J. ENTOMOL. SOc. BRIT. COLUMBIA 114, DECEMBER 2017

Journal of the LIBRARIES

Entomological Society of British Columbia

Volume 114 December 2017 ISSN#0071-0733

Directors of the Entomological Society of British Columbia 2016-2017....................0. 2

Grape leaf rust mite, Calepitrimerus vitis (Acari: Eriophyidae), a new pest of grapes in

British Colatibian...... 2s ont a es eek ee ee A 3

Insect taxa named for the Rev. John H. Keen, early naturalist on the Queen Charlotte

Islands.and at Mietlakatia. itis: © clea else Be A, oo etre: Ferwamars hiwoameghe Doda Eas es 15

Western balsam bark beetle, Dryocoetes confusus Swaine (Coleoptera: Curculionidae:

Scolytinae), in situ development and seasonal flight periodicity in southern British

COMBA oor eer een ee thane Gia nee GL ae ancora ve aN ays

An updated and annotated checklist of the thick-headed flies (Diptera: Conopidae) of

British Columbia; the Yutoon. and Alas 82 Oe so ak dics Ven oun awe 38

Supercolonies of the invasive ant, Myrmica rubra (Hymenoptera: Formicidae) in British

OTT ec eee crc ee nt er ees awe ee ee, ne 56

SCIENTIFIC NOTES

First record of Aedes (Ochlerotatus) spencerii (Theobald) (Diptera: Culicidae) in the Yukon

va onn ah dalll DELETE SENS SO a ie Et eo Bm Ry eee a 65

Cold requirements to facilitate mass emergence of spruce beetle (Coleoptera:

Curculonnige) acuits ms teslaberatony.). lent atiles es oleae is co he pi 68

Production of epicormic buds by Douglas-fir in central British Columbia, Canada,

following defoliation by western spruce budworm (Lepidoptera: Tortricidae) .............. 73

NATURAL HISTORY AND OBSERVATIONS

Archilestes californicus McLachlan (Odonata: Zygoptera: Lestidae):a damselfly new to

Candda cr EL BR A ea Se RD ea See tos Sg tr ee ie

Evidence of established brown marmorated stink bug populations in British Columbia,

CBAOA: oc coccieieshsck donb eat shane a bh Vell « cece: Gee GI a MAIN CURE Bos ela ea hae) 83

An unusual specimen of the subgenus Lasioglossum Curtis from British Columbia, Canada

CER viene ner he i sh es anak hse wath s sak ela amacrine tes Gnle pee clan Sin Pew doe 87

First identifications of aphid and diamondback moth populations on wasabi in British

PRAIA eh acetate hg eae’ bs Ask au va atte Soo epic ares ed ees at odd Shaadi 93

SYMPOSIUM ABSTRACTS

Proceedings of the Pollination: Science and Stewardship Symposium........................ 97

Presentation Arai detes Ba Mia iss Lescadclh aaa CRS Clemens, as SARE ae 101

Symposium abstracts: Biological Control -A Safe Approach to Pest Management ...105

ei cae g denen can cdanadsasresanzencesisaass Inside Back Cover

J. ENTOMOL. SOC. BRIT. COLUMBIA 114, DECEMBER 2017 : e,

DIRECTORS OF THE ENTOMOLOGICAL SOCIETY

OF BRITISH COLUMBIA FOR 2016-2017

President:

Jenny Cory (president@entsocbc.ca)

Simon Fraser University, Burnaby |

Ist Vice President:

Lisa Poirier

University of Northern British Columbia

2nd Vice-President:

Tammy McMullan

Simon Fraser University

Past-President:

Brian Van Hezewijk

Pacific Forestry Centre, Victoria

Treasurer:

Ward Strong (membership@entsocbc.ca)

Ministry Forests, Lands and Natural Resource Operations, Vernon

Secretary:

Tracy Hueppelsheuser (secretary@entsocbc.ca)

B.C. Ministry of Agriculture, Abbotsford

Directors:

Tamara Richardson, Grant McMillan

Graduate Student Representative:

Dan Peach

Regional Director of National Society: Editor, Boreus:

Bill Riel Gabriella Zilahi-Balogh

Natural Resources Canada, Canadian Forest | (boreus@entsocbe.ca)

Service Canadian Food Inspection Agency

Web Editor: Editor, Emeritus:

Brian Muselle (webmaster@entsocbc.ca) Peter & Elspeth Belton

University of British Columbia, Okanagan Simon Fraser University

Campus

Society homepage: http://entsocbe.ca Journal homepage: http://journal.entsocbe.ca

Editorial Committee of the Journal of the Entomological Society of British Columbia:

Editorial Board: Marla Schwarzfeld, Bo

Editor-in-Chief: Dezene Huber Staffan Lindgren, Katherine Bleiker,

__ Gournal@entsocbc.ca) Ward Strong, Lisa Poirier, Lee Humble,

University of Northern British Columbia Bob Lalonde, Lorraine Maclauchlan,

Robert McGregor, Steve Perlman

Copy Editor: Monique Keiran Technical Editor: Jesse Rogerson

J. ENTOMOL. SOC. BRIT. COLUMBIA 114, DECEMBER 2017 3

Grape leaf rust mite, Calepitrimerus vitis (Acari:

Eriophyidae), a new pest of grapes in British Columbia

L. B. M. JENSEN!?”, D. T. LOWERY!2*, and N. C. DELURY!

ABSTRACT

The grape leaf rust mite, Calepitrimerus vitis (Nalepa), was first discovered in the

interior of British Columbia in 2009 on grape leaves from a commercial vineyard —

north of Osoyoos. Bronzing of grape leaves confirmed to be caused by C. vitis in

summer 2009 was followed by severely stunted shoots and distorted leaves in several

vineyards in spring 2010. Numbers and lengths of shoots and fruit clusters were

reduced significantly on vines infested with C. vitis. Earlier studies have shown that

outbreaks of C. vitis result from pesticide sprays targeted to other pests that damage

predator mite populations. Sprays of sulphur-based fungicides early in the season are

the recommended method of control.

INTRODUCTION

Grape leaf rust mite, Calepitrimerus vitis (Nalepa) (Eriophyidae), is a host-specific

pest of grapevines, Vitis vinifera L., (Anonymous 1968; Bernard et al. 2005; Walton et al.

2007) found in most grape-growing regions of the world, including Washington State

since 2002 and Oregon since 2004 (Prischmann and James 2005; Walton ef al. 2007).

Calepitrimerus vitis has not previously been reported on grapevines in British Columbia

(B.C.) and was not found during an extensive survey of vineyard pests conducted in the

Okanagan and Similkameen valleys in 1972 (Madsen and Morgan 1975).

Calepitrimerus vitis has been considered an economic pest of grapes only during the

past four decades, possibly the result of reduced use of sulphur-based fungicides that

provide effective control (Anonymous 1968; Barnes 1970; James 2007) and from

increased applications of pesticides that are harmful to predators that normally keep its

numbers in check (Winkler et al. 1972; Bernard et al. 2005; Schreiner et al. 2014).

Bronzing of grape leaves in late summer can appear significant but is not thought to

affect the current year’s growth or quality of the fruit at fall harvest (Anonymous 2005;

Reinert 2006; James 2007). However, leaf bronzing is a good indicator of the potential

for large overwintering rust mite populations to emerge the following spring and

continue feeding, resulting in damage to the developing buds, shoots and leaves (Bernard

et al. 2005; Prischmann and James 2005; James 2007). Significant economic injury can

occur to grapes if these mites are not properly managed. Feeding of overwintered C. vitis

in spring on developing buds and shoots results in what has been termed short shoot

syndrome or reduced spring growth (Bernard ef a/. 2005; Walton et al. 2007; Schreiner et

al. 2014), which is typified by severely stunted growth, shortened internodes, scarring of

shoots, curled and distorted basal leaves, and reduced fruit set. Severe infestations can

result in abortion of affected bunches and complete crop loss (Walton et al. 2007). The

relationship of impaired and damaged spring growth of grapevines to C. vitis feeding is

not always clear. Several other factors may also be responsible for restricted spring

growth, such as heavy thrips (Thysanoptera) feeding, winter freeze and herbicide damage

(Schreiner et al. 2014).

This paper documents the first confirmed discovery of C. vitis in the Okanagan Valley

of B.C. and the results of subsequent surveys to assess C. vitis abundance and its impact

' Agriculture and Agri-Food Canada, Summerland Research and Development Centre, Box 5000, 4200

Hwy 97, Summerland, British Columbia, Canada VOH 1Z0

2 Shared first authorship

3 Corresponding Author: tom.lowery@agr.gc.ca

4 J. Entomol. Soc. Brit. Columbia 114, December 2017

on vine growth. Grape growing and wine production are important industries in the

Okanagan Valley, with 8,060 acres currently planted in wine grapes, accounting for 84%

of B.C.’s vineyard acreage (British Columbia Wine Institute 2015). In order to provide

advice to growers on the best management practices for this emerging pest, we also

present information on recommended methods of control developed elsewhere that

mostly rely on early season applications of sulphur.

MATERIALS AND METHODS

Identification of Calepitrimerus vitis. Inspection on June 24, 2009, of grapevines at

a commercial vineyard north of Osoyoos (49° 05' 23" N, 119° 30' 39" W) that had heavily

bronzed leaves revealed the presence of eriophyid mites. Samples of these mites were

preserved in 70% ethanol and sent for species verification to Dr. F. Beaulieu, Canadian

National Collection of Insects, Arachnids and Nematodes, Agriculture and Agri-Food

Canada, Ottawa, and to Dr. J. Amrine, West Virginia University.

2009 Summer Survey. A survey for C. vitis was conducted during June 24 to

September 8, 2009, in twelve vineyard cultivar blocks in eight vineyards located

throughout the southern half of the Okanagan Valley from Summerland and Naramata in

the north to Osoyoos in the south (Figure 1), including three blocks (Osoyoos: Shiraz,

Merlot and Cabernet Sauvignon) adjacent to the vineyard block originally found to be

infested on June 24. Two of the original Osoyoos blocks and a heavily infested site in~

Naramata were sampled 2-3 times over the course of the field season. Leaf samples

consisted of 10 randomly selected leaves from each cultivar block. As per the protocol of

Walton ef al. (2007), mites from a sample were transferred to a glass plate by means of a

mite brushing machine (J. G. H. Edwards, Okanagan Falls, B.C.); the glass plate was

previously covered with a thin film of soap to immobilize the mites. The glass plate was

then placed on a grid and examined using a dissecting microscope for counting all mite

species and developmental stages.

2010 Spring Bud Dissections. In early spring 2010, buds from four vineyards found

to be infested with C. vitis in 2009 were dissected, and all mite species under individual

outer bud scales were counted. Vineyard managers of three of the four sampled vineyards

applied sulphur (Kumulus; 80% sulphur; 2.86—11.2 kg/ha; BASF) during the woolly bud

stage of development as per the recommended method of control (Anonymous 1968;

Bernard et al. 2005). Ten randomly selected canes from each sampled cultivar block were

pruned above the third bud and temporarily placed in cold storage for no more than one

week until the assessments. Mite counts were conducted on the most basal bud from each

pruned cane. This procedure was conducted once in March before the sulphur sprays and

again in April approximately two weeks after the sulphur sprays.

