mentioned early eclosion (Melvin 1934;

Kamal 1958; Nuorteva 1977; Nishida 1984; Greenberg 1993).

ACKNOWLEDGEMENTS

This research was supported by a research grant from the Pathology/Biology Section of

the American Academy of Forensic Sciences. We would like to thank Dr. Margaret

Dogterom for the use of her property and Simon Fraser University for the use of its

facilities. We would also like to extend our gratitude to Steve Halford for advice and

assistance, and to Hersimer Johl and Jasmine Wiles. We would like to thank Dr. Lisa

Poirier for her advice and editorial comments.

REFERENCES

Anderson, G.S. 1995. The use of insects in death investigations: an analysis of forensic entomology in

British Columbia over a five year period. Canadian Society of Forensic Sciences Journal 28: 277-292.

Anderson, G.S. 1999. Wildlife forensic entomology: determining time of death in two illegally killed black

bear cubs, a case report. Journal of Forensic Sciences 44: 856-859.

Anderson, G.S. 2000. Minimum and maximum developmental rates of some forensically important

Calliphoridae (Diptera). Journal of Forensic Sciences 45: 824-832.

Anderson, G.S. and V.J. Cervenka. 2001. Insects associated with the body: their use and analyses. In:

W.D. Haglund and M. Sorg, (Eds.) Forensic Taphonomy, The Postmortem Fate of Human Remains.

CRC Press, Boca Raton.

Anderson, G.S. and S.L. VanLaerhoven. 1996. Initial studies on insect succession on carrion in

southwestern British Columbia. Journal of Forensic Sciences 41: 617-625.

Auten, M. 1934. The early embryological development of Phormia regina: Diptera (Calliphoridae).

Annals of the Entomological Society of America 27: 481-506.

Erzinclioglu, Y.Z. 1990. On the interpretation of maggot evidence in forensic cases. Medical Science and

Law 30: 65-66.

Erzinclioglu, Y.Z. 1996. Blowflies. Naturalist's Handbooks 23 Richmond Publishing Co. Ltd., Slough,

UK.

Goff, M.L. 1992. Problems in estimation of postmortem interval resulting from wrapping of the corpse - a

case study from Hawaii. Journal of Agricultural Entomology 9: 237.

Greenberg, B. 1993. Different developmental strategies in two boreal blow flies (Diptera: Calliphoridae).

Journal of Medical Entomology 3: 481-484.

Henssge, C., B. Madea, B. Knight, L. Nokes and T. Krompecher. 1995. The estimation of the time since

death in the early postmortem interval. 2" Edition, Arnold. London.

Kamal, A.S. 1958. Comparative study of thirteen species of sarcosaprophagous calliphoridae and

sarcophagidae (Diptera) I. Bionomics. Annals of the Entomological Society of America 51: 261-270.

Leclercq, M. and F. Vaillant. 1992. Forensic Entomology: An original case. Annales De La Societe

Entomologique De France 28: 3-8.

194 J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

Lord, W. D., M.L. Goff, T.R. Adkins and N.H. Haskell. 1994. The black soldier fly Hermetia illucens

(Diptera: Stratiomyidae) as a potential measure of human postmortem interval: observations and case

histories. Journal of Forensic Sciences 39: 215-222.

Melvin, R. 1934. Incubation period of eggs of certain muscoid flies at different constant temperatures.

Annals of the Entomological Society of America 27: 406-410.

Nishida, K. 1984. Experimental studies on the estimation of postmortem intervals by means of fly larvae

infesting human cadavers. Japan Journal of Legal Medicine 38: 24-41.

Nuorteva, P. 1977. Sarcosaprophagous insects as forensic indicators. pp. 1072-1095 In: C.G. Tedeschi

(Ed.) Forensic medicine : a study in trauma and environmental hazards. Saunders, New York.

Reiter, C. 1984. Zum Wachtstumsverhalten der Maden der blauen Schmeissfliege Calliphora vicina. Z.

Rechtsmed. 91: 295-308.

Wells, J. D. and J. King. 2001. Incidence of precocious egg development in flies of forensic importance

(Calliphoridae). Pan-Pacific Entomologist. In press

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 195

Observations on the behavior of Monochamus scutellatus

(Coleoptera: Cerambycidae) in northern

British Columbia

JEREMY D. ALLISON’ and JOHN H. BORDEN

CENTRE FOR ENVIRONMENTAL BIOLOGY

DEPARTMENT OF BIOLOGICAL SCIENCES, SIMON FRASER UNIVERSITY,

8888 UNIVERSITY DRIVE, BURNABY, BC, CANADA VSA 1S6

ABSTRACT

The location, behavior, and sex were recorded for 329 whitespotted sawyers,

Monochamus scutellatus (Say), on horizontal host logs in a logyard in Ft. Nelson BC.

Over 65% of all males and females observed, and 58% of oviposition, occurred on the

sides of horizontal host logs. This behavior would minimize the costs of desiccation

and slow development of progeny on the upper and lower sections of logs,

respectively. The sex ratio was male-biased throughout the season, rising to 4.1 males

per female on 5 August 2000. Copulation and oviposition peaked on 9 and 21 July,

respectively. By 5 August copulation was no longer observed. A late-season increase in

the proportion of mobile males may represent a change in male reproductive strategy

from selecting a preferred oviposition site and waiting for female arrival, to active

pursuit of increasingly scarce females.

INTRODUCTION

Wood-boring beetles in the genus Monochamus Megerle (Coleoptera: Cerambycidae)

reproduce in stressed, dying or dead coniferous trees throughout North America (Rose

1957). The larvae feed under the bark, in the sapwood and sometimes deep into the

heartwood (Linsley 1961), often boring long tunnels which weaken and degrade the wood

and provide infection courts for wood-rotting fungi (Vallentgoed 1991). Five North

American Monochamus spp. are also known vectors of the pinewood nematode,

Bursaphelenchus xylophilus (Steiner et Buhrer) Nickle (Table 1). Of the eight

Monochamus spp. found in Canada, the whitespotted sawyer, M. scutellatus (Say), is the

most common and has the largest range (Table 1) (Linsley and Chemsak 1984; Gosling

and Gosling 1976).

Female M. scutellatus, deposit eggs singly in niches chewed in the bark; the eggs hatch

in 12 days on average (Rose 1957; Raske 1972; Cerezke 1975). The larvae feed in the

phloem and continue to feed there even after they have bored into the wood, where they

also overwinter. The following spring they continue feeding and mining. Under favorable

conditions M. scutellatus is univoltine. Mature larvae construct a pupal cell close to the

wood surface, pupate and emerge from June through August. If conditions are not

favorable, immature larvae continue to feed and mine throughout the summer and pupate

the following spring, completing their life cycle in two years, although 3-5 year life cycles

have been observed (Raske 1972). Newly-emerged adults engage in a 3-10 day period of

maturation feeding on conifer foliage and shoots before reproducing (Rose 1957; Raske

1972), and return to feed on foliage and shoots throughout their life (Raske 1972).

' Author to whom correspondence should be addressed.

Current address: Department of Entomology, University of California-Riverside, Riverside

CA 92521, USA.

196 J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

Table 1

Species of Monochamus found in Canada, their distribution and host plants. Compiled

from Linsley and Chemsak (1984) and Gosling and Gosling (1976).

Species Distribution Host Genera

*M. carolinensis Eastern United States and Canada to Texas Pinus

(Olivier) and Minnesota

*M. titillator (F.) Eastern United States and Canada to Texas Pinus, Abies, Picea

and North Dakota

*M. scutellatus Newfoundland to Alaska, south to California Pinus, Abies, Larix,

(Say) and east to North Carolina Picea

M. obtusus Casey Washington, British Columbia and Idaho to —— Pinus,

California Pseudotsuga, Abies

M. marmator Southeastern Canada to North Carolina and Abies, Picea

Kirby the Great Lakes

*M. mutator Lake Superior region of Michigan, Pinus, Picea, Abies,

LeConte Minnesota, Ontario and Quebec Larix

*M. clamator California to British Columbia, Rocky Pinus, Abies,

(LeConte) Mountains and Great Basin to Southern Pseudotsuga

Arizona and Honduras

M. notatus Eastern North America to South Carolina Pinus, Picea,

(Drury) west to Montana and British Columbia Pseudotsuga, Abies

*Known vector of the pinewood nematode, Bursaphelenchus xylophilus (Steiner and

Buhrer) Nickle (Linit 1988; Vallentgoed 1991).

Differential ability of male whitespotted sawyers to defend territory at the breeding site

(host logs) may cause a high degree of variation in male mating success (Hughes 1979,

1981). Hughes and Hughes (1987) found that large-diameter trees are more attractive than

small-diameter trees and that females preferred the large-circumference portions of the

bole. They hypothesized that large host logs would produce high quality brood in high

numbers and consequently were preferred ovipositon sites. The sides of fallen logs are

preferred oviposition sites (Rose 1957; Raske 1972; Cerezke 1975), with M. scutellatus

laying eggs in the ratio of 10:3:1 on the sides, top and bottom of horizontal host logs

respectively (Raske 1972).

In 1999, peak adult flight activity for M. scutellatus, in the Okanagan Valley of British

Columbia occured between 17-31 August (McIntosh et al. 2001). In M. clamator

(LeConte)’ males emerged first and peak male emergence preceded peak female

emergence (Ross 1966). Similarly male M. a/ternatus Hope emerged earlier than females

(Togashi and Magira 1981). This protandry, the emergence and reproductive maturation of

males in advance of females (Wiklund and Fagerstrém 1977), is apparently an adaptive

trait (Thornhill and Alcock 1983) that allows early-emerging males to find and defend

preferred oviposition sites (Hughes 1979, 1981), where they await the arrival of females.

We report the results of observations in Ft. Nelson, British Columbia, on M.

scutellatus, demonstrating: 1) male and female preference for resting and ovipositing on

the sides of horizontal host logs; 2) differential mobility and reproductive behavior over

time; and 3) change in the sex ratio over time. We propose some testable hypotheses to

explain our observations.

* Linsley and Chemsak (1984) note that the species designated as Monochamus maculosus

(Haldeman) by Ross (1966) is now considered to be M. clamator (LeConte).

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 197

MATERIALS AND METHODS

Observations were made on 30 June, 9, 21 and 30 July and 5 August 2000 between

1200-1600 h in the logyard of Slocan Forest Products Ltd. Tackama Division, in Ft.

Nelson, British Columbia. The logs inspected were predominately decked white spruce,

Picea glauca (Moench) Voss, black spruce, P. mariana (Mill) B.S.P., subalpine fir, Abies

lasiocarpa (Hook.) Nutt., and lodgepole ping, Pinus contorta var. latifolia Engelmann.

Whenever possible the entire surface of a log was inspected. This was accomplished by

selecting logs that protruded from the end of the log deck. All observed beetles were

identified and sexed in the logyard using elytral and antennal characters (Linsley and

Chemsak 1984). Sample logs were randomly chosen, but inaccessible logs from the

bottom and centre of log decks were excluded. During each visit logs were sampled until

at least 50 beetles were observed (>10 logs sampled/visit). After sampling a log, the

investigator then moved to a log in another location (minimum 2-3 m distant) to reduce the

possibility of observing the same beetle more than once.

Logs were divided so that the sides, top and bottom were each equally represented by

1/3 of the available surface area (i.e. each side equaled 1/6 of the available surface).

Behavior was categorized as stationary, mobile or copulating. For females, an additional

category, oviposition, was characterized as chewing an oviposition niche, or ovipositing in

one. Rose (1957) observed that eggs are not deposited in all oviposition niches; however

we did not discriminate between chewing an oviposition niche and ovipositing. Stationary

beetles were immobile and solitary. Mobile beetles were walking or running on the bark;

their location when first observed was recorded. Copulating beetles were either stationary

or mobile, and were defined as any pair in which the female was mounted by a male,

unless the female was chewing an oviposition niche. In these cases the female was

recorded as ovipositing and the male as copulating.

The Chi-square goodness of fit test was used to test the null hypotheses that male and

female beetles and oviposition niches were randomly distributed on host logs. Chi-square

contingency table analysis was used to test the hypotheses that male and female behavior

and sex ratio were independent of date of observation. In all cases, a=0.05.

RESULTS AND DISCUSSION

Sixty-six and 65 percent of male and female M. scutellatus, respectively, were found

on the sides of logs (Fig. 1), and 58 percent of oviposition also occurred on the sides of

logs (Fig. 1). These results support observations of preferential oviposition on the sides of

horizontal logs by M. scutellatus (Rose 1957; Raske 1972; Cerezke 1975) and indicate that

all types of activity occur mostly on the sides of the logs. Oviposition on the sides of

horizontal host logs may represent a trade-off, which minimizes mortality of eggs and

young larvae from desiccation on the top of logs (Rose 1957), and slow development that

would occur in the cool lower portion of logs (Raske 1972). Ross (1966) allowed M.

clamator to oviposit on ponderosa pine bolts in May and then stored the logs in the shade

or in full sunlight until October of the same year. Of 67 larvae found under the bark of the

shaded bolts, only one had bored into the wood, whereas 22 of the 47 larvae found under

the bark of the bolts in full sunlight had bored into the wood; this developmental state

would enhance their chance of surviving the winter (Raske 1972). Although we did not

record the direction of exposed sides of logs, Post and Werner (1988) observed preferential

oviposition by M. scutellatus on the south facing sides of decked white spruce logs in

198 J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

Location of Solitary or Copulating Beetles Females Ovipositing

or Chewing Niche

70 70 70

60 MALES 60 FEMALES 60

x? = 108.2 x? = 49.6 x? = 6.97

50 df = 2, N = 222 50 df =2,N=107 50 df=2,N=26

P<0.001 P<0.001 0.025 <P < 0.05

Percent observed

Bas

(jo)

30 30 30

20 20 20

10 10 10

0 0 0)

Side Top Bottom Side Top Bottom Side Top Bottom

Figure 1. Locations of male and female Monochamus scutellatus observed on horizontal

host logs and female M. scutellatus observed chewing oviposition niches or ovipositing on

horizontal host logs in the Slocan Forest Products, Tackama Division logyard, Ft. Nelson,

British Columbia. Side, top and bottom each equal 1/3 of the available surface area.

Alaska, a behavior that would maximize exposure of larvae to solar-heat in the short

northern summer.

The observed sex ratio of M. scutellatus on host logs was always male-biased, but

changed significantly over time to favor males four-fold over females by 5 August (Table

2). These observations do not agree with Hughes (1979) who found mostly female M.

scutellatus on host logs. In M. clamator the sex ratio of 164 emergent beetles from fire-

killed ponderosa pine was 1.1 males per female (Ross 1966). One possible explanation for

our observations is that because of the high metabolic cost of oviposition, females spend a

significant portion of their time feeding in the crowns of trees, and thus are found less

frequently on logs than males, which remain to guard their territory (Hughes 1979, 1981).

Although female M. scutellatus live approximately 40 days and males 30 days in the

laboratory (Raske 1972), the costs of oviposition may cause females to die sooner than

males in the field, resulting in the sharp rise in male to female sex ratio in August (Table

2). In northern Ontario males were observed earlier in the afternoon than females (S.

Peddle, 2001, 256 Yorkshire St. N., Guelph, Ontario, NIH 5C4, personal

communication). Conversely, Hughes (1979) reports that few beetles were seen before

1400 h and most of these were females. It is possible that our observed bias in male sex

ratio is confounded by sampling time.

Table 2

Sex ratio of Monochamus scutellatus observed on host logs in the Slocan Forest Products,

Tackama Division logyard in Ft. Nelson, British Columbia. Significant change in sex ratio

with time, y7=12.7, d.f.=4, 0.025<P<0.01.

Date N Sex ratio

males/female

30 June 2000 50 2.6

9 July 2000 80 14

21 July 2000 73 Ds

30 July 2000 75 1.3

5 August 2000 5] 4]

All dates 329 1.9

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 199

The observed proportions of beetles engaged in various behaviors changed

significantly with time for both sexes (Fig. 2). The proportions of beetles copulating

peaked on 9-July and then decreased to zero one month later. Peak oviposition by females

was observed 12 days after the peak in copulation; oviposition persisted at the final

observation on 5 August. It has been demonstrated that some cerambycids prefer specific

host plants (i.e. tall and conspicuous host plants, see Hanks (1999) and references therein).

The high proportion of stationary males early in the summer, except for the peak

copulation period in early July, is consistent with the hypothesis that males secure

territories in preferred oviposition sites and wait for females. When a female landed and

approached within 2-3 cm, a male would dash toward and rapidly mount her. We observed

copulation at all times between 1200-1600 h, whereas Hughes (1979) did not observe

copulation before 1400 h. We hypothesize that the increase in male mobility on 5 August

represents a change in reproductive strategy in late summer, when females have become

scarce and therefore are less likely to enter a given male’s territory. Switching from a

territorial to roaming reproductive strategy could increase the likelihood of contacting and

copulating with a female.

MALES FEMALES

x? = 54.5, df= 8, P<0.001 X? = 33.3, df= 12, P< 0.001

. Stationary = Stationary

40 40

20 20

5 0 )

Be Mobile dl Mobile

(3)

Q 40 40

@ 20 20

c ;

WY)

2 Copulatin Copulati

O 60 p g - opulating

O 40 40

@ 20 20

0 )

ae a) a

Ry yp ys SS Ss

Ovipositing

S 9 ~ 29

ue) v is) Le

@ ss 2)

ey SS SS we

Figure 2. Proportions of observed behavior of male and female Monochamus scutellatus

on horizontal host logs from 30 June — 5 August, 2000, Slocan Forest Products, Tackama

Division logyard, Ft. Nelson, British Columbia.

200 J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

ACKNOWLEDGEMENTS

We thank Sarah Butler and Stuart Campbell for assistance, Peter de Groot, Dave

Moore, James Burns and Dean Morewood for critical reviews and for helpful comments.

This research was supported by the Natural Sciences and Engineering Research Council of

Canada, the Science Council of British Columbia, Forest Renewal British Columbia, the

Canadian Forest Service, Abitibi Consolidated Forest Products Inc., Ainsworth Lumber

Co. Ltd., B.C. Hydro and Power Authority, Bugbusters Pest Management Inc., Canadian

Forest Products Ltd., Gorman Bros. Ltd., International Forest Products Ltd., Lignum Ltd.,

Manning Diversified Forest Products Ltd., Millar Western Forest Products Ltd., Phero

Tech Inc., Riverside Forest Products Ltd., Slocan Forest Products Ltd., Tembec Forest

Industries Ltd., TimberWest Ltd., Tolko Industries Ltd., Weldwood of Canada Ltd., West

Fraser Mills Ltd., Western Forest Products Ltd. and Weyerhauser Canada Ltd.

REFERENCES

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sawyer, Monochamus scutellatus. Canadian Journal of Forest Research 7: 232-240.

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Part II, The Subfamilies Lepturinae and Lamiinae. The Great Lakes Entomologist 10: 1-37.

Hanks, L.M. 1999. Influence of the larval host plant on reproductive strategies of cerambycid beetles.

Annual Review of Entomology 44: 483-505.

Hughes, A.L. 1979. Reproductive behaviour and sexual dimorphism in the white spotted sawyer,

Monochamus scutellatus (Say). The Coleopterists Bulletin 33: 45-47.

Hughes, A.L. 1981. Differential male mating success in the white spotted sawyer, Monochamus scutellatus

(Coleoptera: Cerambycidae). Annals of the Entomological Society of America 74: 180-184.

Hughes, A.L. and M.K. Hughes. 1987. Asymetric contests among sawyer beetles (Cerambycidae:

Monochamus notatus and Monochamus scutellatus). Canadian Journal of Zoology 65: 823-827.

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J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 201

The cabbage seedpod weevil, Ceutorhynchus obstrictus

(Coleoptera: Curculionidae) - a review

H.A. CARCAMO!

AGRICULTURE AND AGRI-FOOD CANADA, LETHBRIDGE RESEARCH CENTRE,

PO BOX 3000, LETHBRIDGE, AB, CANADA TI1J 4B1

L. DOSDALL, M. DOLINSKI

ALBERTA AGRICULTURE FOOD AND RURAL DEVELOPMENT,

304 J. G.O0' DONOGHUE BUILDING, 7000 - 113 STREET,

EDMONTON, AB, CANADA T6H 5T6

O. OLFERT

AGRICULTURE AND AGRI-FOOD CANADA, SASKATOON RESEARCH CENTRE,

107 SCIENCE PLACE, SASKATOON, SK, CANADA S7N 0X2

and J.R. BYERS

AGRICULTURE AND AGRI-FOOD CANADA, LETHBRIDGE RESEARCH CENTRE,

PO BOX 3000, LETHBRIDGE, AB, CANADA T1J 4B1

ABSTRACT

The cabbage seedpod weevil, Ceutorhynchus obstrictus (Marsham), which has recently

become established in southern Alberta, is a serious pest of oilseed rape (Brassica napus

L.) in Europe and the USA and poses a major threat to the economic sustainability of

canola production in western Canada. This paper reviews the biology and control of this

pest and identifies future research needs. Control strategies in Europe and the USA have

so far relied on insecticides because no cultural or biological control methods have been

successful. Research on plant resistance is in progress at several research centres and

could provide the long term solution. Several parasitoid species are known to suppress

populations of the weevil in Europe and are candidates for biocontrol programs in North

America. Current research priorities in western Canada are to quantify the effects of

weevil densities on canola seed yield, to establish economic thresholds and to design

control strategies that integrate chemical, cultural and biological controls. Research

programs should be established to screen a wide range of Brassica germplasm to identify

sources of resistance for use in developing resistant cultivars for western Canada.