Damage Assessment. The effect of early season feeding by C. vitis on developing

shoots and fruit clusters was determined in a block of Chardonnay grapes at Okanagan

Falls that had one section of eight vine rows heavily infested with C. vitis and showing

symptoms of spring feeding damage (i.e., stunted and scarred shoots, small distorted

leaves, brown and shrunken fruit clusters, etc.). The two adjacent damaged and

undamaged sections of the block had been managed in exactly the same manner, except

that the vineyard manager had sprayed the heavily infested rows the previous year with

an undisclosed insecticide to control leafhoppers. No information about the spray

application was provided. The use of a recapture sprayer for the insecticide application

resulted in a clear differentiation between rows with and without C. vitis feeding damage.

Shoot lengths and counts of C. vitis, phytophagous thrips, and predatory mites on

stems and the basal leaf of 20 randomly selected shoots per row were assessed on May

18, 2010, from four of the damaged and four of the undamaged rows. Predatory mites

and phytophagous thrips were not identified to species. Data were collected from the

third to sixth row away from the dividing line between the adjoining damaged and

undamaged sections. Pre-harvest data were collected from half (one arm) of each of 20

J. ENTOMOL. SOC. BRIT. COLUMBIA 114, DECEMBER 2017 5

randomly selected vines per row from those same four rows, with numbers of shoots and

clusters counted and cluster lengths measured on September 15, 2010.

Statistical analysis. Shoot lengths, mite and thrips counts, and pre-harvest data from

damaged and undamaged vine rows were analyzed using one-way ANOVA. All mite and

thrips count data were transformed (V(X + 0.5)) before analysis (Zar 2010). Statistical

tests were performed using JMP Version 10 (SAS Institute Inc. 2013), with all statistical

error rates at a = 0.05.

Keremeos

q Cawston

s Kilometers

Legend

A City/Town

Survey Sites

Figure 1. Locations of vineyards surveyed for Calepitrimerus vitis in the south Okanagan

Valley, B.C. Counts of C. vitis, predatory mites, and tetranychid mites (numbers/leaf) for

corresponding cultivars at each survey site are located in Table 1.

J. Entomol. Soc. Brit. Columbia 114, December 2017

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J. ENTOMOL. SOc. BRIT. COLUMBIA 114, DECEMBER 2017 i

RESULTS AND DISCUSSION

Identification of Grape Leaf Rust Mite. The eriophyid mites found to be the cause

of bronzing or russeting of grape leaves in a commercial vineyard north of Osoyoos in

the south Okanagan Valley in the summer of 2009 were confirmed by two independent

experts to be C. vitis (see methods). This is the first record for B.C. Adults of this

eriophyid mite species, a relative of the grape erineum mite, Colomerus vitis

(Pagenstecher) (Eriophyidae), are approximately 0.15 mm long, light amber in colour,

broader at the front end, and somewhat wormlike in appearance (Lowery 2015; Figures

2, 3). Feeding by C. vitis during the summer causes bronzing or stippling that can appear

similar to spider mite (Tetranychidae) injury. Damage to leaves by C. vitis feeding is

mostly restricted to the upper leaf surface rather than the lower surface and, unlike

tetranychid mites, C. vitis does not produce webbing.

Figure 2. Adult grape leaf rust mite, Calepitrimerus vitis (Nalepa).

2009 Summer Survey. Of the eight vineyards sampled, C. vitis was found in several

geographically separated locations (Table 1). Subsequent sampling has found it to be

widespread in the valley with numbers as high as 998 mites per leaf. Numbers exceeding

3,000 per leaf have been recorded in Europe and Oregon on severely infested grapevines

(Schreiner et al. 2014). The presence of C. vitis in multiple locations suggests that it has

been present in south central B.C. for several years, but widespread movement by wind

and. with human activities (Duffner et a/. 2001) could have resulted in rapid dispersal.

Monitoring of C. vitis on three occasions in one of the surveyed blocks showed that

8 J. Entomol. Soc. Brit. Columbia 114, December 2017

numbers increased by early August and then declined in September (Table 1). Migration

of C. vitis from leaves to their overwintering sites under outer bud scales and bark after

the end of August agrees with reports by Walton ef al. (2007) and Schreiner et al. (2014).

Grape varieties vary in their susceptibility to C. vitis feeding (Anonymous 2005;

Bernard et al. 2005; Schreiner et al. 2014). Combined with cool spring temperatures,

cultivars that develop slowly are exposed to the mites for a longer period and so damage

may be more severe. Cabernet Sauvignon, a later developing cultivar, is reportedly more

susceptible to C. vitis than earlier cultivars such as Chardonnay (Anonymous 2005), but

our survey did not find large numbers of C. vitis on a Cabernet Sauvignon block adjacent

to an infested Shiraz block (Table 1). Of the three cultivars sampled in the original

vineyard, Shiraz leaves had the largest numbers of C. vitis. A Merlot block in Naramata

also had high numbers of C. vitis. A larger study would be required to determine

differences in susceptibility to C. vitis among cultivars under southern B.C. conditions.

There was no clear association between C. vitis numbers and numbers of tetranychid

mites. The highest tetranychid counts (12.8/leaf and 30.2 eggs/leaf) were recorded in a

vineyard (site B) where C. vitis was not found (Table 1). The second highest number of

C. vitis (191) occurred on August 5 in a block of Shiraz (vineyard H) that initially had

few tetranychids (2.0 eggs/leaf), but high tetranychid numbers were recorded from that

same site in August and September. Although low numbers of C. vitis were most often

associated with low numbers of tetranychids, the complexity of mite population |

dynamics combined with differing spray regimes confounds the relationship.

Figure 3. Ove

(Nalepa), feeding under grapevine bud scales in spring.

2010 Spring Bud Dissections. C. vitis were found in large numbers in spring under

the outer bud scales (Figure 3) of vines whose leaves had become bronzed the previous

summer. Monitoring of eriophyid mites is difficult due to their microscopic size and

cryptic nature (Schreiner ef al. 2014). As an alternative to bud dissections, mite counts on

J. ENTOMOL. SOC. BRIT. COLUMBIA 114, DECEMBER 2017 9

double-sided sticky tape applied around the bases of developing shoots has been used to

monitor C. vitis emergence in spring (Bernard et al. 2005), but the method was not found

by Walton eft al. (2007) to provide useful monitoring information. Scouting for signs of

leaf bronzing or stippling in summer followed by an assessment of mite numbers

provides a good indication of the need for control the following spring (Schreiner ef al.

2014). As an alternative to a mite-brushing machine, Schreiner et a/. (2014) developed a

‘rinse in bag’ system that extracted C. vitis from leaves into a small amount of ethanol or

isopropanol for counting.

Sprays of sulphur-based materials during the woolly bud stage of grape development

have been shown previously to effectively control C. vitis and prevent damage to

developing shoots (Anonymous 1968; Bernard et a/. 2005). Our dissections of buds in

spring also indicated that sulphur (Kumulus™) applied by growers at the woolly bud

stage was effective against C. vitis, as none were detected in the sprayed commercial

vineyard blocks two weeks post-application (Table 2). For the cultivar block not sprayed

with sulphur, C. vitis numbers increased ca. 29% over the same two-week time period.

While not as effective, a single application of sulphur in mid-season was reported to

reduce C. vitis populations by approximately 80% (Schreiner et al. 2014). Outbreaks of

C. vitis in Washington State have been attributed to decreased use of sulphur for powdery

mildew control (Prischmann and James 2005; Reinert 2006; James 2007). While reliance

on sulphur sprays in the past for the control of fungal pathogens may also have provided

control of C. vitis, high application rates can be detrimental to predacious mite

populations (McMurtry et al. 1970; James 2007).

Predacious phytoseiid mites are known to provide effective control of eriophyid mites

in the absence of insecticide sprays that are detrimental to their survival (James and

Whitney 1993; Bernard et al. 2005). In the absence of sulphur sprays, conditions are

right for C. vitis outbreaks following sprays that are harmful to mite predators. It was

apparent, for example, that the application of an undisclosed insecticide to part of a

cultivar block resulted in elevated numbers of C. vitis and significant damage to

developing shoots (Figure 4) the following spring. Differences in C. vitis numbers

between the heavily infested Shiraz block and the adjoining Merlot and Chardonnay

blocks (Table 1) possibly reflects differing pesticide applications the preceding summer.

Population levels of C. vitis would also vary depending on the frequency and timing of

sulphur applications in spring.

Damage Assessment. C. vitis may have an uneven distribution even within a

vineyard cultivar block. Assessment in spring 2010 of shoot growth in a section of a

Chardonnay block that had been sprayed with an insecticide the previous summer

showed severe stunting of shoots and distorted leaves (Figure 4). Differences in shoot

lengths for the rows heavily infested with C. vitis were significantly shorter than those

from the adjacent unsprayed rows (F1,76=63.3543, P<0.0001) (Table 3). Examination of

these damaged, or reduced, shoots showed significantly higher populations of C. vitis on

the stems (F1,76=76.5591, P<0.0001) and basal leaf (F1,276=46.1787, P<0.0001) compared

to undamaged shoots from the same vineyard block (Table 3). Numbers of phytophagous

thrips were low, less than one per basal leaf or shoot, and were not found to differ

significantly between the damaged and undamaged shoots (Fi,76=2.1008, P=0.1513)

(Table 3); therefore, thrips are unlikely to have contributed to the damage. Injury was still

measureable at harvest in September, with the C. vitis-infested vines having fewer shoots

(F1,78=5.5237, P=0.0213), fewer grape clusters (F1,73=102.7369, P<0.0001), and shorter

cluster lengths (Fi,7s=18.3596, P<0.0001) (Table 3).

J. Entomol. Soc. Brit. Columbia 114, December 2017

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J. ENTOMOL. SOC. BRIT. COLUMBIA 114, DECEMBER 2017 11

It is worth noting that the damaged shoots had lower numbers of predator mites than

the undamaged shoots, but the difference was not found to be statistically significant

(F 1,76=3.6261, P=0.061). Previous work has shown that preservation of mite predators is

important for the management of C. vitis (James and Whitney 1993; Bernard et al. 2005;

Reinert 2006; James 2007). James et al. (2002) report that effective biological control of

eriophyid mites in Washington vineyards likely depends on a complex of natural enemies

in addition to species of predatory phytoseiid mites. Outbreaks of C. vitis in Australia and

Washington State have been attributed to several causes, including the use of broad-

spectrum, persistent insecticides that harm predators. James (2007) suggested that the

appearance of C. vifis in Washington State might prove advantageous for grapevine

biological control programs, as these mites provide an early season food source for

predacious mites before Tetranychid spider mite species appear later in the season.

Additional study is required to determine the mite predator complex in B.C. and to

establish their role in the sustainable management of C. vitis.

Figure 4. Developing grapevine shoot severely damaged by grape leaf rust mite,

Calepitrimerus vitis (Nalepa), feeding in early spring (left) compared with an undamaged

shoot (right) from the same cultivar block. Note the shortened internodes, brown and distorted

leaves and flower buds, and scarring of the stem on the damaged shoot.