Research on the overwintering ecology and seasonal activity of this weevil is needed to

model how its range is likely to expand to other canola growing regions of Canada and to

enable forecasting of outbreaks.

Key words: Brassica, oilseed rape, Ceutorhynchinae, Ceutorhynchus assimilis,

Ceutorhynchus obstrictus, canola insect pests

' To whom correspondence should be addressed.

202 J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

INTRODUCTION

The cabbage seedpod weevil, Ceutorhynchus obstrictus (Marsham) (Coleoptera:

Curculionidae) was first detected in southern Alberta in 1995 (Butts and Byers 1996). By

2000, the weevil had become a major pest of canola” throughout southern Alberta (Dosdall et

al. 2001) and had spread eastward into adjacent areas of Saskatchewan (Fig.1).

Ceutorhynchus obstrictus poses a serious threat to the canola industry in western Canada and

has prompted provincial and federal entomologists to initiate research programs to develop

effective management strategies. This paper reviews the history and biology of the cabbage

seedpod weevil, its damage, and strategies for its control. We also propose research priorities

that will enable more effective and sustainable management of this pest.

Figure 1. Distribution of the cabbage seedpod weevil in Canada as of 2000.

References: 1) McLeod 1953; 2) Philips 2000; 3) Dosdall et al. 2001; 4) Olfert unpublished

data; 5) Brodeur et al. 2001. Photo by Eric Kokko, AAFC, Lethbridge, AB

* Canola is the term used for cultivars of oilseed rape (Brassica napus L. and B. rapa L.)

that are low in erucic acid and glucosinolate; attributes desirable for the production of food

grade vegetable oil and livestock meal.

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 203

HISTORY

As recommended by Colonnelli (1990, 1993) the name C. obstrictus Marsham will be used

in this paper instead of the synonym, C. assimilis Paykull, used in most previous publications

on this species. The cabbage seedpod weevil (CSW), also known as the turnip seed weevil or

seed weevil in Europe, has been recognized as a pest of crucifers from the Mediterranean to

Scandinavia since the beginning of the 19th century (Bonnemaison 1957). In North America, it

was first recorded near Vancouver, British Columbia tn 1931 (Baker 1936; McLeod 1953) and

is now well established in the interior in both the Okanagan and Creston valleys (Philips

2000). By 1946 it had spread throughout the Pacific North-West and California (Hagen 1946,

Crowell 1952). It is well established in Georgia (Buntin 1990) and Tennessee (Boyd and Lentz

1994) and probably now occurs throughout most of the USA. In 2000 it was found in Quebec

(Brodeur ef al. 2001).

BIOLOGY

Ceutorhynchus obstrictus is univoltine with the adults overwintering under leaf litter in

treed areas, shelterbelts and field margins (Dmoch 1965a). A chill period of about 16 weeks at

4°C is required to break diapause (Ni e¢ a/. 1990). The adults, which are strong fliers, disperse

from the overwintering habitats in spring when air temperatures reach 15°C (Ankersmit and

Nieukerken 1954; Dmoch 1965a). They feed on the buds and flowers of various crucifers for

several weeks before they start to oviposit (Doucette 1947; Dmoch 1965a; Ni et al. 1990). In

southern Alberta they are abundant on early flowering cruciferous weeds, especially flixweed

(Descurania sophia L. Webb) and hoary cress (Cardaria draba L. Desv.), until canola begins

to bloom. Little ovarian development occurs below 10°C or above 25°C (Ni et al. 1990).

Weevil numbers peak when the host crop begins flowering (Dmoch 1965b; McCaffrey et al.

1986). The most common host crops are cultivated crucifers, including canola, other oilseed

rape, cole crops (e.g. B. oleracea L.) and brown mustard (B. juncea L). Yellow (or white)

mustard, Sinapis alba L. is not attacked (Doucette 1947). Females lay eggs singly into young

pods through feeding punctures and usually only one egg is laid per pod unless weevil

densities are high. After oviposition the females brush the pod with the tip of their abdomen,

apparently to apply an oviposition deterrent pheromone (Kozlowski ef a/. 1983). The larvae

undergo three instars and consume three to six seeds each. When mature they chew an exit

hole in the wall of the pod, drop to the ground and pupate in the soil. The new generation

adults emerge from 9-30 days after exit from the pods depending on temperature (Hanson er

al. 1948; Bonnemaison 1955, 1957; Dmoch 1965a). In southern Alberta we (L.D.) have found

that this sometimes takes only 7-10 days. The entire development from egg to adult usually

takes 4-6 weeks (Bonnemaison 1957). The new adults can disperse several km or more in

search of food, especially late maturing crucifers, to accumulate fat reserves before finding

overwintering habitats in early fall (Doucette 1947).

DAMAGE, ECONOMIC THRESHOLDS AND SAMPLING

Cabbage seedpod weevils can significantly reduce the seed yield of canola in several ways.

Feeding by the overwintered adults on buds and flowers in the spring and early summer causes

blossom blasting and pod abortion which may reduce the yield by up to 14% (Coutin ef al.

1974). However, under good growing conditions canola plants can compensate for up to 60%

loss of buds and flowers (Williams and Free, 1978, 1979; Free et al. 1983). Reduced yield can

also occur as a result of interaction between the feeding or oviposition activity of the CSW

adults and other insect pests. In Europe, CSW feeding punctures are used for oviposition by

the pod midge, Dasyneura brassicae Winn. (Free et al. 1983). Seed losses are considerably

204 J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

higher when both pests occur together and as a consequence, the economic threshold for CSW

adults in England is only one per plant (Free ef a/. 1983). The feeding punctures can also be

used by other small insects, such as thrips, to gain access to the seeds; without the weevils,

thrips feed only on the surface of the pods and cause little damage (K.M. Fry, 2000, Alberta

Research Council, personal communication). This interaction might potentially be a problem

in those regions where thrips are sometimes abundant in canola during flowering. Under moist

conditions a reduction in seed yield and quality can also result from fungal pathogens that gain

entry through the feeding punctures or larval exit holes.

The principal damage caused by CSW occurs during the larval stages (McCaffrey ef al.

1986; Buntin 1999). Depending on seed size, three to six seeds are consumed by each larva

(Dmoch 1965a) or about 20 to 30% of the seeds in each pod. With high weevil densities two to

three larvae may infest a pod and consume most of the seeds. Additional losses occur at

harvest because infested pods ripen prematurely and tend to shatter prior to or during harvest.

Late seeded or late maturing canola can be seriously damaged in late summer or early fall

by new generation adults feeding through the pod wall on the immature seeds. Buntin ef al.

(1995) found that in Georgia and Idaho feeding by adults reduced the weight of punctured

seeds by about 16% and the oil content by about 2%. The incidence of damaged seed ranged

from 8 to 17% in untreated fields in Georgia and 5 to 10% in fields treated with insecticide in

Idaho. Although the overall yield loss was less than 2%, a 40% decrease in germination of the

damaged seed and a high incidence of abnormal seedlings, would be of concern for certified

seed producers.

The relationship between weevil densities and yield loss has been little studied although

this relationship is necessary for establishing economic thresholds. Studies in Scandinavia

showed a clear negative relationship between weevil densities and yield (Tulisalo et al. 1976;

Sylven and Svenson 1975). In Tulisalo’s cage study, two weevils per plant reduced yield by

50% and they estimated that one weevil per four plants warranted the use of an insecticide.

They found that at high weevil densities the plants attempted to compensate by producing

more pods, but this was more than offset by a reduction in the average seed weight.

The presence of other pests can also affect the economic threshold for CSW. Free et al.

(1983) determined that, in England, densities below one weevil per plant caused pod

infestation rates of less than 26% and on their own did not warrant control. However, in the

presence of the pod midge, Dasyneura brassicae, losses were much higher and control at

lower weevil densities was warranted. Other studies in France (Lerin 1984) and the USA

(Buntin 1999) have also found that, because of plant compensation, there is little yield loss at

pod infestation levels below about 25%. Although the CSW has been a serious pest of winter

canola in the US Pacific Northwest, no economic threshold has been established. However,

preliminary studies from Idaho indicate that three to six weevils per 180° sweep with a

standard 38 cm diameter sweep net warrants control because at these population levels yields

in unsprayed plots were 15 to 35% lower than in sprayed plots (McCaffrey et al. 1986). A

similar threshold of three to four weevils per sweep is being used in southern Alberta (Dosdall

et al. 2001) until results from current cage and plot studies are available.

Sampling methods vary with the objective of the investigation. For population monitoring,

a sweep net is normally used. Dmoch (1965a,b) determined that 4 samples of 25 sweeps each

estimated weevil populations with adequate accuracy. In plot insecticide trials, the sweep net is

also the usual sampling method and the number of sweeps per plot can be as low as six to eight

in small plots (Buntin 1999). When the crop is fully podded and sweeping becomes difficult

other methods such as dislodging the weevils into buckets or pans have been used (Brown et

al. 1999). Yellow pan traps have been used to study the seasonal pattern of activity of seasonal

activity and to monitor weevil arrival in fields (Bonnemaison 1957). Flight intercept traps have

also been used to provide information about CSW spatial distribution and phenology

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 205

(Ferguson et al. 2000). As suggested by Dolinski (1979) we have found that pitfall traps are

useful for studying the arrival of adults at, and departure from, overwintering habitats. No

studies relating the results of the various trapping or sweeping methods to actual weevil

densities per plant have been published. Because the weevils are concentrated on the buds and

flowers in the uppermost part of the crop canopy, ongoing studies (H.A.C.) are finding the

efficiency of sweep net sampling to be about twice the 10% reported for lygus bugs (Wise and

Lamb 1998).

CONTROL STRATEGIES

Biological control. In Europe, and those parts of North America where it has been

established for some time, the CSW is host to a number of parasitoids. Surveys in Washington

(Hanson et al. 1948; Doucette 1948), Oregon (Doucette 1948), California (Carlson et al.

1951), and British Columbia (McLeod 1953) found 11 parasitoid species associated with this

weevil. The pteromalid wasps, 7richomalus perfectus (Walker) [syn. T. fasciatus (Thomson)]

and Mesopolobus morys L. (syn. Xenocrepis pura Mayr) were the most abundant parasitoids.

Trichomalus perfectus and M. morys were also important parasitoids in northern Idaho,

although the eulophid, Necremnus duplicatus Gahan was present in substantial numbers

(Doucette 1948; Walz 1957). Recently Harmon and McCaffrey (1997) found that the

introduced European braconid, Microctonus melanopus Ruthe, reduced survival of

overwintering adult weevils in Idaho and Washington, with parasitism levels as high as 70%.

In Europe, many parasitoids are known to attack the CSW (Dolinski 1979; Herting 1973;

Kuhlmann and Mason 1999) with the most common being the braconids M. melanopus and

Diospilus oleraceus Haliday, and the pteromalids M. morys and T. perfectus (Kuhlmann and

Mason 1999, Kuhlmann et al. 2001).

Chemical control. Several insecticides control CSW effectively, although none are

currently registered in Canada (Dosdall et al. 2001). Pyrethroids such as deltamethrin and

alphacypermethrin are used in Europe for control of adults when the crop is at the early

flowering stage (Alford et al. 1996). Parathion applied at the end of flowering was reported to

control the larval stage of CSW in the Pacific NW of the USA and was recommended over

endosulfan which was more expensive and less effective against adults (McCaffrey ef al.

1986). In Georgia, Buntin (1999) found that the pyrethroids, bifenthrin, esfenvalerate,

permethrin, and zetacypermethrin controlled CSW on winter oilseed rape more effectively than

the other insecticides, including methyl parathion and endosulfan, currently registered in the

USA. However, only treatment with esfenvalerate increased yield relative to untreated plots

and two applications were required (Buntin 1999). However, preliminary results (H.A.C. &

L.D) indicate that in most fields only one application of insecticide will be needed for control

of CSW on spring canola in Canada.

Insecticide applications should, ideally, be timed to spare parasitoids and minimize

disruption of biological control. Research in the UK has shown that 7. perfectus, an

ectoparasite of CSW larva, arrives in rape fields towards the end of flowering about two weeks

after the weevil (Alford ef al. 1996). Therefore, insecticide applications applied at early

flowering should largely spare the parasitoid (Murchie et a/. 1997). Buntin (1998) found that

the use of esfenvalerate during bloom indirectly reduced T. perfectus numbers because of a

reduction in the number of available hosts, but a greater proportion of the remaining host

larvae were attacked. Another recent UK study (Ferguson et a/. 2000) found that CSW tend to

be spatially aggregated within fields and it might be possible to spot-spray such areas, thereby

reducing mortality of beneficials. Earlier studies had shown that at low populations the weevils

are aggregated along field edges (Free and Williams 1978). As part of an integrated pest

management program developed in France in the 1970's, it was found that spraying only the

206 J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

field borders usually gave adequate control of CSW and enhanced parasitism in the rest of the

field (Jourdheuil et al. 1974).

Cultural control. Cultural control methods for CSW have received little attention. There

are no published studies on effects of rotation, intercropping, planting date or seeding rate.

Because the adults disperse widely (Kjaer-Pedersen 1992), crop rotation is unlikely to reduce

damage. Intercropping should be investigated because it has been shown that interplanting of

canola with barley provides some protection against crucifer specialists such as flea beetles

(Butts et al. 1999). Mixed planting with non-host crops might interfere with the chemical host

finding cues used by CSW (Evans and Allen-Williams 1992). Late planted fields and

experimental plots have been observed to largely escape weevil damage in southern Alberta,

however, too late a seeding date exposes the crop to damage by new generation adults.

Although an increase in seeding rate of canola can counteract damage by root maggots, Delia

spp., (Dosdall et al. 1998) the effect of seeding rate on CSW is unknown.

Buntin (1998) investigated trap cropping as a method of managing CSW in winter oilseed

rape. He used 0.35 ha plots with the peripheral 4.9 m planted with a spring cultivar (trap crop)

and the rest planted with a conventional winter cultivar (main crop). Both were planted at the

same time in the autumn and as a consequence the spring cultivar flowered several weeks

earlier the following spring. Although weevils were more numerous in the trap crop, their

control with esfenvalerate did not prevent damage and loss of yield in the unsprayed main

crop. However, he speculated that trap cropping might work better at the field scale. Buechi

(1990) also failed to reduce losses in oilseed rape (B. napa) by using turnip rape (B. rapa) as

the trap crop. However, he did not spray the trap crop to kill the CSW adults which apparently

preferred to oviposit in the oilseed rape. Ongoing studies in Alberta (Carcamo et al. 2001)

using an earlier flowering Polish cultivar (B. rapa) as the trap crop and a later flowering

Argentine cultivar (B. napa) planted at the same time or staggered planting of the same

cultivar, with the trap crop border being planted | to 2 weeks earlier, show that invading CSW

adults are highly concentrated in the trap strips. Growers may be able to prevent damage to the

main crop if the trap strip is sprayed before the CSW disperse into the later flowering main

crop. Trap cropping has the potential to substantially reduce insecticide use, thereby lowering

production costs and sparing nontarget species, especially pollinators and natural enemies.

Host plant resistance. The development of cultivars of canola with genetic resistance to

the CSW would provide the ultimate solution. In a laboratory assay, Harmon and McCaffrey

(1997) observed reduced feeding and oviposition on excised pods of a B. rapa line compared

to two B. napus lines in choice tests. However, the differences were less pronounced in no-

choice tests and might not be meaningful in the field.

Because yellow mustard (S. a/ba) is immune to CSW attack (Doucette 1947), hybrids of S.

alba and B. napus have been produced with the expectation that these might be resistant to

CSW (Brown ef al. 1997). Although the hybrids were attacked by CSW, fewer larvae

completed development in the hybrids than in the B. napus parent (McCaffrey et al. 1999).

The authors attributed the effect to high concentrations of p-hydroxybenzy! glucosinolate

inherited from the S. a/ba parent. An alternative to developing such hybrids for control of

CSW is to develop cultivars of S. a/ba that produce canola quality oil. Research currently well

underway at the Saskatoon Research Centre (Agriculture and Agri-Food Canada) has made

good progress to developing canola quality S. a/ba lines that are better adapted than canola to

the brown soil zone of Alberta and Saskatchewan and hopefully have retained the resistance to

CSW.

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 207

RESEARCH PRIORITIES

Several basic and applied questions about the ability of the CSW to adapt to the Canadian

prairies need answers to aid the development of sustainable management strategies. In an

earlier review (Dolinski 1979) it was speculated that CSW had limited potential to become a

pest of canola in western Canada because of the cold climate and apparent lack of suitable

overwintering habitats. However, now that we know that it can survive here, at least in some

areas, research is needed on its overwintering ecology to determine its likely range extension.

Systematic surveys need to be conducted to track its spread and identify those areas where

populations are increasing most rapidly. The phenology of CSW also needs to be studied in

more detail to determine if it is, or can readily become, synchronized with that of canola in the

more traditional production areas in the parkland ecoregions in the three prairie provinces and

the Peace River region of Alberta and BC.

Of more immediate concern is the development of economic thresholds specific to the

southern prairies that will enable canola growers to avoid unnecessary spraying. Detailed cage

and plot experiments are required to objectively relate CSW density to seed yield and quality.

Implementation of the established economic thresholds will depend on the adoption of a

standardized sampling protocol. Sweep net sampling is the simplest and, although imperfect,

by far the most practical method. However, conversion factors appropriate for each crop stage

will be needed to relate the sweep net catches to actual CSW densities.

Interactions with other insect pests that may occur at the same time, such as lygus bugs,

thrips, bertha armyworm and diamondback moth need to be investigated so that rational

economic thresholds and IPM strategies for management of the insect pest complex of canola

can be recommended. Insecticides known to be effective, such as pyrethroids, need to be

registered, but registration should take into consideration the impact on pollinators and natural

enemies. This is important because to control CSW the insecticide will probably have to be

applied during flowering.

Studies should also be undertaken to further assess cultural control methods such as

planting date (e.g. fall-planted spring canola), intercropping and trap cropping. Integration of

appropriate cultural practices with longer-term strategies such as biological control and

resistant cultivars should ensure the environmental and economic sustainability of the canola

industry in Canada.

ACKNOWLEDGEMENTS

We thank Andrea Kalischuk for constructive comments on an earlier draft of this article,

Kendra Grams for text processing and Erin Cadieu for graphic support. This paper is

contribution 387-01037 for the Lethbridge Research Centre.

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J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 211

Rearing the cranberry girdler Chrysoteuchia topiaria

(Lepidoptera: Pyralidae) on reed canary grass Phalaris

arundinacea (Festucoideae: Panicoideae)

SHEILA M. FITZPATRICK’, JANIS A. NEWHOUSE,

JAMES T. TROUBRIDGE and KAREN A. WEITEMEYER

AGRICULTURE AND AGRI-FOOD CANADA,

PACIFIC AGRI-FOOD RESEARCH CENTRE, PO BOX 1000, 6947 HIGHWAY 7,

AGASSIZ, BC, CANADA VOM 1A0

ABSTRACT

We report a method of rearing cranberry girdler Chrysoteuchia topiaria (Zeller), a pyralid

that is a serious pest of cranberry Vaccinium macrocarpon Aiton. Fertile eggs from field-

caught females were scattered on reed canary grass Phalaris arundinacea L. planted in

greenhouse flats (50 eggs/flat) kept under fluorescent lights at 16L:8D and temperatures of

22-30°C (day): 19-24°C (night). Under these conditions, survival from egg to adult was

28%. Progeny of these adults entered diapause after exposure to low light (ca. 0.5 lux) as

larvae. Diapause was broken by placing insects in the dark at 4.5-5.5°C for ca. 3 months,

but survival was very poor (8% from egg to adult).

Key words: laboratory colony, laboratory rearing, turfgrass, Vaccinium macrocarpon

Aiton, integrated pest management, subterranean webworm, sod webworm, diapause

INTRODUCTION

Cranberry girdler Chrysoteuchia topiaria (Zeller) is a serious pest of cranberries,

Vaccinium macrocarpon Aiton in North America (Smith 1903). It is also recognized as a pest

of grasses (Ainslie 1916) and coniferous seedlings (Kamm ef al. 1983). Cranberry girdler

belongs to the group of grass-infesting crambids (Pyralidae) commonly called sod webworms,

and is also known as the subterranean webworm (Tashiro 1987). This pest overwinters as

diapausing prepupae in the soil, and moths emerge from late May through early August

(Kamm et al. 1990). Mated females deposit several hundred eggs (Scammell 1917, Kamm

1973b) singly or in groups at the soil surface. Neonate larvae are fragile, remaining near the

surface where they feed on succulent tissue. Older larvae feed on crowns and roots, often

severing them. Cranberry girdler is usually reported to be univoltine (e.g., Kamm ef a/. 1990),

although moths observed flying in late August or September may represent a second

generation (Smith 1903, Fitzpatrick unpublished).

Most studies of biology, chemical ecology and integrated pest management of cranberry

girdler (summarized in Kamm ef a/. 1990) have been done in the field or have used insects

gathered directly from the field (e.g., McDonough and Kamm 1979), because cranberry girdler

is notoriously difficult to rear in the laboratory. The only report of successful rearing from egg

to adult comes from Roberts and Mahr (1986), who obtained at best 17% survival from egg to

adult on pinto bean diet at 16L:8D and 21°C.