In conclusion, our research has demonstrated that reduced spring growth of shoots

having deformed leaves that had often been attributed to herbicide or winter damage,

post-harvest water stress, thrips, and other maladies is in many cases due to feeding

damage from C. vitis. Presence of this eriophyid mite under bud scales and on developing

shoots has been linked to short shoot syndrome of grapevines, an economically important

12 J. Entomol. Soc. Brit. Columbia 114, December 2017

syndrome of grapes in the Pacific Northwest of the United States (Walton et al. 2007;

Schreiner et al. 2014). We determined that C. vitis were distributed widely in the

southern interior of B.C. and observed high numbers feeding on buds and tender shoots

in spring (Figure 3) that resulted in distortion and stunting of shoots (Figure 4; Table 3).

Although growth often improves during the summer, yields will be reduced significantly,

as we have shown.

The recommendation to apply sulphur at the woolly bud stage based on bronzing of

leaves in late summer is supported by our observations. Evidence for potentially

damaging populations of these mites based on bronzing of leaves in late summer is an

indication that control measures should be applied the following spring. Although there is

no reported damage to grapes from C. vitis feeding during the summer, they can be

controlled at this time with foliar sprays of miticides (Anonymous 1968; Walton ef al.

2007; Siquera et al. 2016).

With the documented arrival of C. vitis to the southern interior of B.C., it is important

that growers learn to recognize early signs of severe C. vitis infestations (russeting of

leaves) that indicate the need for timely and appropriate sulphur sprays during early bud

development the following spring. Attempts should also be made to preserve mite

predators by avoiding the use of harmful sprays.

ACKNOWLEDGEMENTS

We extend our gratitude to Dr. F. Beaulieu, AAFC Ottawa, and Dr. J. Amrine, West

Virginia University, for identification of C. vitis. Thank you to M. Weis of AAFC-

Summerland RDC for photographs, P. Bowen and B. Estergaard, AAFC-Summerland

RDC for use of the map, M. Watson of Constellation Brands International, and other

producers for allowing access to their vineyard sites.

J. ENTOMOL. SOc. BRIT. COLUMBIA 114, DECEMBER 2017

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Anonymous. 1968. Pests of the Grapevine. Insect Pest Bulletin 44, N.S.W. Department of Agriculture,

Entomology Branch. 4" edition. 7pps.

Anonymous. 2005. Characteristics of rust mites. Viti-Notes, Cooperative Centre for Viticulture.

www.crcev.com.an/viticare/vitinotes/

Barnes, M. M. 1970. Grape Pest in Southern California. Circular 553, California Agriculture

Experimental Station Extension Service, Division of Agricultural Science, University of California.

Bernard, M. B., P. A. Hore, and A. A. Hoffmann. 2005. Eriophyid mite damage in Vitis vinifera

(grapevine) in Australia: Calepitrimerus vitis and Colomerus vitis (Acari: Eriophyidae) as the

common cause of the widespread ‘Restricted Spring Growth’ syndrome. Experimental Applied

Acarology 35:83—-109.

British Columbia Wine Institute. March 2015. Celebrate the Wines of British Columbia. BCWI media kit

2015.

Duffner, K., G. Schruft, and R. Guggenheim. 2001. Passive dispersal of the grape rust mite

Calepitrimerus vitis Nalepa 1905 (Acari, Eriophyoidea) in vineyards. J. Pest Science 74:1—6.

James, D. G. 2007. Grape rust mites: New enemies (but ultimately friends) invade Washington

vineyards. Vine web — WSU Viticulture and Enology, WSU-Prosser. Available at

winegrapes.wsu.edu/archivedTOM/topic4-07.html.

James, D. G., and J. Whitney. 1993. Mite populations on grapevines in south-eastern Australia:

Implications for biological control of grapevine mites (Acarina: Tenuipalpidae, Eriophyidae).

Experimental Applied Acarology 17:259-270. |

James, D. G., T.S. Price, L. C. Wright, and J. Perez. 2002. Abundance and phenology of mites, |

leafhoppers, and thrips on pesticide-treated and untreated wine grapes in South Central Washington,

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Commercial Growers. BC Wine Grape Council and BC Ministry of Agriculture and Lands.

www.bewgce.org/

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and Similkameen valleys, British Columbia. J. Entomol. Soc. Brit Columbia. 72 (Dec.):9-14.

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Characters and the impact of spray practices. In, Ecology of tetranychid mites and their natural

enemies: A review. Hilgardia 40(11):33 1-390.

Prischmann, D. A., and D. G. James. 2005. New mite records (Acari: Eriophyidae, Tetranychidae) from

grapevines in Oregon and Washington State. Internat. J. Acarol. 31(3):289-291.

Reinert, B. 2006. New mites detected in Washington vineyards. Good Fruit Grower. 57(5):37.

SAS Institute Inc. 2013 JMP Version 10, SAS Institute, Cary, North Carolina.

Schreiner, R. P., P. A. Skinkis, and A. J. Dreves. 2014. A rapid method to assess grape rust mites on

leaves and observations from case studies in western Oregon vineyards. Horticultural Technology

24(1):38-47. http://hdl handle net/1957/47976.

Siqueira, P. R. E., M. Botton, P. R. B. Siqueira, G. S. Peres, and L. L. Soares. 2016. Effect of acaricides

on Calepitrimerus vitis (Nalepa, 1905) (Acari: Eriophyidae) and on the production of vineyards.

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Walton, V. M., A. J. Dreves, D. H. Gent, D. G. James, R. R. Martin, U. Chambers, and P. A. Skinkis.

2007. Relationship between rust mites Calepitrimerus vitis (Nalepa), bud mites Colomerus vitis

(Pagenstecher) (Acari: Eriophyidae) and short shoot syndrome in Oregon vineyards. International J.

Acarol. Vol. 33 No. 4:307—318.

Winkler, A. J., J. A. Cook, W. M. Kliewer, and L.A. Lider. 1972. Grape pests. pps 503-555. Jn, General

Viticulture, Chapter 19. University of California Press.

Zar, J. H. 2010. Biostatistical Analysis, 5‘* Ed. Prentice Hall, Upper Saddle River, NJ.

J. ENTOMOL. SOC. BRIT. COLUMBIA 114, DECEMBER 2017 iS

Insect taxa named for the Rev. John H. Keen, early

naturalist on the Queen Charlotte Islands and at

Metlakatla, British Columbia

SPENCER G. SEALY!

ABSTRACT

The Reverend John Henry Keen (1851-1950) spent nearly 20 years serving Anglican

missions in British Columbia, at Masset on the Queen Charlotte Islands/Haida Gwaii

in the 1890s, and on the adjacent mainland at Metlakatla, during the summer of 1890

and for several years in the early 1900s. Despite leading the busy life of a clergyman,

Keen assembled extensive collections of natural history specimens, particularly of

insects and mammals. He was spurred on by the likelihood that many specimens

would represent species new to science, predictions that were later borne out. Keen

initially sent specimens to the Natural History Museum in London, but later sent most

of them to Dr. James Fletcher, Dominion Entomologist, in Ottawa, who forwarded

many specimens to specialists in the United States and France for identification. Keen

was among the first collectors of natural history specimens on the north coast of

British Columbia and, in recognition of his contributions, eight insect taxa were

named after him, based on the type specimens he collected in this region.

Key words: British Columbia; Masset; Metlakatla; Queen Charlotte Islands/Haida

Gwaii; keeni; type specimens

Among the members of the clergy whose early entomological contributions in British

Columbia were chronicled by Riegert (1991) in Entomologists of British Columbia was

the Rev. John Henry Keen (1851-1950). Keen was an Anglican missionary who served at

missions on the Queen Charlotte Islands/Haida Gwaii at Massett (hereafter the modern

spelling of Masset is followed; 54.0115°N, 132.1472°W) in the 1890s, and on the

mainland coast at Metlakatla (54.3373°N, 130,4447°W) for several years in the early

1900s. Keen arrived in British Columbia in the early summer of 1890 and worked for

several weeks on the mainland (Sealy 2016a, b), where he wasted no time in assembling

the first collection of beetles from this region (Keen 1891). Keen finally arrived in

Masset in mid-September 1890 and began his clerical duties, but also continued what

would become a productive period in the history of natural history in this region (Sealy

2015, 2016a, 2017).

Prior to leaving England, Keen prepared for the upcoming challenges of serving a

people with which he was unfamiliar, but he was also filled with anticipation of working

in a region whose natural history was relatively unexplored. Keen’s letters to Albert K. L.

G. Ginther (1830-1914), Keeper of Zoology at the British Museum (Natural History),

London, from 1875 to 1895 (Gunther 1930), now the Natural History Museum (NHM),

revealed his anticipation of reaching the Queen Charlotte Islands and the discoveries he

felt sure would follow. Shortly before arriving at Masset, Keen wrote to Giinther, “As |

remember you said that almost anything from [the Queen Charlotte Islands] would be of

interest, I shall hope to send you a good deal from time to time” (Keen 1890). Upon his

arrival, he re-iterated his awareness of the value of specimens from this new locality,

again in a letter to Giinther: “The productions of the island, one would think, ought to be

of considerable interest as it is separated from the mainland by a channel [Hecate Strait]

about 60 miles wide” (Keen 1890). Keen’s work on the natural history of the Queen

Charlotte Islands had begun, and recognition of the results soon followed.

| Department of Biological Sciences, University of Manitoba, Winnipeg, Manitoba R3T 2N2, Canada;

spencer.sealy@umanitoba.ca

16 J. Entomol. Soc. Brit. Columbia 114, December 2017

In the years that followed, Keen assembled extensive collections of insects and other

animals, particularly of mammals, and some plants. Although Keen collected the

specimens and made notes on the behaviour and habitat of many of the species (Sealy

2015, 2017), he had to rely on specialists to identify them. Specimens collected at

Metlakatla and during the first year or so of service at Masset were sent to Giinther at the

British Museum (Sealy 2015, 2016c). By late-1892, impatient with delays in receiving

determinations from busy curators in England and the irregular mail service, Keen began

sending most specimens of insects and a few other invertebrates and mammals to the

Central Experimental Farm in Ottawa. There, entomologist James Fletcher (1852-1908)

identified some of the insects, but sent the other specimens to specialists in France and

the United States. Fletcher, a highly acclaimed entomologist (Gibson and Groh 1909),

recognized the seriousness and energy that Keen brought to his observations and

collections and, thus, both contributed importantly to knowledge of the natural history in

a region that was almost completely unexplored biologically at the time.

Keen’s contributions to knowledge of the insect fauna of the coastal region of central

British Columbia took three forms: (1) published lists of beetles identified by specialists,

frequently annotated with notes on behaviour and seasonal habitats (Sealy 2015); (2)

specimens, mostly of beetles, incorporated into catalogues, taxonomic revisions, and

distributional works published by other entomologists and in museum reports (e.g.,

Harrington 1894, Fannin 1898, Kavanaugh 1992); and (3) eight taxa, including two taxa ©

that Hatch (1957b) considered to be nomen nuda, described from specimens collected by

Keen. These specimens were catalogued in the National Collection of Insects (Ottawa),

the Natural History Museum (London, UK), Provincial Museum of British Columbia

(now Royal British Columbia Museum, Victoria), and Biological Survey of the U.S.

Department of Agriculture (U.S. National Museum, Washington).

Insect taxa named for John H. Keen. The dates of collection of Keen’s insect

specimens, types and otherwise, are generally not known except for the year or the

season (“February”, “summer”). It is known with certainty, however, that specimens

were collected on the Queen Charlotte Islands between mid-September 1890 and

late-1898, during Keen’s residency at Masset (Sealy 2016a). Specimens were collected at

Metlakatla in June and July 1890, while Keen waited for the steamer to transport him to

Masset, and again after settling at Metlakatla in late summer of 1899 (Sealy 2016a).

Specimens were collected year-round on the Queen Charlotte Islands, within “a circle of

five miles’ radius from Massett” (Keen 1895), and at Metlakatla, mainly at settlements

along the Nass River (Sealy 2016c). Unlike descriptions of new taxa of mammals from

the Queen Charlotte Islands that were based on Keen’s specimens, in which each was

accompanied by a type specimen and institutional registration number (Sealy 2015,

2017), catalogue numbers of type specimens of insects were generally not available. The

new taxa listed below are presented in chronological order of the date of description,

with the exception of two species considered by Hatch (1957b) to be nomen nuda, which

are presented at the bottom of the list.

Pezomachus keenii (Harrington 1894) (Hymenoptera: Ichneumonidae). Among a

collection of several undescribed ichneumonids assembled by the Rev. G. W. Taylor near

Victoria, British Columbia, and entrusted to W. H. Harrington, were two new species of

wasps collected by Keen near Masset. Regarding the first species, Cremnodes

canadensis, Harrington (1894) noted that it was “[d]escribed from one 9 specimen from

Queen Charlotte Islands, sent by the Rev. J. H. Keen to Mr. Fletcher. A very interesting

wingless species, with rufous head and abdomen, and testaceous thorax and legs ...” This

species now resides in the genus Polyaulon Foerster. It took only seven lines for

Harrington (1894) to describe the second species, this one placed in the genus

Pezomachus, based on four females collected at Masset by Keen, “... after whom I have

much pleasure in naming the species, as a recognition of his efforts to advance our

knowledge of the insect fauna of this distant portion of the Dominion.” Pezozmachus was

synonymized with the genus Gelis by Viereck (1914).

J. ENTOMOL. SOC. BRIT. COLUMBIA 114, DECEMBER 2017 Y

[George W. Taylor (1854-1912) was another among several early clergymen who

collected insects in British Columbia. He became a sought-after expert on the

Geometridae and exchanged specimens with collectors and identified moths for others

(Riegert 1991). William H. Harrington (1852—1918) was one of the founders of the

Ottawa Field-Naturalists Club, a still-active organization that publishes the Canadian

Field-Naturalist. His entomological contributions focused on systematics and economic

entomology, particularly of the Hymenoptera and Coleoptera (Gibson 1918). ]

Platyceropsis keeni (Casey, 1895) (Coleoptera: Lucanidae). This is the first of three

species of beetle that Capt. Thomas L. Casey (1857-1925) named in honour of Keen,

based on a single female specimen (Benesh 1946). Casey noted (1895) “This interesting

species was discovered by Rev. J. H. Keen [at Masset], and the original specimen kindly

given me for description by Mr. [H. F.] Wickham, with permission of Mr. James Fletcher,

of Ottawa. It has recently been taken in abundance.” This species had been collected only

on the Queen Charlotte Islands (Wickham 1899) and, with specimens of Haida keeni (see

below), were part of a collection of 141 species of beetle that Keen presented to the

British Columbia Provincial Museum in Victoria (now Royal British Columbia

Museum). The list of Keen’s specimens was included in Fannin’s (1898) preliminary

catalogue of collections deposited in the Museum. This species belongs to the genus

Platyceropsis (Benesh 1946).

[Henry Frederick Wickham (1966-1933), professor of entomology at the State

University of Iowa (now Iowa State University), was a specialist in the Coleoptera

(Anonymous 1934). He described many new species of beetle, including fossilized

species, and was among several specialists to whom James Fletcher forwarded Keen’s

specimens for identification and whom Keen acknowledged (Keen 1895). Wickham’s

field experience extended to British Columbia, where he collected insects for one month

near Victoria on Vancouver Island in 1889 (Wickham 1890) and, in 1891, collected them

in Alaska and the adjacent portions of British Columbia (Wickham 1893). ]

Oxypsylla keeni (Baker, 1896) (Siphonaptera: Ceratophyllidae). Keen collected

fleas secondarily as they escaped from the pelage of Keen’s Mouse (Peromyscus keeni (=

Sitomys keeni Rhoads) and from mouse nests, while recording notes on this species’

behaviour (Keen 1896; also see Sealy 2015). From “several [male and female] specimens

taken on Sitomys keeni at Masset... in August of 1895, by Rev. J. H. Keen’, Baker

(1896) described a new species, Pulex keeni, and named it in honour of Keen. He also

acknowledged his indebtedness to James Fletcher “... for the opportunity of examining

this very interesting and well-marked form.” Baker (1904) supplemented the original

description of Pulex keeni with figures and re-assigned it to the genus Ceratophyllus.

Jordan (1933) re-assigned Ceratophyllus to the genus Opisodasys. A lectotype male

was designated by Smit and Wright (1978), and the species was placed in the subgenus,

Oxypsylla, by Smit (1983), where it resides today. The preferred host of Oxypsylla keeni

is the deermouse genus Peromyscus in south-central British Columbia, on Vancouver

Island and other coastal islands, including Haida Gwaii, and north along the panhandle of

Alaska (Holland 1985), as well as deermice throughout the northwest United States,

including western Montana and northern Nevada and Utah (Lewis 2008).

Baker benefited further from Keen’s collecting skills, with his description of another

new species of flea taken from a deermouse nest at Masset in 1898. Two females

provided the basis for Baker’s (1898) description of Typhlopsylla charlottensis, based

primarily on account of its reduced eyes. Further study prompted Baker (1904) to expand

“the meagre original description” of this new species, and he re-assigned it to the genus

Ceratophyllus. Rothschild (1915) later designated the genus as Catallagia, thus,

Catallagia charlottensis.

After a furlough in England, Keen resumed his duties in mid-July 1899, this time at

the mission at Metlakatla, where he continued to collect specimens, including fleas.

Holland (1985) included two species that included specimens collected by Keen:

Hystrichopsylla occidentalis occidentalis Holland (from Norway Rat [Rattus norvegicus |

18 J. Entomol. Soc. Brit. Columbia 114, December 2017

at Metlakatla), and Monopsyllus ciliatus protinus Jordan (from Red Squirrel

[Tamiasciurus hudsonicus] at Metlakatla and Tamiasciurus sp. at Inverness).

Haida keeni Keen 1897; Brown 1944 (Coleoptera: Staphylinidae). The history

surrounding the naming of Haida keeni may be unique. Among Keen’s beetle specimens

sent by Fletcher to Mons. Albert Fauvel, of Caen, France, a specialist on the Coleoptera

(e.g., Fauvel 1889), was a specimen collected at Masset on 18 October 1893 (Figure 1;

also figured by Keen (1897) and frontispiece in Campbell 1978: facing p. 1) that was

recognized as a new taxon. Fauvel suggested it be named Haida keeni, the genus

suggested in honour of the Haida people, the traditional inhabitants of Haida Gwaii with

whom Keen was working, and the species after Keen, the collector. But Fauvel never

published a formal description of Haida keeni, although he is sometimes named as the

author of the genus (e.g., Fannin 1898, but see Armett and Thomas 2000). As Hatch

(1957a) noted, this resulted in Keen’s (1897) “interesting but taxonomically most

inadequate remarks constitut[ing] the original description of both genus and species, and

resulted in Keen being in the anomalous position of naming a species after himself!”

Forty-five years later, Haida keeni was accurately described by Brown (1944), based on

three additional specimens collected at Masset in 1893 and catalogued in the National

Insect Collection in Ottawa. But Keen, not Brown, is still recognized as the species’

author (e.g., Hatch 1957b, Campbell 1978, Bosquet et al. 2013; also see Hatch 1957a).

Keen (1895) noted this species is “Not common. Found in moss at roots of trees, in

December.” He did not collect this species at Metlakatla (Keen 1905), although

specimens have been taken subsequently from the mainland, from southeast Alaska to

southwestern British Columbia (Campbell 1978).

Figure 1. Haida keeni registered in the Canadian National Collection of Insects, Ottawa,

Ontario, collected by the Rev. J. H. Keen on 18 October 1893 near Masset, Queen Charlotte

Islands, British Columbia. Photo credit: Serge Laplante, courtesy of the Canadian National

Collection of Insects, Arachnids and Nematodes, Agriculture and Agri-Food Canada, Ottawa.

Atheta (Lamiota) keeni Casey 1910 (Coleoptera: Staphylinidae). Describing this

species (type, USNM 38480 insects, collected at Metlakatla), Casey (1910) noted, “This

strikingly distinct species is dedicated with pleasure to Rev. J. H. Keen, who has made

J. ENTOMOL. SOC. BRIT. COLUMBIA 114, DECEMBER 2017 19

many interesting discoveries among the small clavicorn Coleoptera of the northern coast

of British Columbia.” Considered a valid species, Atheta keeni was designated the type

species of the subgenus Lamiota Casey, although Gusarov (2003), who supplemented

Casey’s description of Atheta keeni with illustrations of body parts, cautioned that the

“Subgeneric assignment of At. keeni and the status of the name Lamiota require further

study.” Atheta keeni is known from Alaska, British Columbia and Oregon (Gusarov

2003).

Gyrophaena keeni Casey, 1911 (Coleoptera: Staphylinidae). The description of the

third species of beetle named for Keen by Casey (1911) was accompanied only by a brief

acknowledgement and statement of the type locality: “British Columbia (Metlakatla), —

Keen.” This species, described from a male specimen, and others in the genus

Gyrophaena proposed by Casey have been confirmed as valid. Seevers (1951) grouped

five closely related species of Gyrophaena in a “Keeni group” (also see Stace Smith

1957), which is composed of fungus-feeding beetles, obligatory inhabitants of the fungi

during the larval and adult stages.

Bryobiotos keeni Fauvel (Coleoptera: Staphylinidae). Hatch (1957b) considered

this species to be a nomen nudum, and thus not described and not valid. Keen (1895)

noted in the entry for this species in his list of beetles from Masset that it was

“Occasional in June under stones on sandy beach, between tide marks. Larvae in same

place.”

Anthobium keeni (Fauvel) (Coleoptera: Staphylinidae). Keen (1895) listed this

species in the genus Lithrimaeum, noting “Several in rotten sea-weed, in June”, but

Hatch (1957b) considered it to be a nomen nudum. Among the locations given by Hatch

(1957b) in the description of his new species, Anthobium sinuosum, was Metlakatla,

where one of the female paratypes was collected, probably by Keen. It is possible that

keeni was placed with the new species.