Scammell (1917) complained that “some species of Crambinae defy all attempts to rear the

larvae’, noting that cranberry girdler was one of these. Our initial attempts to rear this insect

' To whom correspondence should be addressed.

22 J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

(Fitzpatrick unpublished) almost led us to conclude that he was right. Under our conditions,

girdler larvae did not survive on southwestern corn borer diet (Bioserve #F9763B; Bioserve,

Frenchtown, NJ), sod webworm diet (Bioserve #F954B) or on pinto bean diet (modified from

Shorey and Hale 1965) made in our laboratory. Only one of 200 larvae survived to the adult

stage on general insect diet (Bioserve #F9004). We also tried five species of grass reported to

be attractive to girdler larvae (Roland 1990): reed canary grass Phalaris arundinacea L.,

meadow foxtail A/opecurus pratensis L., red top Agrostis alba L., hard fescue Festuca ovina

var. Duriuscula (L.) Koch., and creeping red fescue Festuca rubra L. Of these species, only

reed canary grass sustained enough larvae for a colony. Here we report our method of rearing

cranberry girdler on reed canary grass.

MATERIALS AND METHODS

Source of insects. A modified handheld vacuum (Bioquip, Gardena, CA) was used to

collect mated female moths from commercial cranberry farms in Richmond and Pitt Meadows,

British Columbia, in June and July 2000. Female moths were placed in plexiglass cages (0.3 x

0.3 x 0.3 m) in the laboratory at 16L:8D with temperatures ranging from 22-30°C (day): 19-

24°C (night). Eggs, which are simply released from the ovipositor and dropped, were collected

on sheets of wax paper or aluminum foil. If fertile, eggs changed colour from yellow to orange

within 5-8 days of oviposition.

Rearing Conditions. Reed canary grass Phalaris arundinacea L. was seeded into a 50:50

mixture of potting soil and vermiculite in greenhouse flats (53 cm x 27.5 cm x 6.5 cm deep).

The grass was watered, fertilized with 15-30-15 (N-P-K) as required, and maintained under

fluorescent lights at 16L:8D in the laboratory or in the greenhouse, depending on which site

was available. (At the time of this study, we had limited facilities.) In the laboratory,

temperatures ranged from 22-30°C (day): 19-24°C (night). In the greenhouse, temperatures

ranged from a high of 23°C during the day to a low of 15°C at night. All temperatures were

recorded by Hobo® dataloggers (Onset Computer Corp., Bourne, MA). The reed canary grass

grew for 10-60 days before fertile eggs were scattered onto the flats, and was kept trimmed to

ca. 5-7 cm tall. Patches of grass killed by girdler larvae were reseeded. Girdler prepupae in

cocoons were usually left in flats, which were placed in cages of various dimensions to contain

emerging moths. Cages were made from screen (0.5 mm mesh) and PVC irrigation pipe, or

plexiglass with screened openings. Flats from one rearing were placed in small controlled-

environment chambers at 16L:8D with temperatures ranging from 21.5-24.5°C (day): 15-

16.5°C (night). Light intensity, measured with a Hobo® datalogger, was 28-60 lux in the

controlled-environment chambers. Light intensity in the laboratory and greenhouse was not

measured.

We maintained some mature larvae and prepupae individually to produce unmated moths

for fecundity studies (reported elsewhere). Mature larvae and prepupae in cocoons were

removed from flats of reed canary grass and placed individually in 30-ml clear plastic cups

containing a small amount of the moistened soil:vermiculite mixture and, if larvae were still

feeding, a plug of reed canary grass. Insects in plastic cups were placed in small controlled-

environment chambers under the conditions described above.

Statistics. Some moths were weighed on the day of emergence using a Sartorius

microbalance (Sartorius Canada Inc., Mississauga, ON). To compare weights of adults, t-tests

were performed on raw data (Systat 8.0 1998). Weights are given as mean + standard error.

RESULTS

In mid-July 2000, a subset of 1550 fertile eggs from 131 field-collected females were

scattered on 31 flats (50 eggs/flat) of reed canary grass in the laboratory. About 3 weeks later,

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 13

small patches of dead grass could be lifted to reveal larvae feeding on the crowns and roots. As

feeding and larval development progressed, some larvae crawled out of flats containing only

dead grass. Most of these larvae were collected by hand and placed in flats containing live

grass. Most larvae finished feeding by the third week of August. From 21-30 August, an

uncounted number of girdler cocoons were removed from the flats and placed individually in

30-ml clear plastic cups containing the soil:vermiculite mixture. Cocoons recovered from the

flats were left unopened to promote survival of the prepupa or pupa within. Cups containing

cocoons were placed in the small controlled-environment chambers. Because we could not be

sure that we had recovered all cocoons from the flats of grass, the flats were saved and

maintained in the laboratory.

From | September to 27 October 2000, 268 males and 192 females emerged from the cups

and flats. These individuals completed development without diapause. A subset of 60 unmated

males and 60 unmated females was weighed. Males weighed 12.0+0.3 mg; females weighed

21.640.6 mg (f13.72, df=118, P<0.001). From 8 September through 8 November 2000, a

subset of 1900 fertile eggs from females used in fecundity studies was scattered on 19 flats of

reed canary grass (100 eggs/flat) kept in the greenhouse. After 8-12 weeks, all larvae that

could be found in the flats (570 larvae) were placed individually in 30-ml clear plastic cups

with a plug of reed canary grass rooted in soil. All cups were placed in empty flats stacked

(because space was very limited) in small chambers and repositioned every few days. In the

previous rearing, cups were not tightly stacked because there were fewer cups and more space.

The flats containing soil, dead grass and unrecovered cocoons were saved and also maintained

in small chambers.

By early January 2001, only 32 adults had emerged. Several cocoons were opened to reveal

diapausing prepupae. The measured light intensity reaching larvae inside cups in stacked flats

was ca. 0.5 lux, which was so low that larvae probably did not experience the 16-h

photoperiod as daylight. The light intensity experienced by larvae in cups at the top of the

stack would have been close to the measured light intensity in the small chambers: 28-60 lux.

We speculated that some larvae received enough light to continue their development without

diapausing, but the majority did not. To break diapause, we exposed the cups containing

cocoons and flats containing the remaining soil to simulated winter conditions of 4.5-5.5°C and

total darkness in the small chambers from 5 January until 16 April 2001. On 17 April,

photoperiod was set to 16L:8D and cups were arranged so that each received adequate light.

Over the next 10 days, temperatures were gradually stepped up to 22:16°C. The flats

containing the remaining soil were brought into the laboratory on April 17 (because there was

insufficient space in the small chambers) and exposed to 16L:8D and approximately 22:16°C.

From 4 May to 11 June 2001, 70 males and 61 females emerged from the flats of soil. No

moths emerged from the cups. Males weighed less than those of the previous generation

(9.7+0.2 mg; 5.47, df=128, P<0.001), as did females (18.140.7 mg; 3.88, df=119,

P<0.001). At least 90% of these moths were unmated at the time of weighing. Of the

individuals that never emerged from cups, 60 were pupae, 140 were prepupae, and the rest

died as larvae.

DISCUSSION

When reared on reed canary grass P. arundinacea in greenhouse flats under fluorescent

lights at a photoperiod of 16L:8D and temperatures of 22-30°C (day): 19-24°C (night), girdlers

developed from egg to adult without diapause in 6-10 weeks. Roberts and Mahr (1986)

reported developmental times of 10.4 weeks at 21°C and 7.6 weeks at 24°C for cranberry

girdler reared through one generation without diapause on pinto bean diet.

Earlier observations in 1999 showed that this insect can be reared on reed canary grass

214 J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

under the above conditions through two generations without diapause (Fitzpatrick,

unpublished). However, in late 2000, most of the progeny of non-diapausing girdlers entered

diapause after larvae were reared at 15-23°C for the first 8-12 weeks, then exposed to low light

intensity (ca. 0.5 lux) at 16L:8D and 15-24.5°C. Roberts and Mahr (1986) reared larvae at

16°C and at 21°C without triggering diapause. Our results suggest that diapause is triggered by

photophase experienced by larvae, and is facultative. The only other report on photoperiod in

relation to diapause in cranberry girdler comes from Kamm (1973a), who found that

diapausing prepupae developed more slowly under a 12-h than a 16-h photophase.

In our study, 28% of non-diapausing cranberry girdlers survived from egg to adult. We did

not quantify stage-specific mortality but we observed that late instars often crawled out of flats

that had been overwatered or were very dry or were full of dying grass. These mobile larvae

sometimes drowned under flats or escaped to corners of the laboratory and died. In the field,

late instars may be a dispersing stage.

The survival of diapausing girdlers was very low, only about 8% from egg to adult. We

suspect that the main reason for the additional mortality of diapausing girdlers was inadequate

moisture in the soil, particularly in the 30-ml cups, during the 3-month simulated winter. It is

also possible that larvae did not receive adequate food during development.

In both diapausing and non-diapausing groups, the average weight of newly emerged

females was at least 80% greater than that of males. Adults that emerged after ca. 3 months in

diapause weighed on average 17-20% less than adults of the previous, non-diapausing

generation. This difference in weight may represent a physiological cost of diapause or may

have resulted from insufficient food during larval development or dessication due to

inadequate moisture in the soil during diapause.

In conclusion, cranberry girdler can be reared without diapause in greenhouse flats planted

with reed canary grass (50 eggs/flat) and maintained under fluorescent lights at 16L:8D and

temperatures of 22-30°C (day): 19-24°C (night). Late instars and prepupae in cocoons can be

removed from soil in the flats and maintained individually until adult emergence.

ACKNOWLEDGEMENTS

We thank Sandy Uhazy, Minder Sidhu and Don Middleton for allowing us to collect

girdler moths on their cranberry farms, and Céline Maurice and Carlos Silva for excellent

technical assistance. We thank two anonymous reviewers for helpful comments on the

manuscript. This study was part of a larger one funded by AAFC Matching Investment

Initiatives, the British Columbia Cranberry Growers Association, Ocean Spray Cranberries

Inc., and the Cranberry Institute. This article is contribution # 656 from the Pacific Agri-Food

Research Centre, Agassiz, British Columbia.

REFERENCES

Ainslie, G.G. 1916. Notes on crambids. Journal of Economic Entomology 9: 115-118.

Kamm, J.A. 1973a. Role of environment during diapause on the phenology of Chrysoteuchia topiaria

(Lepidoptera: Pyralidae). Entomologia Experimentalis et Applicata 16: 407-413.

Kamm, J.A. 1973b. Biotic factors that affect sod webworms in grass seed fields in Oregon. Environmental

Entomology 2: 94-96.

Kamm, J.A., G.C. Fisher, L. M. McDonough and C.H. Shanks, Jr. 1990. Cranberry girdler as a pest of grass

seed, cranberry and nursery tree production in the Pacific Northwest, pp. 141-150. In: N.J. Bostanian, L.T.

Wilson and T.J. Dennehy (Eds.). Monitoring and Integrated Management of Arthropod Pests of Small Fruit

Crops. Intercept Ltd., Andover, Hampshire, United Kingdom.

Kamm, J.A., P.D. Morgan, D.L. Overhulser, L.M. McDonough, M. Triebwasser and L. N. Kline. 1983.

Management practices for cranberry girdler (Lepidoptera: Pyralidae) in Douglas-Fir nursery stock. Journal

of Economic Entomology 76: 923-926.

McDonough, L.M. and J. A. Kamm. 1979. Sex pheromone of the cranberry girdler, Chrysoteuchia topiaria

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 215

(Zeller) (Lepidoptera: Pyralidae). Journal of Chemical Ecology 5: 211-219.

Roberts, S.L. and D.L. Mahr. 1986. Development of cranberry girdler, Chrysoteuchia topiaria (Lepidoptera:

Pyralidae) in relation to temperature. The Great Lakes Entomologist 19: 85-90.

Roland, J. 1990. Use of alternative plant species as a monitoring tool for the cranberry girdler (Lepidoptera:

Pyralidae). Environmental Entomology 19: 721-724.

Scammell, H.B. 1917. The cranberry girdler. United States Department of Agriculture Bulletin No. 554.

Shorey, H.H. and R.L. Hale. 1965. Mass rearing of nine noctuid species on a simple artificial medium. Journal of

Economic Entomology 58: 587-594.

Smith, J.B. 1903. Insects injurious in cranberry culture. United States Department of Agriculture Farmer’s

Bulletin No. 178.

Systat 8.0. 1998. SPSS Science Marketing Department, SPSS Inc., 233 South Wacker Drive, 11th Floor,

Chicago, IL 60606-6307.

Tashiro, H. 1987. Lepidopteran pests: Family Pyralidae, pp. 70-90. In: H. Tashiro. Turfgrass Insects of the

United States and Canada. Cornell Unversity Press, Ithaca, New York.

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 I17

Monitoring the seasonal population density of Pandemis

pyrusana (Lepidoptera: Tortricidae) within a diverse fruit

crop production area in the Yakima Valley, WA

A. L. KNIGHT

YAKIMA AGRICULTURAL RESEARCH LABORATORY, AGRICULTURAL RESEARCH

SERVICE, 5230 KONNOWAC PASS RD., WAPATO, WA, UNITED STATES 98951

ABSTRACT

The population dynamics of Pandemis pyrusana (Kearfott) were studied in 60 contiguous

orchard blocks (154 hectares) of mixed fruit production situated in the Yakima Valley,

Washington. Grids of sex-pheromone-baited and liquid-food-baited traps were placed at a

rate of one trap of each type per 2 hectares. Trees within 50 m of each trapping location

were sampled for overwintering and summer generation larvae, and fruit injury prior to

harvest. Larvae from both generations were found in a low proportion of apple (Malus

domestica Borkh.), pear (Pyrus communis L.), and cherry (Prunus avium L.) orchards, but

not in the peach/nectarine (Prunus persica (L.)), apricot (Prunus armeniaca L.), or prune

(Prunus domestica L.) orchards. Larval densities between generations increased 5-fold in

apple and 10-fold in cherry and non-bearing apple. Parasitism of field-collected larvae by

tachinid parasitoids averaged 37% and 23% for each generation, respectively. Low levels of

fruit injury (<< 0.5%) by P. pyrusana were detected in only five apple and pear orchards.

Cumulative moth catch was 10-fold higher in sex-pheromone than food-baited traps. Moth

catch in both types of traps varied significantly among crops. In general, moth catches were

highest in apple and cherry. Cumulative moth catch in both trap types in apple and pear

during the first flight was weakly correlated with levels of fruit injury. In contrast, moth

catch during the second flight was not correlated with fruit injury. The observed low

predictive ability of traps was likely due to trap saturation and contamination with non-target

moths and a general dispersal of moths among orchards throughout the region. The capture

of female moths versus the total of both sexes caught in food bait traps did not improve the

prediction of fruit injury in apple or pear.

Key words: Pandemis, leafrollers, sex pheromone traps, food bait traps, fruit crops

INTRODUCTION

Pandemis spp. (Lepidoptera: Tortricidae) leafrollers are important direct pests of apple,

Malus domestica (Borkh.), from British Columbia to California (Newcomber and Carlson

1952; Madsen et al. 1984; Zalom and Pickel 1988). Two species of Pandemis overlap

geographically, within this range with P. /imitata Robinson predominating in northcentral

Washington and British Columbia and P. pyrusana (Kearfott) in the Yakima Valley,

Washington, Oregon, and California. Levels of fruit injury caused by P. pyrusana have

increased following the adoption of sex-pheromone-based mating disruption of codling moth,

Cydia pomonella L., and the concurrent decreased use of the broad-spectrum organophosphate

insecticides in these programs in Washington (Knight 1995) and California (Walker and

Welter 2001). Management of P. pyrusana has relied on either the use of organophosphate

insecticides, endotoxins of Bacillus thuringiensis Berliner, (Knight et a/. 1998) or the use of

sex pheromones for mating disruption (Knight and Turner 1999).

Management decisions for P. pyrusana in apple involve larval sampling in the spring prior

to bloom (Beers ef a/. 1993). However, this method is labor intensive and ineffective in

218 J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

detecting low-density populations. The use of sex pheromone-baited traps to monitor P.

pyrusana populations has been hampered by a poor relationship between moth catch and the

within-orchard pest density, lack of knowledge of the drawing range of these lures, and limited

understanding of the population dynamics and dispersal patterns of P. pyrusana (Brunner

1999). The adoption of low-load sex pheromone lures (Brunner 1999) and food baits that can

catch both male and female moths may improve the predictive ability of traps (Landolt 2000).

Pandemis \eafrollers have a broad host range that includes both cultivated and uncultivated

plant species (Brunner 1983; Vakenti et a/. 2001). Fruit injury has been reported in a range of

tree fruits in Washington including apple, pear (Pyrus communis L.), cherry (Prunus avium

L.), apricot (Prunus armeniaca L.), prune (Prunus domestica L.), and peach/nectarine (Prunus

persica (L.)) (Newcomer and Carlson 1952; Brunner 1983). Reduced insecticide sprays in

cherry after harvest and within blocks of non-bearing apples can allow the establishment of

refugia for P. pyrusana populations within a region (Brunner and Beers 1990). Development

of an effective area-wide management scheme for tortricid leafrollers such as P. pyrusana may

require the application of season-long control tactics in these crops.

Grids of monitoring traps along with intensive larval sampling have been used successfully

to study the area-wide population dynamics of other tortricids among various crop and non-

crop hosts (Knight and Croft 1987; Knight and Hull 1988). These studies have demonstrated

the patchwork pattern of overwintering pest populations and have clarified patterns of seasonal

adult dispersal across crop types. A similar protocol was used in this study. The objectives of

this study, conducted in 1999, were to measure the population density of P. pyrusana

overwintering and developing during the summer within a variety of tree fruit crops within a

contiguous region in the Yakima Valley, Washington, and to compare the usefulness of either

a low-load sex-pheromone-baited or liquid-food-baited trap to monitor populations and predict

larval densities and fruit injury levels in apple and pear orchards.

MATERIALS AND METHODS

This study was conducted in a 3 km’ contiguous area of tree fruit production (Parker

Heights) situated in the Yakima Valley of Washington (46° 29’N, -120° 24’W). Pandemis

pyrusana had been reported to be an important pest for several apple and pear growers in this

area during 1998. We identified 60 orchard blocks comprising 154 ha of apple, pear, cherry,

peach, apricot, peach/nectarine, and prune production (Table 1). Apple orchards were planted

with four cultivars (% of area): ‘Gala’ (46%), ‘Golden Delicious’ (19%), ‘Red Delicious’

(19%), and ‘Fuji’ (18%). All orchards received typical seasonal spray programs during the

season (Olsen 2001).

Blocks were sampled for first generation larvae twice, in late May and early June, and

again in August for second generation larvae. Five shoots from 10 trees within 50 m of each

trap site were inspected on each date. Pole pruners were used to sample shoots randomly from

the upper canopy. Rolled leaves were partially opened to determine if larvae were present.

Infested shoots were placed in small paper bags, and returned to the laboratory. Recovered

larvae were placed on artificial pinto bean diet (Shorey and Hale 1965) and reared under

constant light at 24°C until adult eclosion to determine species and parasitism rate. Parasitoids

were identified by Robert Pfannenstiel (USDA, ARS, Weslaco, TX). Apple and pear blocks

were sampled for fruit injury just prior to harvest. Thirty fruit selected randomly on 20 trees

within 50 m of each trap site were visually examined.

Two types of traps placed in a regular array (100 — 200 m spacing) were used to monitor

adult P. pyrusana. Low-load (10% of the proprietary standard load) sex-pheromone-

impregnated red septa (Trécé Inc., Salinas, CA) were used in delta traps (Pherocon 6, Trécé

Inc., Salinas, CA). Food bait traps consisted of plastic dome traps (Scenturion Inc., Clinton,

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 219

WA) loaded with 150 ml of 1.0% glacial acetic acid and brown food coloring. All traps were

placed in the field the last week.

Table 1

Composition and density of tree fruit orchards within the Parker Heights study site and the

number of sampling sites associated with sex-pheromone and food-bait traps.

Crop No. orchard blocks No. hectares No. trapping sites

Apple 19 66.0 68

Pear 18 36.4 48

Cherry 14 224 28

Peach 4 17.4 20

Non-bearing apple 2 5.4 6

Apricots 3 2 6

Prune l [2 2

Total 61 153.6 178

of May. Each trap type was placed in orchard blocks at a rate of one trap per 2 hectares for a

total of 178 trap sites. Traps were placed about 2-m high in the canopy. Traps were checked

weekly from early June to early August (first flight) and late August to early October (second

flight). Moths caught in bait traps were returned to the laboratory for identification and P.

pyrusana moths were sexed under a dissecting microscope. Sex-pheromone septa were

replaced every 4 weeks. Sticky trap liners used in delta traps and the liquid bait solution in the

dome traps were replaced as necessary.

Data Analysis. All moth count data were transformed to stabilize variances [square root

(x + 0.01)] prior to analysis of variance (Analytical Software 2000). Moth catches in the single

trap of each type placed in the prune orchard were not included in these analyses. Means were

separated with Fisher’s LSD test where significant differences occurred (P < 0.05). Linear

correlation coefficients were computed among the cumulative mean moth counts per trap

during each flight period, larval densities, and percent fruit injury. A chi-square contingency

test was used to compare the proportion of parasitized larvae among crops during each

generation.

RESULTS

Larval Sampling. Overwintering larvae were found in only 10 blocks within the study site:

apple (4), non-bearing apple (2), pear (2), and cherry (2); and were not found in any blocks of

peach/nectarine, apricot, or prune. While non-bearing apples had the highest mean percentage

of infested shoots (Table 2), the percentage of infested shoots ranged up to 5% in pear and 6%

in apple. Parasitism of overwintering larvae by the tachinids, Nemorilla pyste Walker and

Nilea erecta Coquillett, totaled 37% and varied significantly among crops (x? = 11.47, df= 3,

P <0.01): 12.5% in cherry, 37.8% in apple, 50.0% in pear, and 85.7% in non-bearing apple.

Larval population density within the study site was roughly 3-fold larger during the

summer than the overwintering generation, but densities increased nearly 5-fold in apple and

10-fold in both non-bearing apple and cherry blocks (Table 2). Summer generation larvae were

sampled in three blocks of apple, three blocks of non-bearing apple, and two blocks of cherry.

These included four blocks in which an overwintering population was not previously detected.

In addition, in five blocks, overwintering larval populations were detected but no second

generation larvae were found. The highest mean percentage of infested shoots occurred in non-

bearing apple (Table 2). There were no infested shoots found in pear orchards during the

summer. Apple orchards with the highest densities of summer generation larvae were situated

near two areas containing cherry and non-bearing apple. Tachinids parasitized 23% of the

220 J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

Table 2

Correlations among the overwintering and summer larval densities and fruit injury for P.

pyrusana within tree fruit crops in the Parker Heights study site.

% (+ SE) infested shoots

Correlation coefficients*

Overwintering Summer % (+ SE) fruit

Crop larvae (OL) larvae(SL) injury(FI) OL-SL OL-FI SL-FI

Non-bearing apple 0.50+0.27 5.88+2.10 - 0.83** - -

Apple 0.45+0.18 2.07+0.68 0.06+0.02 0.38 0.18 0.837*

Pear 0.25+0.16 0.00+0.00 0.03+0.02 0.00 0.21 0.00

Cherry 0.28+0.14 2.644+1.28 - 0.15 - -

Peach 0.0 0.0 0.0 - - -

Apricot 0.0 0.0 0.0 - - -

Prune 0.0 0.0 0.0 - - -

“Correlation coefficients followed by ** were significant at P < 0.01.

field-collected summer generation larvae and no difference was found among crops (x? = 3.45,

df=2, P=0.18).

Moth Flight. Moth catch in sex-pheromone-baited traps varied among crops during the

first moth flight (F = 3.55; df = 5,82, P < 0.01) and was higher in apple, pear and cherry than

in peach (Table 3). During the first flight, moth catch in non-bearing apple and apricot did not

differ from the other crops. Cumulative moth catch in sex-pheromone traps increased about

400% between generations but did not vary among crops during the second flight (F = 0.76;

di =). 3257 =458):

Moth catch in the food-bait traps was much lower than in sex-pheromone traps during both

flights (Table 3). The male:female sex ratio was 0.60 and 2.26 during the two moth flights,

respectively. Cumulative male catch varied significantly among crops during both the first

(F = 2.65; df= 5, 82; P < 0.05) and second moth flight (F' = 3.00; df= 5, 82; P< 0.05). Male

moth catch was lowest in apricot during both flights. Male moth catch in cherry during the

second flight was significantly higher than in all other crops except apple (Table 3).

Cumulative female catch varied among crops during the first flight, (F = 3.02; df= 5, 82;

P < 0.01) and counts were higher in apple and cherry than in peach and non-bearing apple

(Table 3). Moth catch in food-bait traps increased about 250% between the two flights.

Cumulative total moth catch in the bait traps varied among crops during both flights (first

flight: F = 2.81; df = 5, 82; P < 0.05 and second flight: F = 2.46; df = 5,82; P < 0.05).

Cumulative moth catch in bait traps during the first flight in apple and cherry were

significantly higher than peach, apricot and non-bearing apple (Table 3). Cumulative catch in

bait traps during the second flight in cherry were greater than those in apricot and pear.

Fruit Injury. Fruit injury by P. pyrusana was detected in only one pear and four apple

blocks within the study site and ranged from 0.1 to 0.5%. Surprisingly, no overwintering or

summer generation larvae were detected in samples collected from three of these five blocks.

All orchards with fruit injury were adjacent to or near blocks of cherry and non-bearing apple.

Correlations Among Population Measures. The overwintering density of larvae in both

apple and pear orchards was not correlated with levels of fruit injury (P = 0.79 and P = 0.77,

respectively) (Table 2). However, summer larval densities in apple were well correlated with

fruit injury (P < 0.001). The percentage of shoots infested by overwintering larvae was not

significantly correlated with percentage of shoots infested with summer larvae in apple

(P = 0.27) or cherry (P = 0.90). However, larval densities in non-bearing apple blocks were

significantly correlated (P < 0.01) (Table 2).

The correlations of cumulative moth catch per trap during each flight period with other

221

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measures of population density varied for each trap type in apple and pear (Table 4).

Cumulative moth catch from both trap types during the first flight was significantly correlated

with fruit injury in apples and pears but this relationship was weak (r’s < 0.50; Table 4).

Restricting the cumulative moth counts to the first 3 or 4 weeks of the season did not improve

these correlations (r’s < 0.45). In comparison, moth catches during the second flight were not

correlated with fruit injury (Table 4). Cumulative moth catch per trap during the first and

second flight were correlated for each type of trap. However, moth catch in sex-pheromone-

baited traps during both flight periods was not correlated with either overwintering or summer

larval densities. In comparison moth catch in the food-bait traps during the first flight period

was correlated with overwintering larval density and summer larval density; and moth catch

during the second flight was correlated with summer larval density (Table 4). Correlations of

cumulative moths and cumulative female moths for each flight with larval densities and fruit

injury were similar (Table 4).

Table 4

Correlations of cumulative moth catch during the 1“ and 2™ moth flight of P. pyrusana in traps

baited with either low-load sex-pheromone lures or a liquid-food bait with selected population

measures across all apple and pear blocks within the Parker Heights study site.

Trap type Flight period Population measure Correlation coefficient*

Low-load 1* flight Overwintering larval density 0.04

Sex-pheromone Summer larval density 0.25

2™ moth flight 0.41**

% fruit injury 0.45**

Overwintering larval density 0.02

. Summer larval density 0.21

% fruit injury 0.18

Liquid-food bait” Overwintering larval density 0.40* (0.44**)

Summer larval density 0.607* (0.611 **)

2™ moth flight 0.32* (0.36**)

% fruit injury 04677 (037"*)

2" flight Overwintering larval density Olay Orth)

Summer larval density 0.40** (0.35**)

% fruit injury 0.07 (0.07)

“Correlation coefficients followed by * were significant at P < 0.05; coefficients followed by

** were significant at P < 0.01.

> Correlation coefficients in brackets are for cumulative female moth catch only.

DISCUSSION

Establishing action thresholds based on the capture of male moths in sex-pheromone-baited

traps has been difficult for many tortricid pests that occur in high densities within orchards

(Madsen and Peters 1976; Minks eft al. 1995; Walker and Welter 1999). Success has been

achieved by reducing the effects of trap saturation by either using only early-season catches

(Knight and Hull 1989) or by reducing the attractiveness of the lure (Faccioli et al. 1993).

However, these approaches have not always improved the performance of sex-pheromone

traps. A significant correlation of peak moth catch and larval density could not be established

for the apple pest, Argyrotaenia citrana (Fernald) with traps baited with lures across a 1,000-

fold range in their sex-pheromone load (Walker and Welter 1999).

The utility of sex-pheromone traps to accurately predict the population density of P.

pyrusana may be limited. Walker and Welter (2001) found a significant but moderate

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 293

relationship (7 = 0.59, P = 0.04) between peak weekly moth catch of P. pyrusana during the

first moth flight and summer larval density in California apple orchards. While Brunner (1999)

suggested that the use of low-load (5%) sex-pheromone lures improved the accuracy of traps in

predicting larval populations and fruit injury, these data have not been published. Results

reported herein found that low-load (10%) sex-pheromone lures were, at best, weak predictors

of local population densities estimated by either larval sampling or levels of fruit injury.

Many factors affect the performance of lure-baited traps and the correlation of moth catch

with local larval population density. Moth capture within traps is influenced by operational

factors including the lure’s emission characteristics and the attractant’s chemical stability, and

the size, geometry, placement, and maintenance of the trap (McNeil 1991). Saturation of the

trap’s catch surface with moths and non-target species can also reduce the effectiveness of

traps to reflect relatively high population densities (Brown 1984). An accurate estimate of low-

density leafroller larval populations is difficult to achieve without extensive host sampling.

Furthermore, P. pyrusana larvae were typically found feeding on shoot terminals in the upper

canopy of trees, and the variability in tree height among orchards and crops may have created a

sampling bias in the estimate of larval density. In general, a greater number of larvae were

detected in apple orchards with smaller canopies - non-bearing apple and younger blocks of

Fuji. The relationship between moth catch and fruit injury is further impacted by a large

number of horticultural and biological factors including cultivar and crop load, tree size and

pruning, and spray practices. Orchards within our study site varied widely for many of these

factors.

Moth catch of P. pyrusana in both types of traps during first flight was a better predictor of

larval populations and fruit injury than moth catch during second flight. This result is similar to

data previously reported for tufted apple bud moth Platynota idaeusalis (Walker) (Knight and

Hull 1988) and A. citrana (Knight and Croft 1987). Both of these studies used grids of sex

pheromone-baited traps within a diverse agricultural setting. Both leafroller pests overwintered

primarily within managed agricultural sites and early-season male moth catches reflected this

local distribution. However, the adults of both species are highly mobile and male moth

catches of the summer generation flight were more homogeneous within the region. Similarly,

counts of male P. pyrusana during second flight were uniformly high among 56 of the 60

orchards in Parker Heights.

While, moth catches during first flight in food bait traps were more closely associated with

spring and summer larval densities than moth catches in sex-pheromone-baited traps, both trap

types were similar in predicting fruit injury. Interestingly, the counts of female moths in the

food bait traps did not improve the prediction of local population density. Liquid-food-bait

traps had a greater number of problems associated with their use than the sex-pheromone traps.

First, the acetic acid mixture evaporated rapidly during the warmer weather in August and

required frequent servicing. The food-bait traps were not selective and caught a large number

of non-target moths (Lepidoptera: Noctuidae) that saturated the traps. Identification and sexing

of P. pyrusana individuals from a large decomposing mixture of insects was time consuming.

The decomposition and fermentation of the mixture was also highly attractive to muscid flies

and may have released volatile chemicals that may have reduced the attractiveness of the bait

to P. pyrusana. Development of dry food baits placed in either sticky or insecticide-treated

traps will likely reduce these problems (Landolt and Alfaro 2001).

Establishing effective management of a polyphagous pest such as P. pyrusana requires a

concerted area-wide program across all potential hosts. Interestingly, populations of P.

pyrusana in Parker Heights were not detected in the eight commercial orchards of

peach/nectarine, apricot, and prune despite these crops’ apparent host suitability (Brunner

1983). Its absence in these crops may have been due to the use of broad-spectrum insecticides

for other key pests in these crops. For example, early-season and summer sprays for green

224 J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

peach aphid, Myzus persicae (Sulzer), peach twig borer, Anarsia lineatella (Zeller), oriental

fruit moth, Grapholitha molesta (Busck), and western flower thrips, Frankliniella occidentalis

(Pergande), are widely used in these stone fruits (Olsen 2001). In contrast, P. pyrusana larvae

were found in over a third of the cherry orchards. Cherry is sprayed early in the season for

phytophagous mites and the black cherry aphid, Myzus cerasi (Fabricus), and receives a series

of cover sprays through June for the western cherry fruit fly, Rhagoletis indifferens (Curran).

However, Washington growers typically do not apply any insecticides to cherry after June. The

role of cherry orchards in serving as refugia for leafrollers was also reported for P. idaeusalis

in a typical mixed-fruit production area in Pennsylvania (Knight and Huli 1988).

Leafroller management in apple and pear tends to ignore the potential role of these extra-

orchard habitats and this allows populations of P. pyrusana to remain established at high levels

within a given region. Even leafroller populations in non-bearing apple blocks are generally

not treated. Instead, leafrollers are managed by individual growers within-season in their

respective orchards with one or more well-timed applications of efficacious sprays.

Implementation of an effective leafroller management strategy, however, is often hampered by

poor spring weather, poor spray timing, the survival of larvae feeding within protected leaf

shelters, and insecticide resistance.

Conversely, the idea of growers working together to implement an effective area-wide pest

management program that suppresses a pest population across all host habitats would seem to

be more effective and has recently been demonstrated for codling moth, Cydia pomonella L.

(Calkins 1998). Similarly, the obliquebanded leafroller, Choristoneura rosaceana Harris was

effectively managed by 13 growers using sex-pheromone-based mating disruption and B.

thuringiensis sprays (Knight et a/. 2001). The effectiveness of the area-wide approach requires

that the pest population density be reduced to low levels through an integration of selective

tactics. The success of the codling moth project was predicated on the clean-up of all problem

sites that harbored high pest populations (Knight 1999). Similarly, effective area-wide

management of P. pyrusana will require that populations in the surrounding cherry and non-

bearing apple blocks be managed successfully.

ACKNOWLEDGEMENTS

I would like to thank Mike Marsello (USDA, ARS, Wapato, Washington) for his technical

assistance in collecting and summarizing the data. I would also like to acknowledge the

cooperation of the 14 Parker Heights growers, and the financial support provided by the

Washington State Tree Fruit Research Commission. This paper was significantly improved by

the many constructive comments made by Art Agnello (Department Entomology, New York

State Agricultural Experiment. Station, Geneva, NY), Jim Hansen (USDA, ARS, Wapato,

WA), Peter Shearer (Department Entomology, Rutgers Agricultural Research Center,

Bridgeton, NJ), and two anonymous reviewers.

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J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 DOF

Fleas (Siphonaptera) from sciurid and murid rodents

on the eastern slope of the Cascade Range,

Kittitas County, Washington

JAMES R. KUCERA!

ASSOCIATED REGIONAL AND UNIVERSITY PATHOLOGISTS, INC.,

SALT LAKE CITY, UT, UNITED STATES

GLENN E. HAAS

557 CALIFORNIA AVE., PMB #7, BOULDER CITY, NV, UNITED STATES 89005

MICHAEL K. MACDONALD

WASHINGTON DEPARTMENT OF TRANSPORTATION, 15700 DAYTON AVE.

NORTH, SEATTLE, WA, UNITED STATES 98133-9710

ABSTRACT

Eight species of rodent fleas [Ctenophthalmidae: Megarthroglossus procus Jordan &

Rothschild; Leptopsyllidae: Peromyscopsylla selenis (Rothschild); Ceratophyllidae:

Ceratophyllus ciliatus protinus Jordan, Eumolpianus eumolpi eumolpi (Rothschild),

Malaraeus telchinus (Rothschild), Opisodasys vesperalis (Jordan), Orchopeas agilis

(Rothschild) and Oropsylla idahoensis (Baker)| were collected in 1993 and 1995 from

seven species of rodents [Neotamias amoenus (J.A. Allen), N. townsendii (Bachman),

Spermophilus saturatus (Rhoads), Glaucomys sabrinus (Shaw), Neotoma cinerea (Ord),

Clethrionomys gapperi (Vigors) and Microtus longicaudus (Merriam)] live and snap

trapped. There were four new county records for M. procus, P. selenis, O. vesperalis

and O. idahoensis, and there were five new host records for the state with M. procus and

C. ciliatus protinus ex G. sabrinus, P. selenis and M. telchinus ex C. gapperi and P.

selenis ex M. longicaudus. Distribution patterns and host preferences in the Pacific

Northwest are discussed.

Key words: fleas, Siphonaptera, rodents, Washington State

INTRODUCTION

The flea fauna of Washington is less well known than its neighbors British Columbia

and Oregon, with only 80 species of fleas recorded while British Columbia has 98 species

and Oregon has 110 (Holland 1985; Lewis et a/. 1988). The number of publications

concerning fleas is smaller for Washington than for Oregon (Lewis ef al. 1988). The

number of locality/county symbols on distribution maps is fewest for Washington

(Haddow et al. 1983; Holland 1985; Lewis ef a/. 1988). Washington ranks last, at least in

part, because of its comparatively small land area (172,266 km’). Oregon is 76,858 km?

larger, and British Columbia (948,600 km’) is 2.25 times larger than Oregon and

Washington combined. Danks (1995) considered range of habitats as well as comparative

land areas and found the same lower ranking for Washington in his tabulations from

' Address correspondence to: 5930 S. Sultan Circle, Murray, UT 84107-6930

D8 J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

published species records in the order Dictyoptera and according to selected families and

genera in the orders Hemiptera, Coleoptera, Diptera, Lepidoptera and Hymenoptera.

As Lewis et al. (1988) concluded from their review of literature, the flea faunas of the

states contiguous with Oregon “... are in much need of additional study.” In the present

study, we contribute new data for eight rodent fleas in Kittitas County, central

Washington, with five new flea-host associations for the state.

MATERIALS AND METHODS

All flea specimens were received from the University of Alaska (Fairbanks) Museum

mammalogy section, preserved in alcohol. All were collected by one of us (MKM) in

Wenatchee National Forest, Kittitas County, Washington during 1993 and 1995. Hosts

were collected with snap traps (woodrats) and Havahart® and Sherman® live traps and

snap traps (flying squirrels, chipmunks and voles). Host specimen number (AF number)

for the University of Alaska mammal collection is designated in brackets. Fleas were

permanently mounted on microscope slides by standard techniques (Lewis et al. 1988). All

flea specimens were deposited in the US National Museum insect collection. The western

chipmunk genus Neotamias, as proposed by Jameson (1999), is adopted here.

SPECIES ACCOUNTS

Ctenophthalmidae

Megarthroglossus procus Jordan & Rothschild, 1915

South Fork Taneum Creek, 1.6 km west of South Fork Meadows, 47°5’49.08”N,

121°0’37.68"W (1150m), 23 Aug 1995 [AF 13631], 12 ex Glaucomys sabrinus (Shaw)

(northern flying squirrel) .

The specimen is identified as M. procus, although “... a definite identification is

possible only if male specimens are available” (Tipton et a/. 1979). Characters such as the

spermathecal form and the lack of a noticeable sinus on the posterior margin of sternum

VII, plus the known host associations, are consistent with this identification. The

morphologically similar M. jamesoni Smit occurs in California and Nevada (Lewis et al.

1988). Megarthroglossus procus is known from Skagit, Whatcom and Yakima Counties

(Tipton et al. 1979). Our specimen adds Kittitas County and a new host for the state.

Megarthroglossus procus is widely distributed in western North America, from the

Pacific Northwest and as far east as western Nebraska. There is an apparent void east of

the Cascades in Washington and Oregon (Mendez 1956; Tipton ef al. 1979).

Megarthroglossus divisus (Baker) is known to replace M. procus in nests of G. sabrinus in

northeastern Oregon (Wilson and Bull 1977; Whitaker ef a/. 1983).

Records from British Columbia, Oregon and California convinced Holland (1949b,

1985) that the true host of M. procus is Tamiasciurus douglasii (Bachman) (Douglas’

squirrel). From the wide variety of recorded hosts, Lewis et al. (1988) were inclined to

name 7. hudsonicus (Erxleben) (red squirrel) as a preferred host. More collecting from G.

sabrinus and especially its nests will probably establish it as another preferred host of M.

procus.

Leptopsyllidae

Peromyscopsylla selenis (Rothschild, 1906)

South Cle Elum Ridge, along US Forest Service road 3350, 47°8°20.10’N,

120°58’12.24"W (1172m), 21 Sep 1995 [AF 5504], 19 ex Microtus longicaudus

(Merriam) (long-tailed vole). South Fork Taneum Creek, 1.6 km west of South Fork

Meadows, 47°5’49.08”N, 121°0°37.68"W (1150m), 17 Aug 1995 [AF 5499], 192 ex

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 229

Clethrionomys gapperi (Vigors) (southern red-backed vole). Same locality but 25 Aug

1995 [AF 13640], 19 ex C. gapperi.

Peromyscopsylla selenis is a common flea found on arvicoline rodents and accidentally

on other hosts in many areas of the western United States and Canada (Johnson and Traub

1954) but sparsely recorded in Washington State. The earliest collections in the state were

in 1935 from Microtus richardsoni (DeKay) (water vole) in Skamania County and in 1938

from M. townsendii (Bachman) (Townsend’s vole) in Skagit County (Hopkins and

Rothschild 1971). Hubbard (1943, 1947) recorded a 1943 collection from M. richardsoni

in Klickitat County. Johnson and Traub (1954) added a record from a Microtus sp. in

Spokane County. The distribution map of Lewis et al. (1988) has marks for three of these

counties plus Whitman County. Our two hosts, C. gapperi and M. longicaudus, are new for

the state in a county not previously known for P. selenis.