CONCLUSIONS

Keen’s collecting career in coastal British Columbia spanned more than 20 years. His

last specimen apparently was a sap-feeding beetle (Fabogethes nigrescens (Stephens))

taken at Metlakatla in 1915 and catalogued in the British Museum (Easton 1955, also see

Hatch 1957b). Osgood (1901), in the first treatise of the fauna and flora of the Queen

Charlotte Islands, extolled Keen’s dedication and far-reaching contributions to natural

history, acknowledging that the little that was known of the vertebrate fauna of the

islands “was entirely due to the zeal of Rev. J. H. Keen ...” Accolades from

entomologists followed. Baker (1904) stated “All of our records [of fleas] for the Queen

Charlotte Islands are due to this gentleman, and his contributions have been most

important ones.” Riegert (1991), in a brief sketch of Keen’s life and accomplishments,

noted that “Our knowledge of the original beetle fauna of the northwest B.C. coast is due

primarily to the painstaking and energetic collecting of this remarkable clergyman.” This

sentiment was echoed by Kavanaugh (1992) who acknowledged that “Development of

our present knowledge of the carabid beetle fauna of the Queen Charlotte Islands began

with the Reverend J. H. Keen, Anglican missionary to the Haida people ...” In a memoir

published following Keen’s death in England in 1950 at the age of 98, Hatch (1957a)

acknowledged that entomologists were aware of Keen’s “short series of papers” on the

beetles of the Queen Charlotte Islands, but he lamented that there was a “... complete

dearth of published information about Rev. Keen.” Hearne (1997) extended Hatch’s

(1957a) brief biography of Keen in a tribute to the “forgotten naturalist” of the Queen

Charlotte Islands. Recently, photographs of Rev. Keen have been uncovered (Sealy

2016a).

20 J. Entomol. Soc. Brit. Columbia 114, December 2017

ACKNOWLEDGEMENTS

Serge Laplante provided the photograph of the type specimen of Haida keeni in the

Canadian National Collection of Insects, Ottawa, Ontario. Daisy Cunynghame and

Lorraine Portch (Natural History Museum & Archives, London, UK) provided copies of

Keen’s correspondence archived in that Institution. Terry D. Galloway (Department of

Entomology, University of Manitoba) pointed out several references and commented on

an early draft of the manuscript. The reviewers, Robert J. Cannings and Jan

Klimaszewski, commented constructively on the manuscript.

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22 J. Entomol. Soc. Brit. Columbia 114, December 2017

Western balsam bark beetle, Dryocoetes confusus Swaine

(Coleoptera: Curculionidae: Scolytinae), in situ

development and seasonal flight periodicity in southern

British Columbia

L. E. MACLAUCHLAN'! and J. E. BROOKS?

ABSTRACT

In situ development and seasonal flight periodicity of the western balsam bark beetle,

Dryocoetes confusus Swaine, was observed in subalpine fir, Abies lasiocarpa (Hook)

Nutt. stands in southern British Columbia for three years between 1998 and 2002. This

study shows developmental differences of western balsam bark beetle in downed and

in standing, live subalpine fir. Larval development was slower in the downed trees.

Recorded daily minimum phloem temperatures were significantly lower for downed

trees than for standing trees during periods of beetle development and flight. There

were no significant differences in the recorded daily maximum phloem temperatures

between standing and downed trees until late summer, when downed trees saw cooler

daily maximum phloem temperatures. This cooler host habitat would provide fewer

degree days for insect development. Three distinct larval instars were identified by

head capsule measurement. There were two flights per season, the first and major

flight occurring from late June to late July, and the other smaller flight occurring in

late August. A combination of minimum daily phloem temperatures reaching 5° C and

maximum daily phloem temperatures approaching 20° C appeared to trigger the onset

of beetle flight, with flight initiated earlier in the season at lower elevations.

Key words: development, instar determination, subcortical temperature

INTRODUCTION

The western balsam bark beetle, Dryocoetes confusus Swaine (Coleoptera:

Curculionidae: Scolytinae), is the most destructive insect pest of mature and over-mature

subalpine fir, Abies lasiocarpa (Hook.) Nutt. in British Columbia (B.C.) (Garbutt 1992).

Western balsam bark beetle is found throughout the range of subalpine fir and is the

dominant successional force in high-elevation ecosystems of the Engelmann Spruce—

Subalpine Fir zone (ESSF) (Stock et al. 1994; Maclauchlan 2016), which include dry,

moist, and wet subzones (Meidinger and Pojar 1991). Tree mortality from this beetle is

first noticed in stands approaching 70—90 years of age (Maclauchlan 2001) and, as stands

age, the aggregated pattern of attack by western balsam bark beetle describes the small-

scale gap dynamic process, which over time releases the next generation of subalpine fir

(Stock et al. 1994). Subalpine fir is susceptible to a wide variety of other disturbance

agents, including two-year-cycle budworm, Choristoneura biennis Freeman, various root

and butt rots, stem rots, animal damage and windthrow (Alexander 1987; Unger 1995;

Parish and Antos 2002). Fire is relatively rare in the wetter ESSF subzones (Anon. 1995),

often seen as small, localized events. Despite these other disturbances, western balsam

bark beetle is one of the primary drivers of succession in both the ESSF and other

subalpine fir-dominated ecosystems throughout B.C. (Maclauchlan eft al. 2003;

Maclauchlan 2016).

Western balsam bark beetle selectively kills small groups of subalpine fir at a

relatively low, but constant, level each year in infested stands (Stock et a/. 1994; Unger

| B.C. Ministry of Forests, Lands and Natural Resource Operations and Rural Development, 441

Columbia Street, Kamloops, B.C. V2C 2T3; Lorraine.maclauchlan@gov.bce.ca

2 Forest Health Management, 466 Central Avenue, Gibsons, B.C. VON 1V1

J. ENTOMOL. Soc. BRIT. COLUMBIA 114, DECEMBER 2017 23

and Stewart 1992; McMillin et al. 2003). Recently killed, red trees occur in groups of

two or more, often spread over hundreds of hectares (Stock 1991). Attack rates of

western balsam bark beetle in southern B.C. can range from 0.7 to 1.6% of subalpine fir

annually in a stand, depending upon ecosystem (Maclauchlan 2016). The beetle attacks

large-diameter, standing live trees and downed subalpine fir. Western balsam bark beetle

consistently attacks trees from the largest-diameter classes in each stand, but the mean

diameter of attacked trees between sites may vary significantly (ranging from 10 cm

diameter at breast height to over 50 cm), indicating that factors other than diameter

contribute to the susceptibility of subalpine fir to western balsam bark beetle (Bleiker ef

al. 2003). Susceptibility has been associated with tree diameter, age, recent radial growth,

and induced resinosis (Bleiker et a/. 2003). Although cumulative mortality can reach

significant levels in chronically infested stands (Garbutt and Stewart 1991; Maclauchlan

2016), western balsam bark beetle may be less aggressive or exhibit unique attack

dynamics compared to other tree-killing bark beetles at epidemic levels. Beetle

populations persist within a stand for many years until most of the mature subalpine firs

are killed (Garbutt 1992; McMillin et al. 2003). This selective and patchy distribution of

mortality suggests that western balsam bark beetle may be limited by the abundance and

distribution of susceptible hosts, as well as the harsh environment in which they live.

Many researchers (Hansen 1996; Gibson et al. 1997; Negron and Popp 2009; Stock et

al. 2013) have concentrated primarily on the flight periodicity and insect activity within

subalpine fir stands. A paucity of work has been done on life-stage development of the

western balsam bark beetle primarily due to the remote nature of most subalpine fir

forests. In B.C., adults generally emerge in late June and fly until late July, locating

suitable host trees through kairomones and primary attraction, at which point the males

initiate construction of nuptial chambers beneath the bark (Bright 1976; Garbutt 1992;

Stock et al. 2013). The species is polygamous, with males often attracting three or more

females in a nuptial chamber. Females mate, then lay eggs in brood galleries that radiate

out from the nuptial chamber, and toward the end of August will construct feeding and

hibernation niches (Bright 1976) for the winter. Mature females may resume laying eggs

the following year within hosts that have adequate phloem resource, whereas females

within trees that are fully occupied with brood may emerge mid-summer to locate new

hosts (Bright 1976). In addition to the main attack flight, comprised of males and

females, an additional smaller flight, largely comprised of parent females (Hansen 1996;

Gibson et al. 1997; Stock et al. 2013), has been observed later in the summer. In this late-

season second flight, the females join existing gallery systems and often create

hibernation niches.

Mathers (1931) first described the life cycle of the western balsam bark beetle in

B.C., demonstrating that it completed its life cycle within two years. Bright (1963)

subsequently speculated that the insect might be capable of completing its life cycle in

only one year in the western and southwestern United States. Whether the brood is

capable of developing to the adult stage in one year has not been shown. Insects have

different strategies to cope with fluctuating weather conditions and phases of growth and

dormancy: some undergo hormonally controlled diapause (obligatory diapause) that

prevents insect development even when environmental conditions are good (Gilbert

1990), while others only undergo diapause when induced environmentally (facultative

diapause). The spruce beetle, Dendroctonus rufipennis Kirby, is known to have a life

cycle that can vary in duration, from one year up to three years, depending on climatic

conditions and suitability of host material available (Knight 1961; Schmid and Frye

1977; Hansen et al. 2001; Bentz et al. 2010). Johansson et al. (1994) describe a flexible

generation time for Dryocoetes autographus (Ratz.), a circumpolar species, that has

expanded its range in Norway north, with the establishment of its host species, spruce.

Swift and Ran (2013) noted that climate change may have a pronounced effect on high-

elevation forests and associated insects. Therefore, a greater understanding of the

developmental requirements for western balsam bark beetle is needed.

24 J. Entomol. Soc. Brit. Columbia 114, December 2017

This study focuses on western balsam bark beetle life-stage development in standing

and down subalpine firs, and flight periodicity over a range of elevations in southern

B.C. We also studied the relationship of temperature to western balsam bark beetle

development in standing and down host material. These trials were conducted in 1998,

1999, and 2002 at various field sites in southern B.C.

MATERIALS AND METHODS

The developmental biology of western balsam bark beetle was investigated in the

field by flight periodicity trapping, sampling of in situ life stages, and weather

monitoring. These studies focused on the two very different stages in the beetles’ life

history: 1) emergence and flight dispersal and, 2) brood production and maturation

within the host tree. On-site weather monitoring was used to determine critical threshold

subcortical (phloem) temperatures for flight and development.

Seven field sites were selected throughout the southern interior of B.C. in subalpine

fir ecosystems with active populations of western balsam bark beetle. Two sites, Cherry

Ridge and Sun Peaks, were used in all three studies, while the other sites were used for

monitoring flight dispersal timing only (Table 1).

One micro-logger—based climate station (Campbell Scientific Inc.) was set up at each

of the Sun Peaks and Cherry Ridge sites to monitor ambient, duff and phloem

temperature of western balsam bark beetle-attacked trees. The Sun Peaks and Cherry -

Ridge climate stations were located on the north aspects of standing trees within close

proximity (+10 m) of other attacked trees, where flight dispersal monitoring and detailed

life-stage studies were conducted. The Sun Peaks climate station was set up on June 18,

1998, and again on June 15, 1999. The climate station was installed next to a standing

tree, and thermocouples were inserted three meters up the bole in the phloem of five

nearby attacked subalpine fir. Thermocouples were inserted into western balsam bark

beetle entrance holes. Thermocouples were also placed in the phloem on downed trees.

Ambient air temperature was recorded at the relative humidity sensor on the climate

station. Another climate station was installed on June 16, 1999, at Cherry Ridge, as per

Sun Peaks. Numerous variables were recorded, but only date and phloem temperature

were used for analyses in this study. On August 5, 1999, the climate station at Cherry

Ridge malfunctioned, and no further weather data were recorded.