Ceratophyllidae

Ceratophyllus ciliatus protinus Jordan, 1929

South Fork Taneum Creek, 1.6 km west of South Fork Meadows, 47°5’49.08”N,

121°0’37.68”W (1150m), 17 Aug 1995 [AF 13575], 13, 329 ex Neotamias townsendii

(Bachman) (Townsend’s chipmunk). Same date and locality [AF13574], 2¢¢ ex N.

townsendii. Same date and locality [AF 13573], 14, 399 ex N. townsendii. Same date and

locality [AF 13626], 14, 19 ex N. townsendii. Same locality but 18 Aug 1995 [AF 13599],

1¢ ex G. sabrinus. Same date and locality as previous [AF 13601], 19 ex N. townsendii.

South Fork Taneum Creek, 0.8 km west of South Fork Meadows, 47°5’51.96’N,

121°0’6.00”W (1152m), 16 Aug 1995 [AF 5474], 14, 229 ex N. townsendii. Same date

and locality as previous [AF 5481], 14 ex N. townsendii. Same locality but 24 Aug 1995

[AF 14912], 19 ex N. townsendii.

This taxon is well known from tree squirrels, chipmunks and some associated small

mammals in western Washington, Oregon and British Columbia (Hubbard 1947; Johnson

1961; Haddow et al. 1983; Holland 1985). The earliest record for Washington is from

Carson, Skamania County, 1939 (Hubbard 1940), and since then King, Kittitas, Pierce and

Yakima Counties were added (Lewis et al. 1988). Lewis and Maser (1981) and Lewis ef

al. (1988) reported large numbers of C. ciliatus protinus on Townsend’s chipmunk in

Oregon. All but one of our 19 specimens came from this host. The exception came from G.

sabrinus, a host record that is new for the state.

Eumolpianus eumolpi eumolpi (Rothschild, 1905)

South Cle Elum Ridge, 1.6 km southwest of Peoh Point along USFS road 3350,

47°8°31.74°N, 120°57°42.54”"W (1411m), 21 Sep 1995 [AF 14925], 229 ex Neotamias

amoenus (J.A. Allen) (yellow-pine chipmunk).

The genus Eumolpianus was erected by Smit (1983) to include the distinctive

chipmunk fleas of the “ewmolpi group,” genus Monopsyllus, of Traub and Johnson (1952)

(see also Johnson 1961). The range of the genus Eumolpianus is roughly that of its

preferred hosts, chipmunks of the genus Neotamias in Canada, the western and northern

United States and Mexico (Johnson 1961; Haddow et a/. 1983; Holland 1985). Of the two

nominal subspecies, E. e. eumolpi occurs in the Pacific Northwest and is one of the most

common fleas of Washington east of the Cascade crest. Since the early collection of 1920

in Adams County (Jellison and Senger 1976), it has been reported from the following 11

counties: Asotin, Chelan, Ferry, Grant, Kittitas, Klickitat, Lincoln, Spokane, Stevens,

Whitman and Yakima (Hubbard 1943; Miller and Drake 1954; Johnson 1961; Jellison and

Senger 1976; Lewis et al. 1988). Nonetheless, distribution maps continue to show no

records in the moister areas west of the Cascades (Johnson 1961; Haddow et al. 1983;

Lewis et al. 1988).

230 J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

Malaraeus telchinus (Rothschild, 1905)

South Fork Taneum Creek, 1.6 km west of South Fork Meadows, 47°5’49.08’N,

121°0°37.68"W (1150m), 24 Aug 1995 [AF 13616], 1¢ ex C. gapperi. South Cle Elum

Ridge, 1.2 km east of USFS road 214 along USFS road 3350, 47°8’20.10’N,

120°58712.24”W (1172m), 21 Sep 1995 [AF 5504], 19 ex M. longicaudus.

This is the more commonly collected species of Malaraeus in the Pacific Northwest;

M. sinomus is unknown in Washington. Malaraeus telchinus is recorded from Clark,

Grant, Klickitat, Lincoln, Skamania, Whitman and Yakima Counties (Hubbard 1947;

Jellison and Senger 1976; Lewis et al. 1988). Malaraeus telchinus is found on a variety of

mice and voles in mesic habitats; however, there are no published records from

Clethrionomys in the State of Washington.

Opisodasys vesperalis (Jordan, 1929)

All ex G. sabrinus. South Fork Taneum Creek, 0.8 km west of South Fork Meadows,

47°5°51.96”N, 121°0’6.00”W (1152m), 10 Aug 1995 [AF 5466], 12. Same locality but 18

Aug 1995 [AF 13588], 19. Same locality but 23 Aug 1995 [AF 35114], 229. South Fork

Taneum Creek, 1.6 km west of South Fork Meadows, 47°5’49.08”N, 121°0’37.68”W

(1150m), 23 Aug 1995 [AF 13622], 229. Same date and locality [AF 13631], 444, 299.

Glaucomys sabrinus is the preferred host of this flea in the Pacific Northwest.

Opisodasys vesperalis is also found on G. volans (L.) (southern flying squirrel) in other

areas of North America. Sparse collection records in British Columbia (Holland 1985) and

Washington (Lewis ef al.1988) probably reflect the need to examine more G. sabrinus and

their nests for fleas. It was previously unknown from Kittitas County, being known in the

state only from Clallam, Cowlitz and Lincoln Counties (Jellison and Senger 1976; Lewis

et al. 1988). The presence of Opisodasys pseudarctomys (Baker), another true Glaucomys

flea that is unknown from the state (Haddow ef al. 1983), may also be established by

further collections from G. sabrinus or especially their nests.

Orchopeas agilis (Rothschild, 1905)

All ex Neotoma cinerea (Ord) (bushy-tailed woodrat). S Fork Taneum Creek, USFS

road 135, in abandoned cabin, 47°6711”N, 120°57’1”W (900m), 25 Sep 1993 [AF 5405],

14,629. 12 Oct 1993 [AF 5427], 14. [AF 5428], 23, 499. [AF 5429], 14, 19.

This parasite of Neotoma spp. was originally described as Ceratophyllus agilis by

Rothschild (1905) from a type series collected from N. cinerea in Banff, Alberta and other

mammals from other localities in Alberta and British Columbia (Holland 1985). Jordan

(1929) reduced this taxon to a subspecies of C. sexdentatus (Baker). Jordan (1933)

published the genus name Orchopeas to replace the preoccupied genus name Bakerella

Wagner. In his review of the fleas of British Columbia, Wagner (1936) listed this flea as

Orchopeas sexdentatus agilis.

Recently, Lewis (1998, 2000) reviewed the genus Orchopeas Jordan and elevated to

species each of the six taxa that had been subspecies of O. sexdentatus. Lewis (2000) also

noted that O. agilis is the member of the O. sexdentatus group with the widest distribution,

ranging from the Yukon Territory through British Columbia and western Alberta, eastern

Washington, Oregon and California, the Basin and Mountain states south into the Mojave

Desert in Nevada, Utah and Arizona, and into the Rio Grande watershed of New Mexico

(Haddow et al. 1983; Holland 1985; Lewis et al. 1988).

Orchopeas agilis is known from several Neotoma spp., but N. cinerea is the only

species in Washington and Canada (Cowan and Guiguet 1965; Ingles 1965; Banfield

1974). Locality records of O. agilis are especially numerous in southern British Columbia

(Holland 1985). In Washington there are records of collections of either O. sexdentatus or

O. s. agilis (both = O. agilis) in eight counties: Benton, Douglas, Franklin, Grant,

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 23]

Klickitat, Lincoln, Spokane and Yakima (Hubbard 1940, 1943, 1947; Bacon 1953;

O’Farrell 1975; Jellison and Senger 1976; Lewis ef a/. 1988). Our new records of O. agilis

confirmed Kittitas County.

In a survey of wild animal diseases in five counties in the Columbia Basin, Miller and

Drake (1954) found O. agilis only on Peromyscus maniculatus (Wagner) (deer mouse).

Only a small, unspecified number of specimens were collected in one or more unnamed

counties. The counties surveyed were Adams, Franklin, Grant, Kittitas and Lincoln.

The nine contiguous counties with records occupy much of the state east of the

Cascades. Orchopeas cascadensis Jordan, the other member of the O. sexdentatus group in

Washington, is known only from west of the Cascades in Clark County (Hubbard 1947;

Lewis et al. 1988; Lewis 2000).

Oropsylla idahoensis (Baker, 1904)

South Fork Taneum Creek, 1.6 km west of South Fork Meadows, 47°5’49.08”N,

121°0°37.68”W (1150m), 16 Aug 1995 [AF 5484], 14, 299 ex Spermophilus saturatus

(Rhoads) (Cascade golden-mantled ground squirrel).

This is the only flea known to parasitize S. saturatus. Early records from this host for

Washington are in three counties: Skamania (1935), Yakima (1938) and Klickitat (1939,

1943) (Hubbard 1940, 1943, 1947). Hubbard (1947) also mentioned locality records in

Skamania and Yakima Counties without giving collection dates and numbers of

specimens. Holland (1949a) recorded early collections from S. saturatus in British

Columbia: Princeton (1939) and Manning Provincial Park (1945). Holland (1985) gave

two additional records for Manning Provincial Park (1953 and 1955). Our new record is in

a county that lies on the same longitude (121°) that runs through contiguous Yakima

County to the south and Manning Provincial Park to the north.

The wider-ranging Pacific Northwest Spermophilus spp. are hosts to other species of

fleas in addition to O. idahoensis (see Wagner 1936; Jellison 1945; Holland 1985; Lewis et

al. 1988). The apparent absence of these fleas on S. saturatus could be a reflection of less

collecting effort, as this ground squirrel has been classified by some authors as a

subspecies of S. /ateralis (Say), a common, wide-ranging western North American ground

squirrel. Spermophilus saturatus is nowhere sympatric with S. /ateralis, being restricted to

a small range on the eastern slopes of the Cascades in Washington and British Columbia

(Cowan and Guiguet 1965; Ingles 1965; Banfield 1974; Tomich 1982; Trombulak 1988).

CONCLUSION

Although no fleas were added to the Washington list, the five new host records suggest

that future surveys should include rodents known to host fleas on the lists for British

Columbia and Oregon. With only one species of flea known for the rodent Spermophilus

saturatus, a thorough survey in this mammal’s small Washington to British Columbia

range is desirable. Mammals other than rodents, such as opossums, shrews, moles, bats,

pikas and carnivores deserve more attention as do domestic mammals and birds, wild and

domestic. Above all, nests of mammals and birds need to be examined for adult fleas and

their poorly known larval stages.

ACKNOWLEDGEMENTS

Professor J. Cook of the University of Idaho (Pocatello) bestowed this collection to us

from the University of Alaska. We are also grateful to Amy Runck (University of Alaska,

Fairbanks) for providing and clarifying data. The comments of two anonymous reviewers

improved the manuscript.

232 J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

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J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 235

Use of Japanese-beetle traps to monitor flight of the Pacific

coast wireworm, Limonius canus (Coleoptera: Elateridae),

and effects of trap height and color

DAVID R. HORTON and PETER J. LANDOLT

USDA-ARS, 5230 KONNOWAC PASS Rad.,

WAPATO, WA, UNITED STATES 98951

ABSTRACT

Japanese-beetle traps were used to monitor flight of the Pacific coast wireworm,

Limonius canus LeConte, in an agricultural field in northern Oregon. Overwintered

' beetles first appeared in traps in mid-March 2000 and 2001, and were collected until

mid- to late-May both years. Most (93%) of the females collected at the beginning of the

flight period had been inseminated, which may indicate that mating takes place very

soon after beetles emerge from the soil. Sex ratios favored males at the beginning of the

flight period and favored females at the end of the flight period. Lower temperatures in

April 2001 compared to those in 2000 may have caused a delay in timing of the peak

catch (relative to timing in 2000) by almost 3 weeks. A count of over eight beetles per

trap per 7-day sampling interval was obtained during the week of peak catch in April

2000. Traps were hung at three heights:1.5, 0.9, and 0.3 m above ground. Catch

decreased with increasing trap height. Traps that had been painted yellow collected

more beetles than traps painted white, which in turn collected more beetles than traps

painted red, green, dark blue, or black. Two other elaterids, Crenicera pruinina (Horn)

and Cardiophorus montanus Bland, were also trapped during the study.

Key words: Limonius canus, click beetles, monitoring, flight, trap

INTRODUCTION

The Pacific coast wireworm, Limonius canus LeConte (Coleoptera: Elateridae), inhabits

irrigated soils of western North America, where it is a pest of grain and vegetable crops

(Gibson 1939; Lane and Stone 1960). Like other wireworm species (Stone 1941; Lane and

Stone 1960), L. canus requires several years to complete its life cycle (Landis and Onsager

1966). The insect overwinters in the soil primarily in the larval stage as a mix of ages. Final-

instar larvae pupate in summer, and the pupae eclose into the adult stage in fall and winter.

Adults remain in the soil during the winter, emerging the following spring apparently in

response to increasing soil temperatures. Life span of the overwintered (post-emergence) adult

appears to be no longer than 2 months (Toba 1986).

There is a critical need for research on this and other wireworm pests that could lead to

advances in managing these insects (Jansson and Seal 1994). Research into new ways of

sampling populations is needed, to allow studies of population biology and to assist growers in

making management decisions (Jansson and Seal 1994). Much of the previous research done

on sampling these insects has concentrated on the larval stage (e.g., Onsager 1975; Toba and

Turner 1983; Williams ef a/. 1992). Considerably less study has focused on the adult click

beetle, despite the fact that the adult stage is responsible for spreading wireworm infestations

(Boiteau et al. 2000).

This study tests whether traps that were developed to monitor Japanese beetle, Popillia

Japonica Newman (Coleoptera: Scarabaeidae), might be used to sample adult L. canus.

236 J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

Japanese-beetle traps have several advantages over other traps used to sample click beetles

(including sticky traps, window interception traps, and water pan traps), in that they are simple

to set out in the field, are easily monitored, and require no adhesive materials such as

tanglefoot to trap the insect. We used these traps to document the flight period of male and

female L. canus in an agricultural field located in north-central Oregon, and determined if

seasonal trends in trap catch were associated with air temperature. Second, traps were placed

at different heights to learn if catch depended upon height of the traps above ground, as shown

for other Elateridae (Boiteau et a/. 2000). Third, we compared traps of different colors to

monitor their effectiveness. Finally, we dissected female beetles trapped on one date at the

beginning of the female flight period to determine if they were mated. Other wireworm species

mate very soon after emergence from the soil in spring (Stone 1941; Zacharuk 1962), but it is

not known whether L. canus exhibits similar behavior.

MATERIALS AND METHODS

All studies were conducted between March and May, 2000 and 2001 at the Agricultural

Research and Extension Center, Oregon State University, Hermiston, Oregon. Yellow,

four-vaned Japanese-beetle traps (Trécé, Salinas, CA) were hung on metal poles in a fallow

field adjacent to a circle of irrigated wheat. The same location was used both years of the

study. Ten poles were set out in a line at spacings of approximately 20 m. Three traps at

different heights were hung on each pole: 1.5 m, 0.9 m, and 0.3 m above ground. Vegetation in

the surrounding field and immediately beneath the traps was cut by a mower or by hand to

prevent plants from obscuring the lowest traps. Clear glass jars were used as the collecting

reservoirs beneath the traps. Jars were emptied every 7 d between March and May. Click

beetles were counted, identified, and categorized to sex. Air temperature data were obtained

from a weather station located at the study site and maintained by Experiment Station

personnel.

Trap catch data were compared among trap heights using analysis of variance, with each

pole being considered a block. To determine if trap catch increased or decreased as a linear or

curvilinear function of height, linear and quadratic contrasts were extracted in each analysis.

A second study was done to determine the effects of trap color on catch. The visible

surfaces of yellow Japanese-beetle traps were painted with one of six colors (Krylon High

Gloss paints, Sherwin-Williams, Cleveland, OH): Sun Yellow Gloss, Glossy White, Emerald

Green, Banner Red, Gloss Black, and True Blue Gloss. Traps were hung at 0.3 m above

ground on fence lines at the Hermiston Experiment Station during the same time interval as the

height tests. In 2000, traps were placed at the northern edge of the station adjacent to plots of

wheat and potatoes. Catch of L. canus was small here, so in 2001 the traps were moved 200-

500 m to the south, where they were hung 0.3 m above ground on fence lines adjacent to an

irrigated circle of wheat. Six traps of each color were set out in a randomized complete-block

design with six replicates. Adjacent traps within a block were approximately 5 m apart.

Adjacent blocks were separated by at least 20 m. Traps were emptied weekly, and click beetles

were counted, identified, and sexed. Data were analyzed using analysis of variance followed

by an LSD-test to separate treatments (trap colors).

To determine whether the first female L. canus collected in traps in 2001 had been

inseminated, we dissected beetles that had been collected on 1 May during the height test. On

24 April, 2001, water was added to each collecting jar on each trap to prevent beetles from

mating while in the jar. Water was used rather than a preservative to avoid the possibility that

odors from the preservative might affect trap catch. The collecting jars received water from

rainfall at irregular intervals over the course of the studies, so the presence of water in the jars

was not something unusual. Jars were collected 1 week later and samples were taken to the

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 237

laboratory. Female L. canus were dissected in alcohol to remove the internal reproductive

organs, using the drawings of Becker (1958) and Zacharuk (1958) as visual reference. The

spermatophoral receptacle (Zacharuk 1958) was then examined under a dissecting microscope

to determine whether a spermatophore was present. If no spermatophore was found, the

internal organs were teased apart with dissecting needles, crushed on the microscope slide, and

examined for sperm using a compound microscope at 100-400x (in other Elateridae absence of

a spermatophore does not indicate lack of mating, as the spermatophore is slowly broken down

in the female; Zacharuk 1958). Decomposed beetles were not dissected.

Voucher specimens have been deposited with the M.T. James Entomology Museum,

Washington State University, Pullman.

RESULTS

The largest weekly catch of L. canus in the height test was about three-fold higher in 2000

than 2001 (Fig. 1). Beetles began showing up in traps in mid- to late-March apparently in

response to warming temperatures (Fig. 1). We captured beetles well into May both years. The

peak in captures occurred almost 3 weeks later in 2001 (1 May sample) than in 2000 (12 April

sample). Moreover, the peak catch contained a much larger percentage of females in 2001 than

2000 (see below). Lower temperatures in late March and early April 2001 relative to

conditions in 2000 may have caused reduced levels of flight activity in 2001 compared to

levels over the same time interval in 2000 (Fig. 1).

300

250

200

—-—- High temp.

—@e— Click beetles

Beetles per 30 traps

(snisja9) aunyeradwwa} ybiy Ajleq

10 20 30 10 20 30 10 20 30

March April May

Figure 1. Numbers of L. canus collected in 30 traps (10 poles x 3 traps per pole) per week

(solid lines) and daily high air temperatures (dashed lines); Hermiston Agricultural Research

and Extension Center, Hermiston, Oregon. Horizontal dotted line at 20° C included in each

panel to provide perspective.

238 J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

Males predominated in the March and April samples, whereas females were more

abundant than males in samples obtained in May (Figs. 2-3). Samples obtained during peak

flight had a much higher proportion of males in 2000 (88.6% male [217/245 beetles]; 12 April

sample) than in 2001 (50.6% male [43/85 beetles]; 1 May sample). Total numbers of male

beetles collected over the sampling period was larger in 2000 than in 2001 (x = 50.4 beetles

per three traps in 2000 vs 15.6 beetles per three traps in 2001; F = 131.5, df= 1,18, P< 0.001;

numbers pooled for the three traps on the same pole and summed across dates). Totals for

females were similar between years (X = 14.7 beetles per three traps in 2000 vs 10.1 beetles

per three traps in 2001; F = 3.2, df= 1,18, P = 0.09). In both years, numbers decreased with

increasing height of the trap (Figs. 2-3).

Ga Males

E-ZZZA Females

Weekly total catch per 10 traps

March 29 April 12 April 26 May 10

Figure 2. Weekly total catch of L. canus per 10 traps at each of three heights; 2000 data.

Mean number of beetles per trap (sexes combined; summed over 7-week sampling period):

28.9 (0.3 meters), 24.9 (0.9 meters), and 11.3 (1.5 meters); F = 29.2; df = 2,18; P < 0.001

(contrasts: linear effects of height, P < 0.001; quadratic effects of height, P = 0.03).

Gu Males

Females

Weekly total catch per 10 traps

30 Z

Mar. 27 Apr.10 Apr.24 May8 May 22

Figure 3. Weekly total catch of L. canus per 10 traps at each of three heights; 2001 data.

Mean number of beetles per trap (sexes combined; summed over 10-week sampling

period):13.0 (0.3 meters), 8.7 (0.9 meters), and 4.0 (1.5 meters); F = 16.6; df = 2,18; P<

0.001 (contrasts: linear effects of height, P < 0.001; quadratic effects of height, P = 0.88).

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 239

Click beetles other than L. canus collected in the traps included Ctenicera pruinina

(Horn), Cardiophorus montanus Bland, and unidentified Dalopius and Melanotus (Table 1).

Of these, C. pruinina (Great Basin Wireworm) is a known pest of vegetable and grain crops in

the Pacific northwest. Most of the beetles were collected in the lower and middle traps, as with

L. canus. Sex ratios were male-biased (Table 1).

Few L. canus were collected in the 7-week test of trap color done in 2000 (Table 2). The

traps were moved to a different area of the station in 2001 with better results. In 2001 (10

weeks), traps that had been painted yellow caught significantly more L. canus than white traps

(Table 2), and either color caught more beetles than the blue, green, red, or black traps.

Samples had male-biased sex ratios in all trap colors (Table 2). Other species collected

included C. pruinina and C. montanus.