In 1998, three 8-funnel (8 plastic funnels aligned vertically over each other) Lindgren

multiple-funnel traps (Lindgren 1983) were erected between June 18 and August 31 at

Sun Peaks and monitored regularly for western balsam bark beetle flight activity. Traps

were placed along an elevation gradient (1,450 m; 1,650 m; 1,850 m), and all traps were

positioned just inside the stand edge. Each trap was hung on an aluminum pole, with the

top of the trap approximately two meters above the ground. Traps were baited with the

commercially available (4)-exo-brevicomin (release rate 0.4mg/24 h) bait for western

balsam bark beetle (supplied by Phero Tech Inc., Delta, B.C., Canada, and now available

through Distributions Solida Inc., Scotts Miracle-Gro Company).

In 1999, trapping trials were established at Sun Peaks and Cherry Ridge sites to

follow the flight timing and activity of western balsam bark beetle. Four traps were hung

at Sun Peaks over an elevational range at 100-150 m intervals (1,450 m—1,850 m), and

two traps were hung at 1,650 m at Cherry Ridge. Weekly trap catches were collected

from June 16 to October 9 at Sun Peaks. At Cherry Ridge, collections were made at

irregular intervals from June 15 to October 30.

In 2002, a more comprehensive trapping trial was conducted, using nine sites (three

traps per site) in six geographic locations (Table 1) within four ESSF subzones. Lindgren

funnel traps were set up in a triangular formation, with the traps being approximately 20

metres apart. Traps were established from May 29 to June 21, with regular collections

beginning June 19 until September 27. During the peak flight period, trap collections

were made more frequently, up to three times per week, until the peak flight was over.

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All insects collected from the trapping trials were stored in zip-lock bags, labeled, and

frozen until processed in the laboratory, where each sample collection was then counted

and the western balsam bark beetles were sexed.

Trap catch results were compared to daily weather patterns at sites with climate

stations. The maximum and minimum phloem temperatures were plotted against trap

catch to interpret the relationship between insect flight and subcortical temperature.

The literature describes western balsam bark beetle as having a two-year life cycle

(Mathers 1931; Bright 1976). To capture all life stages (from initial attack through newly

emerged adults), sample trees meeting specific criteria were selected in 1999 at the Sun

Peaks and Cherry Ridge study sites. External signs and symptoms, such as foliage fade,

and presence of entrance holes and frass on the bole, were used to select candidate trees

and ascertain year of attack. Western balsam bark beetle tree baits ((+)-exo-brevicomin;

release rate 0.4mg/24 h) were attached to two standing live and two freshly felled

subalpine firs in early June 1999 to induce western balsam bark beetle attack. Downed

trees were felled into the stand, where they were well shaded, and no limbs were

removed. Both standing and down (natural blowdown) subalpine fir attacked by western

balsam bark beetle in 1997 or 1998 were selected for sampling in 1999 to determine if

there were obvious developmental differences between these two host scenarios. Six

trees at Sun Peaks and three trees at Cherry Ridge were suitable for sampling (Table 2).

All sample trees were in close proximity (+10 m) to the climate stations. Sampling took

place at weekly intervals between June 15 and September 20, 1999. A rigorous sampling —

procedure was followed at each sampling date to help interpret progression of attack and

tree symptoms. For each sample tree, numerous foliar attributes and bole symptoms were

recorded, but only the ones used in the analysis are described. In some cases, particularly

in the trees that were attacked in 1997, a comment was made at each dissection as to the

abundance of exit holes. A ladder was used to access western balsam bark beetle attack

found higher on the bole, up to 3.5 meters. A 20-cm x 20-cm template was centered over

an entrance hole, and the bark was carefully removed to expose the gallery system.

Gallery systems (Figure 1) were described to help elucidate the stage in the life cycle and

productivity. All western balsam bark beetle life stages present (eggs, larvae, pupae, and

parent and teneral adults) were collected and placed in vials of 70% ethanol for future

processing in the laboratory.

Head capsule measurement is a commonly used method to determine the instar of

immature insects (Bleiker and Régniére 2014). The head capsules of all western balsam

bark beetle larvae collected in the field were measured using a dissecting microscope.

Every measurement was taken at 4.5 X magnification, which yielded a micrometre

measurement of 0.022 mm. Measurements were taken across the widest portion of the

sclerotized head capsule. All head capsule measurements were sorted in ascending order

and plotted to display frequency distribution. The lowest frequency class between peaks

on a histogram can be used as the cut-points for each instar and are often visually

determined (Logan et al. 1998; Bleiker and Régniere 2014). From these frequency

distributions, delineation of instars was determined by visually identifying cut-points.

Each instar was assigned a head capsule size range. All larvae collected were then given

an instar designation and these data were then compared to field temperatures and date of

field sampling. These comparisons provided the interpretations of life-stage occurrence

and duration in standing and down trees.

RESULTS

Very few beetles were caught in 1998 at the Sun Peaks site (170 beetles over 75

days), with the highest trap catches occurring July 6 and July 10. In 1999, at the Sun

Peaks site, peak trap catch occurred between July 27 and August 8, with 64 beetles

collected July 27 and 40 beetles collected August 8. There were additional small trap

catches until early September (158 beetles in total). In 1999, at the Cherry Ridge site, 605

J. ENTOMOL. SOC. BRIT. COLUMBIA 114, DECEMBER 2017 27

western balsam bark beetles were caught in two funnel traps, with the maximum number

of beetles collected on August 3, similar in timing to insects collected at the Sun Peaks.

Figure 1. Photograph of a western balsam bark beetle gallery system, showing central nuptial

chamber, four egg galleries and two parent females.

In 2002, insects were collected regularly from six additional sites, then counted and

sexed (Figure 2). In total, in 2002, 348 trap samples at nine sites were collected and

assessed, with 4,897 western balsam bark beetles caught from June 24 to September 10.

Significantly (t test, p<0.05) more males than females were trapped early in the flight

season (Figure 2). After July 9, 2002, equal or fewer male than female beetles were

trapped. |

Western balsam bark beetles were collected from traps beginning in mid- to late June,

with the exception of one high-elevation site at Spius Creek, west of Merritt, B.C., where

no beetles were collected until mid-July (Figure 3). At Sun Peaks, where there were three

trapping sites on an elevational gradient, beetle flight occurred earlier at the lower-

elevation site, gradually increasing in insect numbers later in the season at the higher-

elevation site. Smaller numbers of beetles were caught at the high-elevation site at the

onset of the flight period, with the majority trapped from July 9-20 (Figure 3). Although

sites at or above 1,600 meters in elevation had high trap catches (Figure 3), there was no

significant difference (p>0.05) in mean trap catch numbers at the different elevations

(1,450 m—1,600 m; 1,600 m—1,750 m; 1,750-1,900 m). There was no significant

difference in trap catch numbers between the ESSF subzones, where the traps were

located.

A second lesser flight beginning in late August was evident at Spius Creek, Torrent

Creek, Sun Peaks, and Apex in 2002. At Sun Peaks, beetles were caught in varying

numbers throughout the main flight period and into the second. Beetles were trapped in

high numbers at all three Sun Peaks sites, with the highest catch at the mid-elevation site

28 Je Entomol. Soc. Brit. Columbia 114, December 2017

(1,535 m) from June 24 to July 9. An elevational cline was observed at Sun Peaks, with

most beetles caught in the upper-elevation site (1,850 m) (July 9 through July 19), when

trap catches at the mid- and low-elevations sites were declining (Figure 3). This second

flight was small and comprised approximately 7% of the total number of insects trapped

at all study sites.

| ~ atotal females

2000 ee CE OT

| Total beetles caught = 4897

1500 -

No. D. confusus

“2A. jon 2-Jul te “46-Jul 23- Jul 30-Jul 6-Aug 13-Aug20-Aug27-Aug 3-Sep 10- Sag

Collection date

Figure 2. Total number of western balsam bark beetles collected during the 2002 flight period

from 348 traps at nine sites in the southern interior of B.C., from June 24 to September 10.

Asterisk above bars indicates significantly (t test, p<0.05) more males than females were

trapped. Total number beetles caught = 4897.

Not one of the four baited subalpine fir trees was successfully attacked in 1999.

Numerous beetles had initiated attack on the baited trees, as evidenced by the presence of

entrance holes and frass. However, minimal egg galleries and brood were found in

samples. From the nine non-baited trees attacked in 1997 and 1998, 7,257 life stages

were dissected and preserved for further study in the laboratory (Table 2).

Head capsule sizes ranged from 0.276 mm to 1.058 mm wide. Three distinct peaks

emerged from the measurements. First instar larvae ranged from 0.276 mm to 0.437 mm

(0.379 + 0.005 mm, mean + S.E.); second instar larvae from 0.460 mm to 0.644 mm

(0.549 + 0.002 mm); and third instar larvae from 0.667 mm to 1.035 mm (0.826 + 0.001

mm), with only two larvae with head capsules measuring 1.058 mm. Our data clearly

show three distinct larval instars based on head capsule size frequency distribution, and

there are not enough individuals from this study to confirm the possibility of a fourth

instar. Figure 4 illustrates the distribution of head capsule widths for larvae collected

from the attacked sample trees at Cherry Ridge and Sun Peaks sites. In total, 5,052

western balsam bark beetle larval head capsule widths were measured.

A summary of life stages found in 1999 from subalpine fir attacked in 1997 and 1998,

at Sun Peaks and Cherry Ridge, are presented as proportional data in Table 2. Although

life stages were dissected out of and counted from the 1997-attacked trees at Sun Peaks,

new adults had emerged prior to the onset of sampling. Therefore, emergence holes were

noted, but exact counts were not possible at that time.

In 1999, at Cherry Ridge, 3,442 life stages were dissected from three trees attacked in

1998. Two trees were standing attack, while the third tree was down on the ground.

Although the two standing trees contained variable numbers of beetles, the proportion of

each life stage dissected from the trees was similar (Table 2). This was in contrast to the

life stages dissected from the downed attacked subalpine fir, where, by the end of the

summer, only 4.5% of the insects dissected from this tree were teneral adults, compared

J. ENTOMOL. SOC. BRIT. COLUMBIA 114, DECEMBER 2017 29

to 18.3% and 21.7% of those dissected from the standing attacked trees. A similar pattern

existed in the 1998-attacked trees from Sun Peaks, where development was slower and a

smaller proportion of insects reached the teneral adult stage in the attacked downed trees

(Table 2, Figure 5).

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al i ve ne ri x ll a e e- oe oP Pe SH we ogg

Figure 3. Comparison of western balsam bark beetle a catches at nine sites in the

southern interior of B.C. (2002) from June 24 to August 30. Number of beetles caught

(N) is shown for each location.

There was a demarcated transition between life stages in all three standing trees

shown in Figure 5. Late-instar larvae were present June through mid-July, followed by a

3-4 week transition to pupae. At the end of August and into September, the majority of

life stages identified were teneral adults. In contrast, brood in down trees shown in Figure

5 displayed much slower development. Late-instar larvae were the predominant life stage

found throughout August in downed trees. The transition from larvae to teneral adults

was much slower in downed trees than in standing trees (Figure 5).

Seasonal temperature data collected from the phloem of trees by climate stations at

Sun Peaks and Cherry Ridge in 1998-1999 were summarized into hourly and daily

minimums, maximums and averages. The Sun Peaks climate station malfunctioned in

1999, therefore temperature data were not collected until July 25. Figure 6 shows

minimum and maximum daily phloem temperatures at Cherry Ridge from May 20 to

August 31, 1999. The recorded temperatures were similar until early July, when the

minimum temperatures collected from the phloem of standing trees became significantly

higher (t-test p<0.05) than the phloem temperatures recorded on the downed tree. No

beetles were caught in baited traps until the minimum daily temperature in the phloem

reached approximately 5° C. The same trend was observed from temperature data

collected from Sun Peaks. The difference in minimum daily phloem temperatures

between standing and downed trees was greater than the difference in maximum daily

phloem temperatures on standing and downed trees. Only in mid- to late August did

maximum daily phloem temperatures diverge significantly between standing and downed

trees (Figure 6).