Thirty-nine female L. canus were dissected from the 1 May, 2001 samples in the height

test. Of these females, 14 contained a spermatophore, 22 contained sperm but no

spermatophore, and in 3 females we failed to find either a spermatophore or sperm.

Table 1

Click beetles other than L. canus collected in Japanese-beetle traps during height test (summed

over sampling dates); M - males, F - females.

1.5 meters 0.9 meters 0.3 meters

MF MF M_F

29 March - 10 May 2000

Cardiophorus montanus Bland 0 l (ee i.

Dalopius sp. l 0 | 0 3.1

27 March - 22 May 2001

Ctenicera pruinina l 0 6 0 2 O

Cardiophorus montanus l 0 5 0 10 4

Melanotus sp. 0 0 0 l 0 O

Table 2

Click beetles collected in Japanese-beetle traps of different colors. Numbers are totals for the

7-week (2000) or 10-week (2001) sampling periods. M - males, F - females. Six traps per

color. Analysis of variance for the 2001 L. canus data (summing data for sexes) showed

significant differences among colors (F = 13.9; df = 5,25; P < 0.001). An LSD-test showed

significantly (P < 0.05) higher catches in yellow traps than in white traps, which in turn had

significantly more beetles than green, black, blue, or red traps.

Yellow White Green’ Black Blue’ Red

M_F M FM FM FM F MF

29 March - 10 May 2000

Limonius canus 5 4 0 > 0 2 0 1 0 1 0O

Ctenicera pruinina 0 0 0 0 0 0 0 270 0) 30

Cardiophorus montanus 5 2 | 2 | 4 | 5 1 0 0

27 March - 22 May 2001

Limonius canus 43 12 24 11 14 3 #13 4 11 +5 9 5

Ctenicera pruinina 0 0 0 0 4 1 0 O20 I

Cardiophorus montanus 1 0 0 0 1 0 O 1 0 1 0 0

—

cn)

240 J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

DISCUSSION

There is a need for both basic and applied research on the Elateridae to improve our

understanding of these insects but also to provide the knowledge necessary to develop more

effective management programs for the pest species (Jansson and Seal 1994). In many pest

Elateridae, the larval stage has received considerably more study than the adult stage (Boiteau

et al. 2000). This bias may reflect the difference in longevity between larvae and adults, and

also the fact that adult click beetles do little damage. Thus, for many pest Elateridae, we still

lack basic information about the biology of adults, including details about phenology, mating

behavior and sex pheromones, fecundity, egglaying, feeding habits, and dispersal.

Several of these research topics, particularly phenology and dispersal, require having tools

that can be used in the field to monitor movements by the adult beetle. A variety of techniques

have been used to monitor flying click beetles, such as sweep nets (Shirck 1942), water pan

traps (Iwanaga and Kawamura 2000), window interception traps (Boiteau ef al. 2000), sticky

traps (Furlan 1996), and funnel-vane traps (Iwanaga and Kawamura 2000). For species that

spend most or all of their adult lives on or near the ground, adult beetles have been trapped

using mats of cut vegetation (Gough and Evans 1942; Roebuck ef al. 1947) and pitfall traps

(Doane 1963). These different sampling methods have been used to determine direction of

flight (Lafrance 1963), dispersal distances (Doane 1963), phenology (Lafrance 1963),

reproductive status of females (Shirck 1939), and response to chemical attractants (Iwanaga

and Kawamura 2000).

We tested whether traps that were designed to monitor Japanese beetles could be used to

monitor flight of L. canus and other Elateridae. Japanese-beetle traps are commonly used to

monitor flight activity of Coleoptera other than the Japanese beetle, particularly other

Scarabaeidae (Crocker et al. 1999). Metzger and Sim (1933) listed species and numbers of

Elateridae that were caught in Japanese-beetle traps placed in an area of New Jersey, and

showed that large numbers of a species of Melanotus were collected in the traps. The

Elateridae that we collected were composed mostly of L. canus, but we also collected small

numbers of other species. The low numbers of these other species were probably due to low

densities in the study area rather than to trap inefficiency. That is, larval collections made at

the station over the last several years have been mostly L. canus (unpubl.).

Limonius canus adults were active between mid-March and mid- to late-May (Fig. 1).

Emergence of the adults in March was probably in response to warming soil temperatures, as

suggested for other Elateridae (Lafrance 1963). Flight appears also to have been affected by

temperature, because trap catch dropped substantially in late March and early April 2001

coinciding with 2 weeks of maximum air temperatures below 20° C (Fig. 1). Shirck (1939)

stated that cool conditions prevented flight by a closely related species, the sugarbeet

wireworm (Limonius californicus (Mannerheim)), and Doane (1961) showed that numbers of

Ctenicera destructor (Brown) captured in funnel traps dropped in cool weather. Because of the

cooler temperatures in 2001, the peak catch of L. canus occurred almost 3 weeks later in 2001

than 2000 (Fig. 1). Also, total trap catch over the duration of the study was lower in 2001 than

2000, largely because of the reduced flight activity of male L. canus during April 2001. The

sex ratios (Figs. 2-3) show either that males emerged earlier in spring than females, or that

males were more likely than females to engage in flight during March and early April. Others

have shown that male Elateridae emerge in spring before females (Stone 1941; Shirck 1942;

Zacharuk 1962). Male Limonius agonus (Say) emerge | to 3 days earlier than females (Begg

1962). Seasonal totals of beetles in the height test were composed of relatively more males in

2000 (77.4% of beetles) than in 2001 (60.8%), probably due to the reduced catch of males in

April 2001 associated with cooler temperatures.

Efficient use of Japanese beetle traps to monitor L. canus requires information about the

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 4]

roles of trap color and placement in affecting catch. Although based upon relatively small

numbers of beetles, traps that had been painted yellow collected significantly more L. canus

than traps painted white, which in turn collected significantly more beetles than traps painted

green, red, dark blue, or black. These results are similar to those of Furlan (1996), who showed

that yellow or white sticky traps collected more click beetles (Agriotes ustulatus Schiller) than

red, black, or green traps. Other phytophagous Coleoptera show a preference for yellow over

other colors (Fleming et a/. 1940; Cross et al. 1976). We also showed that more L. canus were

collected in traps set at 0.3 m and 0.9 m than at 1.5 m above ground (Figs. 2-3). Boiteau et al.

(2000) used interception traps to collect over 40 species of Elateridae, and noted that trap

catch decreased with increasing height of the traps. The exact relationship between trap height

and trap catch varied with species. Furlan (1996) showed that sticky traps placed just above

the tops of vegetation collected more A. ustulatus than traps at higher elevations.

Lastly, almost all of the females collected at the beginning of their flight period had been

mated (at least 36 of the 39 females that were dissected contained sperm or a spermatophore).

Limonius canus evidently mates soon after emergence in spring, as noted for the congeneric L.

californicus (Stone 1935, 1941). Click beetles in other genera also mate soon after emergence

from the soil (Cohen 1942; Zacharuk 1962). In C. destructor, the female may be mated while

still below the soil surface (Zacharuk 1962). However, because we do not know the age of the

females that were trapped in this study (1.e., we have no data on emergence), and because we

do not know the amount of time between the day that a female was mated and the day that she

was trapped, we cannot say when, following emergence from the soil, mating occurred. Begg

(1962) showed that female Limonius agonus was most dispersive relatively late in the

Oviposition period, and it may be that L. canus has a similar life history. If so, the female

beetles that we collected may have been mated long before being trapped.

ACKNOWLEDGMENTS

We thank Deb Broers, Dan Hallauer, Toni Hinojosa, Richard Lewis, and Tamera Lewis

for assistance in the field and laboratory. We are also very grateful to Dr. Paul Johnson (South

Dakota State University) for his generous help in providing species’ identifications. The

comments of Brad Higbee and Tom Weissling on an earlier version of this manuscript are

appreciated. This research was partially supported by a grant from the Washington State

Potato Commission.

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Ohira (Coleoptera: Elateridae). Applied Entomology and Zoology 35: 283-285.

Jansson, R.K. and D.R. Seal. 1994. Biology and management of wireworms on potato, pp 31-53. In: G.W.

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J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 243

Qualitative analyses of larval oral exudate from eastern and

western spruce budworms (Lepidoptera: Tortricidae)

L.M. POIRIER

BIOLOGY PROGRAM, FACULTY OF NATURAL RESOURCES AND ENVIRONMENTAL

STUDIES, UNIVERSITY OF NORTHERN BRITISH COLUMBIA,

3333 UNIVERSITY WAY, PRINCE GEORGE, BC, CANADA V2N 4Z9

J.H. BORDEN

CENTRE FOR ENVIRONMENTAL BIOLOGY,

DEPARTMENT OF BIOLOGICAL SCIENCES, SIMON FRASER UNIVERSITY,

8888 UNIVERSITY DRIVE, BURNABY, BC, CANADA VSA 1S6

ABSTRACT

A two-choice feeding bioassay was used to investigate the effects of dilution,

centrifugation, storage and autoclaving on the repellency of the oral exudate of eastern

and western spruce budworms, Choristoneura fumiferana (Clem.) and C. occidentalis

Free., to their respective conspecifics. The exudate from insects reared on either

artificial diet or foliage was active at a volume equal to the amount emitted by one

larva when disturbed with a pipet in the laboratory, but repellency was lost at lower

doses. Centrifugation did not partition the exudate into active and inactive fractions.

Exudate from both diet- and foliage-reared insects was active for at least 48 h at room

temperature. However, after being frozen for one month, exudate from diet-reared

insects was still active, while that from foliage-reared insects was not.

Key words: Choristoneura fumiferana, Choristoneura occidentalis, spruce budworm,

oral exudate, regurgitant, epideictic pheromone

INTRODUCTION

Many insects release oral secretions when they are disturbed or handled (Davies and

McCauley 1970, Corbet 1971, Eisner et a/. 1974). While these secretions may serve to

protect the emitters from predators and parasitoids (Eisner et al. 1974), they may also be

used as epideictic pheromones (Prokopy 1981), to repel competing individuals from

potentially overcrowded resources (Corbet 1971). Larvae of the spruce budworm and the

western spruce budworm, Choristoneura fumiferana (Clem.) and C. occidentalis Free.,

respectively, have been shown to produce an oral exudate that repels conspecifics (Poirier

and Borden 1995). In two-choice feeding bioassays, larvae of both species avoided feeding

stations that had been treated with conspecific exudate (Poirier and Borden 1996). Larvae

of both species responded to con- and heterospecific oral exudate in the same manner, as

did laboratory-reared and wild-caught larvae. Larvae reared on artificial diet responded to

exudate from both diet- and foliage-reared larvae, whereas foliage-reared larvae responded

only to exudate from other foliage-reared larvae (Poirier and Borden 2000).

When the bioactive components are known, it may prove possible to exploit the activity

of the oral exudate in the management of spruce budworms. Isolation and identification of

the active components (Brand ef a/. 1979) can be facilitated by an understanding of the

properties of the exudate. We investigated the threshold concentration for repellency,

whether or not the active components are carried in the exudate in suspension or solution,

244 J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

the thermostability of the repellent, its persistence under bioassay conditions, and the

duration the exudate may be stored without losing its bioactivity.

METHODS

Insects. Laboratory-reared insects of both species were obtained as eggs or diapausing

second instars from the Canadian Forest Service, Sault Ste. Marie, Ontario, from colonies

maintained under laboratory conditions for many years at high population densities. Wild

C. occidentalis larvae were collected in June 1994 near Kamloops, BC by L.M.P.

All insects were reared in the laboratory at approximately 24°C, 60% RH, and a

photoperiodic regime of 16L:8D. Diet-fed larvae of both species were reared on an agar-

based artificial spruce budworm diet (Bio-Serv Inc., Frenchtown, New Jersey) in 30 ml

disposable vials. Two larvae were kept in each vial. Foliage-fed C. occidentalis larvae were

reared on potted 3-year-old Douglas-fir, Pseudotsuga menziesii (Mirb.) Franco.

Bioassay. Each experiment (Exp.) employed a two-choice, diet-station bioassay,

modified from Poirier and Borden (1996) to preclude any interaction between organic

solvents and the diet stations, if such solvents were to be used during chemical procedures.

Circular glass cover slips (18 mm diam.) were attached to the ceiling of the petri dish

bioassay chambers using chloroform, so that diet stations on the cover slips had their

centers 3 cm apart. Exudate or water treatments (2 yl) were applied to the glass in a ring

around the outer edge of one cover slip; third- to fifth- instar larvae produce an average of

2 wl of oral exudate when disturbed with a pipet in the laboratory (Poirier and Borden

1996). The other cover slip was left untreated. A drop of molten artificial diet was then

applied to the center of each cover slip within, but not contacting, the exudate treatment

ring. Separate dishes, with one station treated with 2 ul of distilled water, served as

controls. A conspecific third- to fifth-instar larva, randomized with respect to age, was then

placed in the center of the bottom of each petri dish chamber, and left undisturbed for 24 h

under the above rearing conditions. These test larvae were taken from the same food source

as the larvae that provided the exudate. At the end of 24 h, the stations were checked for

signs of feeding or establishment, such as feeding cavities or silk feeding tunnels. Larvae

that did not establish on a feeding station were included in the sample, unless they had

molted or pupated over the 24 h test period. Earlier studies by Poirier and Borden (1996,

2000) showed that non-feeding larvae provide important information about the degree of

repellency of various treatments. Larvae were thus categorized as not feeding, feeding on

the untreated station, or feeding on the treated station.

For each experiment, the numbers of larvae in the three categories were compared

between experimental and control dishes using Fisher's Exact Test for a 2 X 3 contingency

table, a =0.05 (Steel and Torrie 1980, Mehta and Patel 1983; Schlotzhauer and Littell

1987). Possible variation due to larval age was not taken into account.

Threshold for Bioactivity. Exp. | used third- to fifth-instar diet-reared C. fumiferana.

Exp. 2 used third- to fifth-instar wild-collected C. occidentalis, reared on foliage. Exudate

was collected by drawing it into a 5 pl micropipet after touching a larva with the pipet. The

exudate from several larvae was pooled in a glass vial kept on ice. The exudate was then

diluted 10-, 100- and 1000-fold by serial dilution in distilled water. One station in each

bioassay dish was treated with 2 ul of distilled water (control dishes), 2 ul of undiluted

exudate or 2 ul of one of the three exudate dilutions. Twenty dishes were prepared for each

of the five treatments. An uninduced test larva, ie. a larva that had not been induced to

produce exudate, was placed in each dish, and left undisturbed for 24 h.

Solubility. Exp. 3 used third- to fifth-instar diet-reared C. occidentalis larvae, from the

laboratory colony. Exp. 4 used third- to fifth-instar wild-collected C. occidentalis, reared

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 245

on foliage. For both experiments, exudate was collected from larvae using a 5 ul

micropipet, and pooled in three Eppendorf tubes kept on ice. Two tubes were centrifuged

at 13,000 rpm for 5 min in a Micro-Centaur benchtop Eppendorf centrifuge. The

supernatant was drawn off one tube with an Eppendorf Pipetman automatic pipet and

retained, and distilled water was added to the pellet fraction to bring it back to the pre-

centrifugation volume. In the second tube, the pellet was resuspended in the supernatant

with an automatic pipet. The third tube was left untreated. One station in each bioassay

dish was treated with 2 ul of either distilled water (control dishes), whole exudate,

supernatant, pellet suspension, or centrifuged and then reconstituted exudate. Forty dishes

(replicates) were prepared for each of the five treatments. An uninduced test larva was

placed in each dish, and left undisturbed for 24 h.

Storage Duration. Exp. 5 used third- to fifth-instar diet-reared C. fumiferana. Exp. 6

used third- to fifth-instar wild-collected C. occidentalis larvae, reared on foliage. On three

consecutive days, exudate was collected from larvae using a 5 pl micropipet, and pooled in

a glass vial kept on ice until the exudate was applied to the feeding stations. One feeding

station in each dish was treated with 2 ul of either distilled water or exudate. Test larvae

were introduced to the exudate-treated dishes. Exudate was added to the exudate-treated

dishes 0, 24 or 48 h prior to the introduction of test larvae. The water-treated control dishes

were tested immediately (0 h delay). The treatments were staggered so that larvae were

added to all dishes on the same day. The dishes for 24 and 48 h delayed testing were held

under the above rearing conditions until larvae were added. Twenty dishes (replicates)

were prepared for each of the four treatments in each experiment. An uninduced test larva

was placed in each dish, and left undisturbed for 24 h.

In Exp. 7, third- to fifth-instar, diet-reared C. fumiferana larvae were used. Exudate was

collected from larvae using a 5 yl micropipet, and pooled in glass vials kept on ice. One

vial was used immediately. The other vials were stored at approximately -4°C for one week

or one month. One feeding station in each dish was treated with 2 pl of either distilled

water (control dishes), fresh exudate, week-old exudate, or month-old exudate. Twenty

dishes were prepared for each of the four treatments. An uninduced test larva was placed in

each dish, and left undisturbed for 24 h.

Thermal Stability. All insects used in Exp. 8 were third- to fifth-instar diet-reared C.

occidentalis larvae from the laboratory colony. Oral exudate was collected from larvae

using a 5 pl micropipet, and pooled. Half the exudate was autoclaved for 15 min at 15 kPa

and 122°C. One feeding station in each dish was treated with 2 ul of distilled water

(control dishes), 2 ul of fresh exudate, or 2 ul of autoclaved exudate. Twenty dishes

(replicates) were prepared for each of the three treatments. An uninduced test larva was

placed in each dish, and left undisturbed for 24 h.

RESULTS

The results of all eight experiments are given in Table 1.

Threshold for Bioactivity. When the exudate was diluted by any of the three dilution

rates, the numbers of larvae in the three response categories were not significantly different

between experimental and control dishes in either experiment (Exp. 1, 2). Only larvae in

dishes treated with undiluted exudate were significantly deterred from feeding on the

treated station, with most larvae establishing on the untreated feeding station. Results were

similar for C. fumiferana reared on artificial diet (Exp. 1) and C. occidentalis reared on

foliage (Exp. 2).

Solubility. In Exp. 3, diet-fed larvae were significantly deterred from feeding only on

diet stations treated with whole or reconstituted exudate. There was no difference between

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

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control dishes and dishes treated with supernatant or the pellet resuspension. However, in

Exp. 4, the numbers of foliage-fed larvae in the three response categories were significantly

different from the control dishes in all treatments.

Storage Duration. In Exp. 5, more diet-reared larvae fed on the untreated than

exudate-treated station for all delay times, demonstrating persistent bioactivity of exudate

from diet-reared larvae for at least 48 h under laboratory conditions. However, in Exp. 6,

the bioactivity of exudate from foliage-reared larvae persisted for >24 h, but <48 h. When

the exudate from diet-reared larvae was frozen at -4°C, bioactivity persisted for at least a

week, but less than one month (Exp. 7).

Thermal Stability. The numbers of larvae in the three response categories were

significantly different from the control dishes in the fresh exudate treatment only (Exp. 8),

indicating that autoclaving for 15 min destroyed all deterrent activity.

DISCUSSION

Poirier and Borden (1995) showed that each larva of C. fumiferana and C. occidentalis

produced about 2 ul of oral exudate per induction. The results of Exp. 1 and 2 show clearly

that application of one larval equivalent of exudate is necessary to elicit a response from

test larvae in the laboratory bioassay. Because this biologically realistic dose is necessary

for bioassays, it is probable that large amounts of starting material will be needed for

chemical fractionation, bioassay and analysis (Brand et al. 1979), if the active constituents

in the exudate are to be identified.

The results from Exp. 3 and 4 were not conclusive. Centrifugation did not destroy the

bioactivity of exudate from either diet- or foliage-reared larvae, since reconstituted exudate

was as deterrent as whole exudate. Exudate from diet-reared insects may have had some of

the repellent components carried as a suspension, but the particulate matter alone was not

repellent. Exudate from foliage-reared insects appeared to have the repellent components

equally distributed between the pellet and the supernatant, with both fractions being as

repellent as whole exudate. It is possible that the centrifugation used was not sufficient to

separate the materials in the exudate, and that better results might have been achieved by

using higher centrifugation speeds or a longer duration. However, since the exudate

apparently consists of a partially digested substrate (Poirier 1995), it is also possible that a

portion of the repellent component is still associated with the particulate matter suspended

in the exudate, while the remainder is free in solution. Diet-reared larvae may produce

lower concentrations of repellent than foliage-reared insects or may lack an independently

repellent component that is present in the regurgitate of foliage-reared larvae (Poirier and

Borden 2000). In either case, for diet-reared larvae, dividing the exudate between two

fractions could reduce the concentration in one or both fractions below the response

threshold.

Exp. 5, 6 and 7 indicate that exudate from both diet- and foliage-reared larvae

deteriorates over time, although the breakdown appears to be faster in the latter case. While

this deterioration may be mediated by microbial activity, it seems likely that the repellent

components will also be subject to the action of larval enzymes contained in the

regurgitant. The retention of residual foliar enzymes in the regurgitant could account for

the shorter longevity of repellency in the regurgitant of foliage-reared larvae than in that of

diet-reared larvae. It is possible to store the exudate at -4°C for at least a week, but storage

at a lower temperature may increase longevity by halting further enzymatic activity.

Autoclaving appears to destroy the bioactivity of the exudate from diet-reared larvae

(Exp. 8). Because a number of materials may be significantly altered by high temperatures,

this result does little to indicate the type of compound involved in the repellency. However,

250 J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

the bioactive material may prove to be unstable in certain laboratory procedures, e.g. gas

chromatography, that involve heat.

These experiments provide considerable insight into the nature of the bioactive

constituent(s) in the larval oral exudate of these two Choristoneura spp. It persists for up to

48 h at room temperature, and survives storage when frozen at -4°C for at least one week.

However, it is thermally unstable under pressure at 122°C. When centrifuged, the bioactive

component(s) from foliage-reared larvae does not partition completely, suggesting that

some fraction remains associated with the particulate matter in the exudate, while some is

in solution. Lastly, below a biologically realistic concentration of one larval equivalent,

bioactivity is lost. Because diet can influence bioactivity (Poirier and Borden 2000), and

because the components of the artificial diet are known, modification of this diet could

potentially provide further insight into the nature of the bioactive component(s).

ACKNOWLEDGEMENTS

This research was supported in part by the Natural Sciences and Engineering Research

Council of Canada, The Canadian Forest Service, Phero Tech Inc., the Coast Forest and

Lumber Sector, the Cariboo Lumber Manufacturer's Assoc., the Interior Lumber

Manufacturer's Assoc., Canadian Forest Products Ltd., International Forest Products Ltd.,

Northwood Pulp and Timber Ltd., TimberWest Ltd., Tolko Industries Ltd., Western Forest

Products Ltd. and Weyerhaeuser Canada Ltd. The authors are grateful to Drs. G.J.R. Judd,

K.N. Slessor and A.S. Harestad for their advice and support and for review of the

manuscript.

REFERENCES

Brand, J.M., J.C. Young and R.M. Silverstein. 1979. Insect pheromones: a critical review of recent

advances in their chemistry, biology, and application. Progress in the Chemistry of Organic Natural

Products. 37: 1-190.

Corbet, S.A. 1971. Mandibular gland secretion of larvae of the flour moth, Anagasta kuehniella, contains

an epideictic pheromone and elicits oviposition movements in a hymenopteran parasite. Nature 232:

481-484.

Davies, R.W. and V.J. McCauley. 1970. The effects of preservatives on the regurgitation of gut contents

by Chironomidae (Diptera) larvae. Canadian Journal of Zoology. 48: 519-522.

Eisner, T., J.S. Johnessee, J. Carrel, L.B. Hendry and J. Meinwald. 1974. Defensive use by an insect of a

plant resin. Science. 184: 996-999.

Mehta, C.R. and N.R. Patel. 1983. A network algorithm for performing Fisher's exact test in r x c

contingency tables. Journal of the American Statistical Association. 78: 427-434.

Poirier, L.M. 1995. Bioactivity and characterization of spruce budworm larval oral exudate. Ph.D. thesis,

Simon Fraser University, Burnaby, British Columbia.

Poirier, L.M. and J.H. Borden. 1995. Oral exudate as a mediator of behaviour in larval eastern and western

spruce budworms (Lepidoptera: Tortricidae). Journal of Insect Behavior. 8: 801-811.

Poirier, L.M. and J.H. Borden. 1996. Repellency of oral exudate to eastern and western spruce budworm

larvae (Lepidoptera: Tortricidae). Journal of Chemical Ecology. 22: 907-918.

Poirier, L.M., and J.H. Borden. 2000. The influence of insect source and rearing medium on the

intraspecific repellency of larval oral exudate in eastern and western spruce budworms (Lepidoptera:

Tortricidae). The Canadian Entomologist. 132: 81-89.

Prokopy, R.J. 1981. Epideictic pheromones that influence spacing patterns of phytophagous insects. Pp.

181-213 In: D.A. Nordlund, R.L. Jones and W.J. Lewis (Eds.), Semiochemicals: Their Role in Pest

Control. Wiley, New York.

Schlotzhauer, S.D., and R.C. Littell. 1987. SAS system for elementary statistical analysis. SAS Institute,

Cary, NC.

Steel, R.G.D., and J.H. Torrie. 1980. Principles and procedures of statistics: a biometrical approach.

McGraw-Hill, Toronto.

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 251

Verbenone interrupts the response to aggregation

pheromone in the northern spruce engraver, [ps perturbatus

(Coleoptera: Scolytidae), in south-central

and interior Alaska

EDWARD H. HOLSTEN

U.S. DEPARTMENT OF AGRICULTURE, FOREST SERVICE, PACIFIC NORTHWEST

RESEARCH STATION, ANCHORAGE, AK, UNITED STATES 99503

ROGER E. BURNSIDE

STATE OF ALASKA, DEPARTMENT OF NATURAL RESOURCES,

DIVISION OF FORESTRY, ANCHORAGE, AK, UNITED STATES 99501

and STEVEN J. SEYBOLD

DEPARTMENTS OF ENTOMOLOGY AND FOREST RESOURCES,

UNIVERSITY OF MINNESOTA, ST. PAUL, MN, UNITED STATES 55108

ABSTRACT

Field tests of verbenone, a potential antiaggregation pheromone of the northern spruce

engraver, [ps perturbatus (Eichhoff), were conducted in south-central and interior

Alaska in stands of Lutz spruce, Picea x/uizii (Little), and white spruce, P. glauca

(Moench) Voss, respectively. Addition of 84%-(—)-verbenone at a high release rate to

the three-component aggregation pheromone of /. perturbatus (racemic ipsenol,

racemic ipsdienol, and 83%-(-)-cis-verbenol), significantly reduced trap catches. The

results of this study, combined with previous results on the presence of verbenone in

extracts of volatiles collected from feeding /. perturbatus and GC-EAD data, are

consistent with antiaggregant behavioral activity of verbenone for /. perturbatus.

Key words: Bark beetles, /ps perturbatus, semiochemicals, pheromones,

antiaggregation pheromones, verbenone, white spruce, Picea glauca, Lutz spruce,

Picea xlutzii, Alaska

INTRODUCTION

The northern spruce engraver, /ps perturbatus (Eichhoff) (Coleoptera: Scolytidae), is

distributed transcontinentally in the boreal region of North America, generally following

the distribution of white spruce, Picea glauca (Moench) Voss (Bright 1976; Wood 1982;

Robertson 2000). In Alaska it colonizes standing white spruce and Lutz spruce, Picea

x/utzii Little, that are stressed by natural disturbances such as drought, flooding, wind, ice,

and snow damage. Human activities such as logging and right-of-way clearance also

provide significant amounts of potential host material (Holsten and Werner 1987; Holsten

1996, 1997, 1998). Normally, endemic populations may infest individual standing spruce

trees, but during warm, dry summers following mild winters, engraver beetle populations

can increase significantly and kill groups of standing spruce trees. Historically, only limited

tree mortality has been caused by this beetle in Lutz spruce forests in south-central Alaska.

However, damage caused by /. perturbatus and other Jps spp. may assume greater

252 J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

economic importance as more habitat is provided through climate change and human

activities (Robertson 2000). For example, in 1996 more than 47% of the residual spruce in

a thinned area near Granite Creek in south-central Alaska became infested with /.

perturbatus and I. tridens (Mannerheim). Spring drought conditions as well as the recent

overall “warming” of the Kenai Peninsula apparently led to this rapid increase in Ips

activity in 1996 (Anon. 1999; Holsten 1996, 1997, 1998).

Increased tree mortality in Alaska caused by /ps spp. has stimulated research on new

management tactics utilizing semiochemicals. From 1977 through 1992, field tests of the

efficacy of various bark beetle semiochemicals showed that ipsdienol (2-methyl-6-

methylene-2,7-octadien-4-ol) and 2-methyl-3-buten-2-o0l were generally attractive to /.

perturbatus. \psenol (2-methyl-6-methylene-7-octen-4-ol), 3-methylcyclohex-2-enone, and

verbenone (4,6,6-trimethylbicyclo[3.1.1]hept-3-en-2-one) were found to be generally

inhibitory (Werner 1993). Seybold and co-workers found that 1 perturbatus produce

>99%-(-)-ipsenol, ~90%-(+)-ipsdienol, and cis-verbenol (unpublished data), while Holsten

et al. (2000) found that a combination of racemic ipsenol, racemic ipsdienol, and 83%-(—)-

cis-verbenol' was attractive to /. perturbatus in field assays.

The presence of verbenone in an extract of volatiles trapped from the headspace above

male and female /. perturbatus feeding on P. x/utzii, and verbenone’s activity in coupled

gas chromatographic-electroantennographic detection (GC-EAD) assays suggest that

verbenone may be behaviorally active for /. perturbatus. Verbenone collected from the

volatile headspace above feeding male and female /. perturbatus may be synthesized by the

insects, by microbes, or through autooxidation of host a-pinene (Seybold et al. 2000).

Verbenone has been shown to interrupt aggregation in other /ps spp. and in a variety of

other scolytids (Borden 1997). For example, laboratory bioassays with /. paraconfusus

Lanier (McPheron et al. 1997) showed that increasing concentrations of verbenone resulted

in slower responses by beetles reaching an attractant source of naturally produced male

pheromone volatiles. In limited field studies of verbenone’s effect on aggregation of /.

perturbatus, Werner (1993) showed a 19% reduction in trap catch when verbenone was

added to ipsdienol.

Building on semiochemical studies by Holsten et a/. (2000) and Werner (1988, 1993)

and in response to increased /ps activity in south-central Alaska in 1996, efforts to apply

Ips attractants and antiaggregants for population manipulation (Shea 1994; Salom and

Hobson 1995) have been renewed in south-central and interior Alaska.We tested the

efficacy of one enantiomeric blend of verbenone as an antiaggregant for /. perturbatus at

two locations in Alaska.

MATERIALS AND METHODS

Study area characteristics. In the south-central Alaska site (Kenai Peninsula, 150 km

south of Anchorage), characterized by a transitional climate, traps were placed among Lutz

spruce trees located at 250 m elevation in the Granite Creek campground area. This stand

contained trees that were about 90 years old with a mean diameter at 1.3 m height of 7.5

cm, a mean height of 10 m, and a stand density of about 600 per ha. Shrub cover was

sparse, consisting mostly of blue-joint reedgrass, Calamagrostis canadensis (Michx.)

Beauv., and Salix spp.

In the interior site, characterized by a continental climate, traps were placed among

white spruce trees located at 500 m elevation near Tok. This stand contained trees that

' The enantiomeric composition of cis-verbenol in Holsten et al. (2000) was incorrectly

reported as 83%-(+). It was 83%-(-).

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 O53

originated from a fire about 80 years ago and had recently been thinned to about 1400

stems per ha, including 20 stems per ha of quaking aspen, Populus tremuloides Michx..

White spruce trees had a mean diameter at 1.3 m height of 9.5 cm and a mean height of 10

m. Shrub cover was sparse with only green alder, A/nus crispa (Ait.) Pursh, crowberry,

Empetrum nigrum L., Labrador tea, Ledum groenlandicum L., mountain cranberry,

Vaccinium vitis-idaea L., and highbush cranberry, Viburnam edule (Michx.) Raf.,

occupying the site.

Trap Placement and Semiochemicals. Twelve-unit, multiple funnel traps (Lindgren

1983) were hung from branches of non-host or dead spruce trees, or on nylon rope

suspended between host trees. Traps at both sites were hung at least 10 m apart (Bakke ef

al. 1983) with collection containers 0.3 m aboveground. Traps were baited with

semiochemicals (Table 1) dispensed from polyethylene bubble cap release devices

(PheroTech, Inc., Delta, BC, Canada’). To reduce cost, racemic semiochemicals were used

where available. The three-component attractant of ipsenol, ipsdienol, and cis-verbenol was

used because it is more attractive than ipsdienol alone (Holsten et a/. 2000). Each

component of the attractant and verbenone were released from separate bubble caps.

Beetles were collected from traps weekly from late May through July. Trapped insects

were placed in labeled plastic bags and frozen for later identification and counting.

Table 1

Release rates of synthetic semiochemicals used in /ps perturbatus trapping studies, Alaska,

2000'.

Enantiomeric Bubble cap Bubble cap

Semiochemical composition load (mg) release rate

(mg/day)”

Ipsdienol Racemic 40 0.2

Ipsenol Racemic 20 0.2

cis-verbenol 83%-(-) iS 0.6

Verbenone 84%-(—) 790 8

'All semiochemicals have chemical purity >98 percent.

? Release rates determined at 22°C.

Experimental Design and Statistical Analyses. Treatments were completely randomized

in each field test and were initially replicated at least ten times at each location. However,

in some instances trap catches were discarded from the experiment because neighboring

trees became infested with / perturbatus and these natural aggregations influenced trap

catches. Thus, the number of replicates varied from 9 to 10 (Granite Creek) or 7 to 10

(Tok) (Table 2). Treatments were: 1) Attractant (ipsenol + ipsdienol + cis-verbenol), 2)

Attractant + high dosage (two bubble caps) of verbenone’, 3) Attractant + low dosage (one

bubble cap) of verbenone, 4) Verbenone alone (low and high dosages used at Granite

Creek; low dosage used at Tok), and 5) Unbaited traps as controls. Statistical analyses

* The use of trade or firm names in this publication is for reader information and does not

imply endorsement by the U.S. Department of Agriculture of any product or service.

* As the enantiomeric composition of verbenone associated with feeding /. perturbatus has

not yet been determined, we used “standard” verbenone bubble caps (PheroTech Inc.)

typically applied for mountain pine beetle, Dendroctonus ponderosae.

254 J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

were completed using “Statistix 7” software’. Numbers of /. perturbatus caught by each

treatment were first examined by the Shapiro-Wilk Test to determine whether data

conformed to a normal distribution. Since they did not, data were transformed using the

natural log + 1 before being subjected to ANOVA followed by Tukey’s (1953) comparison

of means test (<= 0.05). Untransformed means are reported in the results.

Table 2

Effect of 84%-(—)-verbenone on the response of /ps perturbatus to an attractant composed

of racemic ipsdienol, racemic ipsenol , and 83%-(—)-cis-verbenol in multiple funnel traps,

Granite Creek and Tok, Alaska, 2000’.

No. of beetles caught (mean + SE)?

Treatment Granite Creek (n°) Tok (n)

Attractant 288.8 + 66.7 *(10) 131 51008507)

Attractant + Low Verbenone 112.0 + 16.9 *(10) 194.6 + 47.0 °(7)

Attractant + High Verbenone 46.5+8.7° (9) 136.2 +23.5° (8)

High Verbenone 12+04° (9)

Unbaited trap 16 0:8 “C10) 5.9 +-3.3 (10)

' Granite Creek trapping study had one additional treatment; high release rate of verbenone

alone (two release devices).

* Mean and standard error values followed by same letter within each column are not

significantly different, P< 0.05, Tukey’s comparison of means test.

> Number of replications

RESULTS

The overall effect of treatment was significant at both locations (Granite Creek, F =

98.2, df = 5,52, P < 0.001; Tok, F = 64.6, df = 4,36, P < 0.001). The ternary blend of

racemic ipsdienol, racemic ipsenol, and 83%-(-)-cis-verbenol was significantly more

attractive to /. perturbatus than the unbaited trap (Table 2). Because sexes of J. perturbatus

cannot be differentiated by external morphology, the sex ratio of the captured beetles was

not determined. Verbenone released at a high rate reduced mean trap catches at both Tok

and Granite Creek by a factor of five relative to the attractant (Table 2). Addition of

verbenone at a low release rate also significantly reduced trap catches of /. perturbatus at

Tok (Table 2). There was no significant difference at either location between the responses

to verbenone alone and to the unbaited control (Table 2).

DISCUSSION

The dose-dependent effect of verbenone on the response of /. perturbatus to its

attractant (noted at Granite Creek) has also been demonstrated for other species of Ips

(Miller et al.1995, McPheron ef al.1997). With 1. perturbatus, Werner (1993) showed a

19% reduction in trap catch when verbenone [87%-(—) and 5 mg/day] was added to

racemic ipsdienol (0.2 mg/day). However, we have demonstrated higher reductions (62%

to 84%) in trap catches when verbenone [84%-(—) and 3.5 or 7 mg/day] was added to the

three-component attractant. The enantiomeric blends and release rates of verbenone used

by us and by Werner (1993) were similar, while the ipsdienol used in each study was

* “Statistix 7”, 2000, Analytical Software, PO Box 12185, Tallahassee FL 32317-2185.

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 255

identical in chemical composition and release. Differences in the effect of verbenone on

trap catches between our study and that of Werner (1993) could have been due to variation

in [. perturbatus populations, to differences in trapping technique, or particularly to

differences in attractant used as the positive control for interruption. In as much as Werner

(1993) only used one of the three components we used in our attractant, it is possible that

his 19% reduction might have been higher if all three pheromone components had been

used.

We now have identified an antiaggregant for /. perturbatus (verbenone) that is

associated with feeding /. perturbatus adults (unpublished data). Although we do not know

the exact enantiomeric composition of verbenone associated with /. perturbatus or the

timing of its release during host colonization, we have achieved a significant reduction in

trap catches using the commercially available, and relatively inexpensive verbenone bubble

cap containing 84%-(—)-verbenone. If verbenone is released late in the colonization

process, it may function naturally as an antiaggregation pheromone to minimize further

attacks on host material. As has been demonstrated for /ps pini (Say) in lodgepole pine,

Pinus contorta latifolia (Engelmann) Critchfield (Borden et al. 1992; Devlin et al. 1994),

treatment of white and Lutz spruce logging debris with commercially available verbenone

may reduce the level of colonization of host material and population increase by this forest

pest.

ACKNOWLEDGMENTS

We thank John Hard, USDA Forest Service PNW Research Station (ret.), Kathy

Matthews and Ken Zogas, Forest Health Management, State and Private Forestry, Alaska

Region, Anchorage, Alaska, for help with the installation of the field studies and trap

collections, personnel from Tanana Chiefs, Fairbanks, Alaska, for assistance in the field

studies conducted in Tok, and K.E. Gibson, A.D. Graves, P.J. Shea, B.L. Strom, and two

anonymous reviewers for review of an earlier draft of this paper. Manuscript preparation

and laboratory research on /. perturbatus by S.J. Seybold were supported by Cooperative

Agreements #PS W-0028-CA and #USDA/FS-00-CA-11272138 between the USDA Forest

Service Pacific Southwest Research Station and S.J. Seybold.

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Borden, J.H., D.R. Devlin, and D.R. Miller. 1992. Synomones of two sympatric species deter attack by the

pine engraver, [ps pini (Coleoptera: Scolytidae). Canadian Journal of Forest Research 22: 381-387.

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Devlin, D.R. and J.H. Borden. 1994. Efficacy of antiaggregants for the pine engraver, /ps pini (Coleoptera:

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Holsten, E.H. 1996. Jps tridens: a pest of managed stands? FHP Biological Evaluation R10-TP-64.

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TP-71. Anchorage, AK: U.S. Department of Agriculture, Forest Service, Alaska Region. 7 pp.

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McPheron, L.J., S.J. Seybold, A.J. Storer, D.L. Wood, T. Ohtsuka, and I. Kubo. 1997. Effects of

enantiomeric blend of verbenone on response of /ps paraconfusus to naturally produced aggregation

pheromone in the laboratory. Journal of Chemical Ecology 23: 2825-2839.

Miller, D.R., Borden, J.H., and B.S. Lindgren. 1995. Verbenone: Dose-dependent Interruption of

pheromone-based attraction of three sympatric species of pine bark beetles (Coleoptera: Scolytidae).

Environmental Entomology 24: 692-696.

Robertson, I.C. 2000. Reproduction and developmental phenology of Jps perturbatus (Coleoptera:

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of bark beetle infestations—proceedings of an informal conference. In: Annual meeting of the

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GTR-318. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Research

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Seybold, S.J., J. Bohlmann, and K.F. Raffa. 2000. Biosynthesis of coniferophagous bark beetle

pheromones and conifer isoprenoids: Evolutionary perspective and synthesis. The Canadian

Entomologist 132: 697-753.

Shea, P.J. (Technical Coordinator). 1994. Proceedings of the symposium on management of the western

bark beetles with pheromones: Research and Development; 1992 June 22-25; Kailua-Kona, Hawaii.

Gen. Tech. Rep. PSW-GTR-150. Albany, CA: U.S. Department of Agriculture, Forest Service.

Pacific Southwest Research Station. 53 pp.

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Werner, R.A. 1993. Response of the engraver beetle, Jps perturbatus, to semiochemicals in white spruce

of interior Alaska. Res. Pap. PNW-GTR-465. Portland, OR: U.S. Department of Agriculture, Forest

Service. Pacific Northwest Research Station. 9 pp.

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J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 257

Geographic and temporal distribution of Agriotes obscurus

and A. lineatus (Coleoptera: Elateridae) in British Columbia

and Washington as determined by pheromone trap surveys

BOB VERNON

AGRICULTURE AND AGRI-FOOD CANADA, PACIFIC AGRI-FOOD

RESEARCH CENTRE, AGASSIZ, BC, CANADA VOM 1A0

ERIC LAGASA

WASHINGTON STATE DEPARTMENT OF AGRICULTURE,

OLYMPIA, WA, UNITED STATES 98504-2560

HUGH PHILIP

BC MINISTRY OF AGRICULTURE, FOOD & FISHERIES,

KELOWNA, BC, CANADA V1X 7G5

ABSTRACT

Initial and peak catch of male Agriotes obscurus (L.) in pheromone traps in the lower

Fraser Valley of BC occurred 15.6 and 18.7 days, respectively, before A. lineatus (L.).