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J. ENTOMOL. SOC. BRIT. COLUMBIA 114, DECEMBER 2017 31

Some early flight of western balsam bark beetle at Cherry Ridge may have been

missed due to traps not being in place until June 15, 1999. Sustained catches of western

balsam bark beetle occurred from early July through September. The combination of

exceeding a 5° C subcortical (phloem) daily minimum temperature and reaching or

exceeding 20° C subcortical maximum temperature appeared to initiate beetle emergence

and flight. There may have been minimal flight prior to trap placement when both these

parameters were met (Figure 6; June 10—June 17). Trap data from sites of similar

elevation did not have significant trap catches prior to July.

900

G75 9 ban a i eee eee

QE Ls seer ee eee

Gh oP oD Or pO QV Yh Dh Gan ae LY _ LP QP PLP \

d: Sg d: do: ° d: d: d: d: : cS: d: d: dv: cs: te. \-

Head capsule size (mm)

Figure 4. Frequency distribution histogram and estimation of larval instar for western balsam

bark beetle. A total of 5,052 head capsules were measured.

DISCUSSION

Our study confirmed the primary flight of western balsam bark beetle occurred from

late June through July, depending on site and elevation (Hansen 1996; Gibson ef al.

1997; McMillin et al. 2001; Negron and Popp 2009; Stock et a/. 2013). A much smaller,

secondary flight occurred later in the season, comprised primarily of females re-emerging

from the original host, either to initiate new brood galleries or to create hibernation

niches (Maclauchlan, personal observations). Mathers (1931) reported a secondary flight

occurring in mid- to late July; however, at the elevations monitored in this study, we only

saw this flight beginning in mid-August. Others have also reported a later start to the

secondary flight (Hansen 1996; Gibson et al. 1997; McMillin et al. 2001; Negron and

Popp 2009; Stock et al. 2013). Bark beetle flight and development are highly responsive

to temperature (Hansen ef al. 2001; Gaylord et a/. 2008) and latitude (Williams ef al.

2008; Bleiker and van Hezewijk 2016). Mathers’ (1931) sites were at more northerly

latitudes than our study areas, which could explain the difference in flight periodicity.

Our results show that the western balsam bark beetle is flexible and may initiate flight

earlier if weather conditions support beetle activity. This was seen at the Sun Peaks site,

where beetle flight was initiated two weeks earlier in 1998 than in 1999. The former year

was a record year for high temperatures in southern B.C. and across Canada

32 J. Entomol. Soc. Brit. Columbia 114, December 2017

(Environment and Climate Change Canada, Government of Canada. Canada’s Top Ten

Weather Stories of 1998 (https://ec.gc.ca/meteo-weather/default.asp?

lang=En&n=3DED7A35-1 Accessed on April 7, 2017).

SP Standing-2 1998 attack | SP Down-3 1998 attack

cistinstar g@2ndinstar m3rdmstar spupae zteneral i oistinstar @2ndinstar m3rdinstar spupae ateneral

No. insects

oN ER &

88sss 8

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ee ee ee en ee ee ae

8 8

No, insects

& 8

160 -

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@ 100 + -

2 +

2 0+

40 +: ae we Se Sa Oe Oe ae ee Ee ee Cee een

eo ee oes . oe pcceecccnnnngeoel a

S Se we g oS > LR Rk 2 ee

oop a PP PP, Pw, fe SPP PPL > ee PA, fe

eal 5. Western balsam bark beetle life stages dissected in 1999 from six 1998-attacked

subalpine fir at Sun Peaks (SP) and Cherry Ridge (CR). Each sample measured 20 cm x 20

cm and was centered over an entrance hole.

Beetles generally initiated flight once minimum daily phloem temperatures reached

or exceeded 5° C and maximum daily phloem temperatures approached 20° C or greater.

Maximum daily ambient temperature as a threshold for flight in bark beetles has been

determined in several studies (Stock 1991; Hansen 1996; Gaylord et a/. 2008), and our

determination of given site parameters is well within previously published data. Our

observations and data clearly indicate that both minimum and maximum phloem

temperatures are critical components to western balsam bark beetle flight activity and

play an integral role in initiating emergence, flight and dispersal, as well as being an

important factor in physiological development.

Subcortical, or phloem, temperature is likely an important factor for insect

development. Western balsam bark beetle is active under the bark very early in the

season, as evidenced by sawdust and frass being pushed out of entrance holes and

movement of life stages under the bark when excised (Maclauchlan, personal

observations). This activity has been observed as early as April, with flight not occurring

for another two or more months (Maclauchlan, personal observations). This early activity

highlights the fact that the physiological development of western balsam bark beetle

proceeds within very different temperature limits than is required for flight initiation.

Early season subcortical activity may allow female beetles from the late-season flight to

J. ENTOMOL. SOC. BRIT. COLUMBIA 114, DECEMBER 2017 33

initiate brood production very early the following summer, allowing brood increased

developmental time. With the early onset of warm weather in spring and often long,

extended summers, this potentially gives western balsam bark beetle better

developmental conditions and ultimately could provide an opportunity to shorten their

life history to one year, as suggested by Bright (1963).

on on oe ow Standing Standing-5 <+s++8« Down-4

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PB BP Poh Pr SP CF LP Ur US

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Date

Figure 6. Minimum and maximum temperatures collected from thermocouples inserted in the

phloem on three sample trees at Cherry Ridge research site, between May 20 and August 31,

1999. Arrow indicates date of first trap catch (July 8). Traps were established June 15 and

checked weekly.

The field collections confirmed three distinct larval instars based upon their

respective head capsule size range. The head capsule size range and average we found for

first and second instar larvae fall within the range found by Stock (1981). However, our

data do not clearly indicate a fourth instar; whereas Stock (1981) detected two

overlapping head capsule populations, that he identified as third and fourth instars.

Several larvae from controlled temperatures rearing conducted by the authors

(unpublished data) had very large head capsules (greater than 1.035 mm), hinting at the

possibility of a fourth instar; however, it was an uncommon occurrence in the field. The

occurrence of fourth-instar larvae was much more prevalent in a controlled temperature

setting (unpublished data) than in our field collections of life stages. Stock’s (1981) data

34 J. Entomol. Soc. Brit. Columbia 114, December 2017

were also obtained from controlled temperatures rearing. Our field data suggest that

diurnal fluctuations of subcortical temperature influence physiological development and

may prevent a final molt to fourth instar. Late-instar larvae often have higher temperature

thresholds for development than early instars do, preventing progression to cold-

susceptible advanced life stages before the onset of winter (Safranyik and Wilson 2006).

Under field conditions, the western balsam bark beetle may not have an obligate fourth

instar. |

Our results clearly elucidate that western balsam bark beetle developed more slowly

in attacked down trees than in attacked standing trees. We hypothesize that this is due to

host finding, host suitability, and a number of climatic factors. Host suitability and attack

success may depend on timing of the tree falling (seasoning) (Dyer 1967), placement of

the tree on the ground (touching the ground or somewhat elevated), and amount of shade

or direct sunlight on the bole (Schmid and Frye 1977) that could affect heat sums needed

for beetle development. Blowdown events are relatively uncommon in subalpine fir

stands, unlike the regular and often large-scale blowdown events seen in mature spruce

stands (Woods ef al. 2010). Thus, encountering downed trees is a relatively rare event for

this bark beetle, and its search patterns for down material may not be as discerning as for

vertical hosts. The highest frequency of blowdown is seen at stand edges (e.g., clearcut

edges) and in natural openings (personal observations). Stand edges or more open stand

scenarios might afford better ambient conditions for beetle development. Although ©

downed trees may offer the beetles more moderated conditions in late fall through early

spring due to the protective insulating characteristics of snow cover, this same snow

cover is often retained longer into the spring, depending upon log placement in a stand,

thereby keeping logs cool and delaying the onset of beetle development. Also, cooler

temperatures in downed trees occur earlier in late summer than in standing trees (Figure

6). Although a good resource from the point of view that the beetles encounter little or no

host resistance, it appears that beetle development is prolonged, potentially increasing

vulnerability to parasites, predators, woodborer activity, and host deterioration.

Proportionally, four to five times the number of brood in standing trees attained the

teneral adult stage prior to winter, compared to down trees at both sites. By the end of

summer, less than 10% of insects dissected from 1998-attacked downed trees had

reached the teneral adult stage. There was no indication that attack density differed

between standing and downed trees (Table 2) (Maclauchlan 2003), and large numbers of

larvae developed in the downed trees. However, the cooler and shorter season available

to beetles in downed trees suggest they are less suitable hosts for western balsam bark

beetle.

Both standing and downed subalpine firs were baited in 1999 to induce attack.

Although beetles initially responded to the bait and began excavating nuptial chambers,

the weekly sampling demonstrated that none of the trees was successfully mass attacked.

This has been observed in the past, where trees baited for western balsam bark beetle,

both standing and down, have high levels of unsuccessful attack, compared to natural

attacks occurring in close proximity (Maclauchlan et al. 2003; personal observation).

Bleiker et al. (2003) determined that western balsam bark beetle had discerning host-

selection capabilities, and parameters associated with host quality that the beetles are

able to detect may assist in subsequent developmental success.

Our results clearly revealed a two-year life cycle for western balsam bark beetle in

southern B.C. Additional work is needed to determine if western balsam bark beetle

larvae or teneral adults require a cold period, or if they can undergo continuous

development, as seen in the spruce beetle (Schmidt and Frye 1977), if growing seasons

lengthen. The presence of fourth-instar larvae in laboratory rearing suggests that a cold

period may be needed or conversely that western balsam bark beetle has evolved a

mechanism to postpone the pupation or eclosion process until warmer temperatures

trigger it (Johansson et al. 1994). Information on the response of this insect to changing

J. ENTOMOL. SOC. BRIT. COLUMBIA 114, DECEMBER 2017 a

habitat conditions as our high-elevation forest habitats continue to warm may be useful in

future forest planning, harvest and management.

ACKNOWLEDGEMENTS

We thank K. Buxton, T. Rimmer, K. Hicks and S. Moraes for field assistance. We also

thank R. Adams for installation of the climate stations. The project was supported in part

by funding from the Federal/Provincial Forest Resource Development Agreement

(FRBC) (1996-1997), Science Council of B.C. FR-96/97-422 and the British Columbia

Ministry of Forests, Lands and Natural Resource Operations Forests Sciences Program.

We thank B. Zimonick for editorial comments and two anonymous reviewers for detailed

and thoughtful suggestions that greatly improved the manuscript.

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38 J. Entomol. Soc. Brit. Columbia 114, December 2017

An updated and annotated checklist of the thick-headed

flies (Diptera: Conopidae) of British Columbia, the Yukon,

and Alaska

JOEL F. GIBSON!

ABSTRACT

The thick-headed flies (Diptera: Conopidae) are rarely observed parasitoids.