Both A. obscurus and A. lineatus were taken in pheromone traps from each of 77 fields

monitored throughout the lower Fraser Valley during 2000 and 2001, expanding the known

ranges of both species. No specimens of either species were taken in pheromone traps set

at 56 sites distributed throughout the Okanagan, Similkameen and Nichola Valleys in 2000,

but one specimen of A. lineatus was found in a private collection that had been captured

near Merritt BC. This is the first record of A. lineatus in BC outside of the lower Fraser

Valley. Of nine counties surveyed in Washington State in 2000, A. obscurus was taken in

traps at several sites in Whatcom county, especially along the Canada/US border, and A.

lineatus was taken at several sites in Whatcom, Snohomish and Pierce counties.

Key words: wireworms, Elateridae, A griotes lineatus, Agriotes obscurus, pheromone

traps

INTRODUCTION

In 1952, a proceedings of the Entomological Society of British Columbia was published

to commemorate the fiftieth anniversary of the society. In that issue, an article entitled, ‘List

of the Elateridae of British Columbia’, was published by M.C. Lane, an entomologist from the

Bureau of Entomology and Plant Quarantine in Walla Walla, Washington. Among the 150

species known to be in BC at that time, Lane listed the dusky click beetle, Agriotes obscurus

(L.) (Coleoptera: Elateridae), and the lined click beetle, A. /ineatus (L.), as being present on

Vancouver Island but not on the mainland (Lane 1952). These species had just been

discovered in 1949 by King (1950) at Cobble Hill, near Victoria, and shortly thereafter, 4.

obscurus was found on the mainland at the eastern end of the lower Fraser Valley near Agassiz

(King et al. 1952). By 1980, A. obscurus larvae and related damage had been reported from

several farms in Surrey, about 70 km west of Agassiz, and A. lineatus had spread from

Vancouver Island to Vancouver on the mainland (Wilkinson 1980). In a recent survey

conducted in the lower Fraser Valley in 1996 and 1997, A. obscurus was taken in pitfall traps

258 J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

from several locations between Delta and Agassiz (Vernon and Pats 1997). The easternmost

record of A. obscurus was in Laidlaw, between Hope and Agassiz. The survey also showed

that A. /ineatus was now established in Delta, which was the only region where the two species

overlapped in pitfall traps in the lower Fraser Valley. A single specimen of A. obscurus caught

in a pitfall trap in a field of raspberries in Lynden, WA in 1997 was the first recorded

occurrence of this species in Washington State (Vernon and Pats 1997). Other than the

Lynden capture, neither species has been reported outside of the lower Fraser Valley in BC, or

elsewhere in Washington.

The initial discoveries of A. obscurus and A. lineatus in BC (King 1950; King et al. 1952)

were of particular importance at that time, since both were introductions from Europe, and

both were considered among Europe’s most destructive insects (Eidt 1953). It is believed that

both species were introduced to BC from Europe around 1900 (Wilkinson 1963), although the

actual time and means of introduction are not known for certain. It has been hypothesized that

A. obscurus larvae may have been introduced to the Agassiz area on hops with soil brought

from Europe (A.T.S. Wilkinson, personal communication). Introductions of A. obscurus, A.

lineatus and A. sputator (L.) to the east coast of Canada in the 1800s have been attributed to

the dumping of soil ballast from ships coming from Europe (Eidt 1953). Instances of ballast

dumping are also recorded in areas of Puget Sound and Portland Oregon in the US (Lindroth

1957), and in Departure Bay just north of Nanaimo on Vancouver Island (Scudder 1958).

Once established in Canada, the dispersal of A. obscurus and A. lineatus was believed to

have been slow due to the 4-year life cycle of these species (Wilkinson ef al. 1976). Following

its discovery in Agassiz, BC in 1952 (King et al. 1952), a subsequent delimitation survey

showed that the area affected was confined to less than 320 ha, which was bounded by the

Fraser River on the south and east and by heavily wooded terrain on the west and north.

(Wilkinson 1957). In Nova Scotia, introduced Agriotes spp. were initially restricted to the

vicinity of old ports that were also confined by water and forests (Eidt 1953). It was also

believed by workers studying these species in Nova Scotia and BC that dispersal was slow

because the adults had not been observed to fly (Wilkinson et al. 1976; Eidt 1953). With the

gradual urbanization of BC and Nova Scotia, however, several new manmade avenues of

dispersal appear to be accelerating the movement of wireworms into new areas. The

movement of wireworm-infested soil for example, either as topsoil for landscaping purposes,

or in soil associated with sod farms or ornamental plants has likely played a role in spreading

both species throughout the lower Fraser Valley of BC and beyond since the early reports

(R.S. Vernon, personal observation).

When organochlorine insecticides commonly used to control wireworms were withdrawn

from BC and elsewhere, Wilkinson (1980) predicted that European wireworms would

eventually become a serious threat to agriculture in BC. This prediction has come true,

particularly within the past decade where wireworm damage has increased dramatically in

small fruit, vegetable, ornamental and forage crops throughout the Fraser Valley. Wireworms

have been particularly damaging to potatoes in the lower Fraser Valley, where holes and scars

have reduced marketable yields on conventional farms, and where entire fields of organic

potatoes have been rendered unmarketable (R. S. Vernon, personal observation). Wireworms

are also a major concern to the strawberry industry, where considerable seedling mortality can

occur to newly established plantings, and where wireworms in mature plantings will enter fruit

in contact with the ground to become contaminants during processing. The damage is always

associated with A. obscurus or A. lineatus or both (Vernon and Pats 1997), but because these

species are extremely difficult to distinguish from each other using larval characteristics alone

(Wilkinson 1963), it is not known what species are damaging crops in certain areas.

The recent survey of A. obscurus and A. lineatus distribution in the lower Fraser Valley

and Lynden Washington by Vernon and Pats (1997), relied primarily upon historical Elaterid

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 259

collections at the Pacific Agri-Food Research Centre (AAFC) at Agassiz, and pitfall traps

placed in a total of 12 field sites during 1996 and 1997. In 1999, pheromone traps for A.

obscurus and A. lineatus were developed that are much more convenient and effective at

trapping adult male click beetles than pitfall traps (R. S. Vernon, unpublished data). In 2000,

these pheromone traps were used to survey the Okanagan, Similkameen and Nichola Valleys

of BC and western Washington for A. obscurus and A. lineatus. The traps were also used in

strawberry and potato fields throughout the lower Fraser Valley during 2000 and 2001, and the

spatial and temporal occurrence of A. obscurus and A. lineatus beetles in these three survey

areas is described in this paper.

MATERIALS AND METHODS

Pheromone trapping. The traps used in these surveys were Vernon Beetle Traps

(PheroTech Inc., Delta, BC) baited with bubble cap lures containing pheromone blends for A.

lineatus and A. obscurus (LaGasa et al. 2000). The pheromone lure formulations remain

proprietary information at the present time. The trap, constructed of durable polyvinyl

chloride (PVC), is designed to capture and confine adult male beetles that are attracted to the

internal pheromone lure and fall in after ascending shallow ramps. No killing agent or

preservative was used inside the traps, which relied on regular servicing to provide specimens

in good condition. Traps were placed at ground level, with entry ramps flush with or slightly

covered by adjacent soil to provide unimpeded beetle entry. Sample collection involved

removal of one of the ramp inserts and shaking the trap contents into a tray.

Lower Fraser Valley surveys. In the spring of 2000, pheromone traps for A. obscurus

and A. /ineatus were installed in 17 strawberry fields in the lower Fraser Valley from Delta to

Chilliwack (Fig. 1). Trap placement, spacing and numbers were dependent on field size and

shape, but generally 10 traps were installed along each of six evenly spaced rows of

strawberries to achieve good field coverage. Traps were consecutively numbered (e.g. | to

60) with odd- numbered traps baited with A. /ineatus lures and even-numbered traps with A.

obscurus lures. This was done for both species throughout the growing season in the

westernmost seven fields (Richmond, Delta and Surrey), whereas in the easternmost 10 fields

(Aldergrove, Abbotsford and Chilliwack), A. /ineatus traps were not installed until late May.

Traps were checked on a weekly to biweekly basis from mid-April to mid-July, during which

time all beetles were removed and saved for identification.

Additional pheromone traps purchased by growers were installed by private consultants in

headland areas surrounding 18 potato fields in Delta in 2000. Each field had a single pair of

A. obscurus and A. lineatus traps, with 10 m between the paired traps. These traps were

checked weekly from 13 April to 3 July, during which time all beetles were removed and saved

for identification.

In the Spring of 2001, pheromone traps for A. obscurus and A. lineatus were installed in

50 strawberry fields in the lower Fraser Valley, again from Delta to Chilliwack but also in

areas north of the Fraser River that had never been surveyed (Fig. 1). Seven of the fields

monitored in 2001 had also been monitored with pheromone traps during the 2000 growing

season. Each field had five pairs of A. obscurus and A. lineatus traps. Trap pairs were located

midway along each side of each field, about 10 m in from the field edges with about 10 m

between paired traps. Another pair of traps was located in the approximate center of each

field. Traps were checked on a biweekly basis from mid-March to mid-July, during which

time all beetles were removed and saved for identification. A 10-ha field of pasture in Surrey,

and a |-ha fallowed field in Agassiz were also monitored in 2001 for A. obscurus and A.

lineatus using the pheromone traps, with four of each trap placed at random locations inside

the fields.

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

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Interior BC surveys. From 19-26 May, 2000, six pairs of A. obscurus and A. lineatus

traps were established in the Similkameen Valley of BC between Keremeos and Osoyoos.

From 26 May to 23 June, 29 pairs of traps were established in the Okanagan Valley between

Osoyoos and Salmon Arm, and 13 pairs of traps between Salmon Arm and Kamloops from 23

June tol0 July. An additional eight traps were established between Merritt and Kamloops in

the Nichola Valley from 23-30 June. The traps were placed amongst grass in roadside ditches

along highways and country roads. Paired traps were placed either on the same side of the

road, and spaced 7-10 m apart, or placed on opposite sides of the road, and spaced 15-20 m

apart.

Washington State surveys. Traps were initially installed from mid-April to early-May,

2000, and were checked as frequently as possible until removal in July or early August. Trap

checking intervals varied from weekly in priority areas of Whatcom county, to a month or

more in southwestern Washington counties. At sites where high levels of the target beetles

were collected in the first trap checks in Whatcom County, traps were subsequently relocated

to more southern locations to gather additional delimiting information. Trapping sites in the

northern counties of Whatcom and Skagit were established in an approximate grid pattern,

with between 3.2 and 6.4 km between traps (Fig. 2).

The physical criteria for trap sites included: proximity to areas of turf, pasture, or other

grassy locations, which are considered favored wireworm habitat; and protected situations

where traps would be less likely disturbed or damaged. Outside of the northern counties, traps

were located near ports or nurseries where the target species may have been introduced

through shipping ballast or infested nursery stock.

RESULTS

Initial and peak emergence of A. obscurus and A. lineatus.

Fields of strawberries that were monitored simultaneously for A. obscurus and A. lineatus

in the Surrey region of the lower Fraser Valley in 2000 suggested that the peak activity period

of adult A. obscurus males preceded that of A. /ineatus. Unfortunately, numbers of A.

obscurus taken in pheromone traps in the Surrey fields (installed 13 April) were already high

when traps were first inspected on 19 April (range: 1.1-8.5 beetles per trap per day), indicating

that the initial emergence period of A. obscurus had been missed. For A. obscurus, the highest

recorded catches in traps occurred between 19-27 April in the Surrey fields (range: 1.6-8.5

beetles per trap per day), and between 19 April and 4 May in the Aldergrove, Abbotsford and

Chilliwack fields (range: 1.3-20.8 beetles per trap per day). Numbers of A. /ineatus in the

Surrey fields were quite low in traps on 19 April (range: 0-0.11 beetles per trap per day) with

peak catches occurring on 16 June (range: 0.9-1.9 beetles per trap per day). Trapping for A.

lineatus began too late in the Aldergrove, Abbotsford and Chilliwack fields to determine the

initial or peak activity periods.

Since it appeared that the initial emergence period of A. obscurus and A. lineatus was

missed in 2000, sampling began between 14-22 March in the 2001 strawberry field surveys.

Dates of first catch of A. obscurus varied considerably between the various regions monitored

in the lower Fraser Valley in 2001 (Table 1). The very earliest catch of A. obscurus occurred

on 23 March in five out of eight fields in Chilliwack at the eastern end of the lower Fraser

Valley (mean of eight fields = 28 March) with mean initial catch in other regions ranging from

3-28 April. Mean peak catch of A. obscurus ranged from 3-19 May (Table 1). The dates of

initial catch of A. obscurus are similar to those reported in Europe (i.e. United Kingdom,

France, Switzerland and Poland), where initial captures generally occurred in the last week of

March or first week of April (Cohen 1942).

The very earliest catch of A. /ineatus in pheromone traps was again in Chilliwack in two

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 263

out of six fields on 17 April (mean of six fields = 24 April) with mean initial catch in other

regions ranging from 24 April to 5 May (Table 1). Mean peak catch of A. /ineatus among

regions ranged from 25 May to 2 June (Table 1). The difference in mean initial and mean

peak catch between A. obscurus and A. lineatus in the lower Fraser Valley was 15.6 and 18.7

days, respectively.

Table 1

Mean number of Julian days required for initial and peak catches of A. obscurus and A.

lineatus in strawberry fields monitored with pheromone traps in eight regions of the lower

Fraser Valley in 2001. Regions are arranged from east to west.

Julian date of Julian date of

# initial catch it peak catch

Region Monitored Fields' A. obscurus A. lineatus Fields' A. obscurus A. lineatus

Chilliwack 8:6 86.5 MS 7 8:8 193 145.0

Sumas Prairie De) 105.8 117.6 5:4 122.8 148.0

Deroche 4:4 104.8 119.0 4:4 1325 150.0

Abbotsford 4:4 110.0 122.0 0:4 - 150.0

S. Aldergrove 17 106.0 119.1 eg 123.0 i335

N.Aldergrove 8:8 93.0 19'S 8:8 131.5 149.0

Surrey 35 107.6 119.6 5 131.6 149.0

Delta 6:6 117.8 125.0 7:7 ee 147.0

Mean 103.9 119.5 130.2 148.9

'The number of fields in each region with mean A. obscurus catch > 0.1 (left number) or A.

lineatus catch > 0.1(right number) beetles per trap per day, during initial or peak trap catch.

Geographic distribution of A. obscurus and A. lineatus.

Lower Fraser Valley. In the initial pheromone trap survey of 17 strawberry fields in

2000, A. obscurus and A. lineatus were both found in all fields sampled in Richmond (one

field), Delta (two fields), Surrey (four fields), Aldergrove (two fields) Abbotsford (two fields)

and Chilliwack (six fields) (Fig. 1). Of 18 headland areas surrounding potato fields monitored

with pheromone traps in Delta in 2000, both species were found in all but one field (i.e. on

Westham Island) which failed to capture any A. obscurus.

The 2000 survey findings were verified and expanded upon in the survey of 50 strawberry

fields in 2001, where both species were taken in pheromone traps in all fields monitored in

Delta (8 fields), Surrey (5 fields), Aldergrove (16 fields), Abbotsford (4 fields),

Mission/Deroche (4 fields), Sumas Prairie (5 fields) and Chilliwack (8 fields) (Fig. 1). Both

species were also taken in pheromone traps placed in a fallow field in Agassiz. The catches of

A. obscurus in Richmond and Mission/Deroche, and catches of A. /ineatus in Aldergrove,

Abbotsford, Mission/Deroche, Sumas Prairie, Chilliwack and Agassiz are the first records of

these species in these areas, and both species now appear to coexist throughout the lower

Fraser Valley.

The mean number and relative proportions of A. obscurus and A. lineatus in traps in

geographically distinct groups of fields varied somewhat throughout the lower Fraser Valley in

2001 (Fig. 1). The data show that numbers of A. obscurus caught in pheromone traps were

generally lower relative to A. /ineatus in the western half of the Valley, but were higher in most

of the sampled areas in the eastern half of the Valley. In the field sampled in Agassiz (north

east of Chilliwack) in 2001, the proportion of A. obscurus was very high (0.98).

Okanagan Valley, BC. No specimens of A. obscurus or A. lineatus were taken in

pheromone traps at any of the 56 sites sampled in the Okanagan, Similkameen or Nichola

Valleys of BC between 19 May and 10 July in 2000. However, based on the observations

264 J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001

above which showed that A. obscurus emerges and peaks earlier than A. lineatus (Table 1), the

traps that were set in the Nichola Valley from Merritt to Kamloops between 23-30 June, and

those set between Kamloops and Salmon Arm between 30 June and 10 July, may have missed

the A. obscurus and possibly the A. /ineatus adult generations. The negative results in traps

set in the Okanagan and Similkameen valleys, however, suggests that neither species has been

introduced to those regions.

Merritt, BC. A single specimen of A. /ineatus was found in the collection of Professor

J.H. Borden (Simon Fraser University), which was caught between 1990-93 in a bark beetle

pheromone trap at Miner Creek in an actively logged and reforested area 25 km southwest of

Merritt (collector, Alejandro Camacho-Vera). This is the first record of A. Jineatus in the

interior of BC outside of the lower Fraser Valley.

Washington State. Of the eight counties monitored in 2000, A. obscurus was collected

only in Whatcom county, where 18 out of 22 sites were positive and catches were highest near

the Canadian border (Fig. 2). At these sites, between 68 and 76 adult 4. obscurus were

captured during 7-day trapping intervals in late April. Agriotes lineatus had a broader

distribution, being captured in Whatcom (11 of 15 sites), Snohomish (5 of 8 sites) and Pierce

(2 of 7 sites) counties. The highest average number of beetles per positive trap was in

Snohomish county (21.2 beetles), followed by Whatcom county (2.5 beetles) and Pierce

county (1 beetle).

The disparate collections of A. /ineatus in this survey, occurring in three counties

separated by counties without collections, suggest the possibility of a disjunct population of

that species in parts of the Puget Sound area. The extent of collections that were recorded in

this survey, occurring as far south as the Fife area in Pierce County, clearly demonstrates that

A. lineatus is currently established in areas outside of the previously known infested areas of

BC. Whether the detected populations in Snohomish and Pierce Counties represent natural

spread from BC or are the result of independent introductions was not determined from this

survey.

DISCUSSION

Since their hypothesized introductions to Vancouver Island (A. obscurus and A. lineatus)

and Agassiz (A. obscurus) about a century ago (Wilkinson 1963), both species are now found

throughout the lower Fraser Valley of BC, and at least A. /ineatus is probably established near

Merritt in the interior. Both species are also well established in areas of Washington state.

The fact that A. /ineatus was not found in pitfall traps at most sites east of Delta BC in the

survey by Vernon and Pats (1997) shows the value of using pheromone traps as a delimiting

survey tool for these species. Pheromone traps indicated that both species were present in

virtually all fields monitored throughout the lower Fraser Valley, and the large numbers of

beetles caught in some fields (e.g. 60 traps placed in one Chilliwack field in 2000 caught over

25,000 A. obscurus) indicates that both species are very well established. Because both

species are polyphagous, and will feed among the roots of many crops, it is almost a certainty

that they are being introduced to new areas through the movement of soil or via the

transplanting of plants from infested areas. Agriotes spp. (either A. /ineatus or A. obscurus)

wireworms were found in soil surrounding cedar seedlings originating from the Chilliwack

area that were awaiting planting at a Harrison Lake park (just north of Agassiz), and were also

found in topsoil moved from a Chilliwack farm to a residence in Rosedale in 1996 (R:S.

Vernon, personal observation). The abundance of A. /ineatus and A. obscurus in the lower

Fraser Valley and Washington state, and the distant movement of soil or plants with soil (i.e.

ornamentals or seedlings for reforestation) will likely spread these pests rapidly to new regions

of BC, Washington and beyond.

J. ENTOMOL. SOC. BRIT. COLUMBIA 98, DECEMBER 2001 265

The observation that the time of emergence and peak catch of A. obscurus preceded that

of A. lineatus by 15.6 and 18.7 days, respectively, will help to better streamline surveys and

interpret data in the future. These observations are also important in the development of pest

management strategies that target the adult stage of these species.

ACKNOWLEDGEMENTS

We thank Elaine Goudie, Anita Behringer, Bill Hedges, Patrick Hertzog, Harold

Kamping, Jon Mullen, and ES Cropconsult Ltd. for setting and inspecting the pheromone

traps, and Todd Kabaluk for figure preparation. This project was funded in Canada through

collaborative research agreements between the Fraser Valley Strawberry Growers Association,

the BC strawberry processors, Investment Agriculture Foundation, the Lower Mainland

Horticultural Improvement Assn., PheroTech, Inc. and the Potato Industry Development

Committee with matching funds from the Matching Investment Initiative of AAFC, and in the

US by a 2000 Cooperative Agricultural Pest Survey grant from the USDA APHIS Western

Region.

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NOTICE TO CONTRIBUTORS

The Journal of the Entomological Society of British Columbia is an official annual publication of

the Entomological Society of British Columbia. This journal publishes original research articles and

notes dealing with all facets of the study of arthropods, the content of which is of regional origin,

interest, or application. All papers and notes published in the Journal of the Entomological Society

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