Confirmed hosts include many species of bees and wasps. Often collected from

flowers, conopids may serve as either pollinators or pollinator predators. The last

detailed checklist of the Conopidae of British Columbia was published in 1959.

An updated checklist for British Columbia, the Yukon, and Alaska is presented

based on over 1,000 specimens and specimen records. Geographical distribution,

using an ecoprovince approach, is documented for each of 26 species in the

region. Host, plant association, and hilltopping behavioural records based on past

literature and new observations are also included. An identification key to all

species recorded is included.

Key words: parasitoid, biogeography, plant associations, host associations,

Nearctic

INTRODUCTION

Conopidae (thick-headed flies) is a small, rarely collected family within the

acalyptrate Diptera. Many species are noted for their mimicry of wasps and bees. Adult

female conopids deposit eggs within living hosts using modified abdominal structures,

often in midflight. The larvae develop within the host, slowly consuming tissue, until the

host succumbs. Pupation occurs inside the host. Adult eclosion from the host’s corpse

usually follows an overwintering period. Various species of Hymenoptera are reported as

hosts, but confirmation of host status by rearing is rare (Gibson et al., 2014). The

possible impact of Conopidae on pollinator communities has been the focus of some

research (e.g., Schmid-Hempel and Schmid-Hempel 1996, Gillespie 2010, Malfi and

Roulston 2013). Conopids are also regularly collected from flowers but their role as

pollinators, or even their degree of plant specificity, is poorly documented. Studies

investigating specific flower associations or possible roles as pollinators have been few

and limited (Freeman 1966, Maeta and Macfarlane 1993).

Other aspects of Conopidae life history are understudied. Some species engage in

hilltopping behaviour (Mei et al. 2010), where males gather on hilltops or other

prominent geographical features to await females. The degree to which hilltopping

strategies are used by different species of Conopidae is poorly known.

Worldwide, more than 800 species of Conopidae — organized into six subfamilies and

59 genera and subgenera — are currently described (Gibson and Skevington 2013).

Species live in every region and continent except Antarctica and the Pacific Islands.

Williston (1882, 1883, 1885) described a large number of the western Nearctic species

and summarized the current knowledge in a series of papers. Later studies of Nearctic

species include Van Duzee’s (1927) review of California Academy of Sciences (CAS)

specimens and Parsons’ (1948) analysis of material from Harvard’s Museum of

Comparative Zoology (MCZ). The tireless work of Sid Camras includes revisions of

! Entomology, Royal BC Museum, 675 Belleville St., Victoria BC V8W 9W2; (250) 356-8242,

jgibson@royalbcmuseum.bc.ca

J. ENTOMOL. Soc. BRIT. COLUMBIA 114, DECEMBER 2017 39

individual North American genera (Camras 1943, 1944, 1953, 1955, 1957), regional

analyses (Camras and Hurd 1957), and continental catalogues (Camras 1965).

The Canadian summary of insect diversity (McAlpine et al. 1978) lists 30 species of

Conopidae, with fifteen more likely to be discovered or described. The most recent

review of the Conopidae fauna of British Columbia was that of Smith (1959), whose

checklist, based on 104 specimens from the Spencer Entomological Museum (University

of British Columbia) collection, included eighteen species in six genera. Smith did not

draw any conclusions about the intraprovincial distribution of each species. Neither

Insects of the Yukon (Danks 1997), nor Arctic Arthropods (Danks 1981), mentions

Conopidae. An updated list of conopid species in the northwestern Nearctic, along with a

summary of all known host and plant associations is necessary to assess the true

biodiversity and ecological impact of this family in the region.

MATERIALS AND METHODS

Taxonomic classification follows that of Gibson and Skevington (2013).

Morphological features are as described in Gibson and Skevington (2013) and Gibson et

al. (2013). Previous records of Conopidae confirmed from British Columbia, Alaska, and

the Yukon are tallied. Records include specimens loaned to the author as well as digital

databases of specimens with confirmed identifications. All notes regarding collection

date, plant associations, rearing from hosts, or hilltop locations are recorded. Previous

literature was examined to gather data on species range, host records, and plant

associations. Specimens examined, or confirmed specimen records, are listed in each

species account and are housed in the following collections: University of Calgary

Museum of Zoology, Calgary, Alberta (BDUC); California Academy of Sciences, San

Francisco, California (CAS); Canadian National Collection of Insects, Arachnids, and

Nematodes, Ottawa, Ontario (CNC); University of Guelph Insect Collection, Guelph,

Ontario (DEBU); Essig Museum of Entomology, University of California, Berkeley,

California (EMEC); Royal BC Museum, Victoria, British Columbia (RBCM); Royal

Ontario Museum Entomology Collection, Toronto, Ontario (ROME); Royal

Saskatchewan Museum, Regina, Saskatchewan (RSM); Spencer Entomological

Collection (Beaty Biodiversity Museum), University of British Columbia, Vancouver,

British Columbia (SEM); University of Alaska Museum Entomology Collection,

Fairbanks, Alaska (UAM); National Museum of Natural History, Washington, D.C.

(USNM); William F. Barr Entomological Collection, University of Idaho, Moscow, Idaho

(WFBM); Wallis-Roughley Museum of Entomology, University of Manitoba, Winnipeg,

Manitoba (WRME); James Entomological Collection, Washington State University,

Pullman, Washington (WSU).

An ecoprovince approach, similar to that of Ratzlaff (2015), is employed.

Ecoprovinces are defined as those in Demarchi (2011), Smith et al. (2004), and Gallant et

al. (1995). Ecoprovinces have been used in other recent insect checklists for British

Columbia (Scudder and Cannings 2009, Ratzlaff 2015) to summarize insect distributions.

With this method, the province is divided into area based on climatic, topographic, and

geological similarity (Demarchi 2011). There are ten ecoprovinces in BC (Fig. 1), each of

which could be considered unique sets of habitats. The presence of each conopid species

within these ecoprovinces has been recorded.

RESULTS AND DISCUSSION

Analysis of 1,016 specimens and specimen records has produced a list of 26 species

of Conopidae in British Columbia (Table 1). Six of these species also occur in Yukon,

and three of them are known in Alaska. This represents nine species added to Smith’s

(1959) checklist and, for sixteen more species, geographical ranges in BC, Yukon, and

Alaska are expanded. A complete list of all specimens included in the analysis —

40 J. Entomol. Soc. Brit. Columbia 114, December 2017

including collection localities, collectors, dates, and repositories — has been uploaded to

figshare (DOI: 10.6084/m9.figshare.5373361).

Figure 1. British Columbia ecoprovinces as adapted from Scudder and Cannings (2009).

Ecoprovince abbreviations: GD — Georgia Depression; CM — Coast and Mountains; SI —

Southern Interior; SIM — Southern Interior Mountains; CI — Central Interior; SBI — Southern

Boreal Interior; NBM — Northern Boreal Mountains; BP — Boreal Plains; TP — Taiga Plains.

Say (1823) describes Physocephala marginata from Missouri. Parsons (1948) reports

the range of this species as Kansas and Texas and East to Massachusetts, with isolated

individual specimens from Quebec, Ontario, Wyoming, and the Nicola Valley, British

Columbia. The inclusion of British Columbia in the range of P. marginata in this paper

and inclusion in Smith’s (1959) subsequent list is likely based on a single misidentified

specimen. Analyses recovered no specimens from British Columbia, Yukon, Alberta,

Saskatchewan, or Manitoba. This species likely does not occur in Canada west of

Ontario. (Note — this species has not been included in the summary of species in Table 1).

Confirmed rearing records from other regions were found for nine species. Plant

associations were recorded for all but one species. Hilltopping behaviour, including

observations of hilltopping in BC, was recorded for five species. Detailed discussion of

patterns of distribution and ecological associations follows a complete species checklist.

Key to species of Conopidae found in British Columbia, Yukon, and Alaska

r Labella subequal in length to prementum, filiform, at least partly fused, folded

back along prementum; metatibia without apical shiny patch; vein CuA2

straight; female abdominal tergite and sternite 5 separate; male cerci broadly

SEN ree in ae ay sae Rey Saat Cul any ORLY Aaa CET one chad raieasnaiaael 2

- Labella, shorter than prementum, broad, separate for entire length, projecting

forward from apex of prementum; shiny patch present near apex of metatibia;

J. ENTOMOL, SOC. BRIT. COLUMBIA 114, DECEMBER 2017 4l

8 et

vein CuA2 curved along its length; female abdominal tergite and sternite 5

fused; male cerci attached by narrow, sclerotized stalk ................cc eee eee eens 16

Ocellar and postocellar bristles absent; basisternum broad; veins Sc and Ri

separate for entire length; female abdominal tergite and sternite 6 completely

separate; female abdominal segment 7 laterally compressed along entire length;

phallus visible, elongate, ribbon-shaped, setose along entire length................

WP PREY i SINTER satel ees OR Dalmanniinae ... Dalmannia...3

Ocellar and postocellar bristles present; basisternum short, narrow, single

sclerite; veins Sc and R; fused before reaching costa; female abdominal tergite

and sternite 6 at least partly fused; female abdominal segment 7 rounded at least

in basal half; phallus usually not visible externally ................. Myopinae ... 5

Small species; total body length <4mm. Scutellum and femora entirely

GHSCR eee I ae a Dalmannia vitiosa Coquillet 1892

Larger; total body length >6mm. Scutellum and femora with some yellow ...... 4

Dorsal hairs black. Wings smoky ............. Dalmannia blaisdelli Cresson 1919

Dorsal hairs white. Wings hyaline ................ Dalmannia picta Williston 1883

Stout, reddish flies with a large white head; gena at least one third of total head

height; anterior margin of subcranial cavity rounded; costa uniform thickness

SAH CRAPS TOMO titty Bnei cds ea cnc opens oan Gude aada vee Myopa ... 6

Small (<10mm total length) black, shining flies; gena less than one third of head

height; anterior margin of subcranial cavity straight; costa thickened at endpoint

of SerRs Mitiiien wowed. tie. aa ee ee ae Thecophora ... 12

Wings. with: spots or With cross-Veins ClOUGOR osc sisson sien SG PNAS sins oe 7

Wings-completely byalines:::, .facioaie diatiol, wiod, Jats) eabesii emia... 8

Wings with only cross-veins clouded .................. Myopa vicaria Walker 1849

Wings with distinct spots in addition to clouded cross-veins ..................055

sia bacPote SATS gh MDA ANE MRBU MIDs nets stoi cibena BODE Catan ReMi Da Sate Myopa willistoni Banks 1916

Abdonién-almost entirely biaeki: laiios caaebee elias An otierst bids. is... 9

Abdomen almost entirely red; with or without pollinosity ...................0068. 10

Abdomen dorsally with long, black hairs ........... Myopa longipilis Banks 1916

Abdomen dorsally with short, pale hairs .............. Myopa vesiculosa Say 1823

Abdomen with dense, long black hairs dorsally; (this species is easily confused

with M. clausa and M. rubida) .............c00008 Myopa curticornis Kréber 1916

Abdomen with sparse, short black hairs dorsally ............... cee cee eee seen eeeee es 11

Abdomen dorsally with extensive pollinosity; (this species is easily confused

with M. curticornis and M. rubida) .......... cscs eceeees Myopa clausa Loew 1866

Abdomen dorsally without extensive pollinosity; (this species is easily confused

with M:clausa and..Md: CUrtiCornis) ics inis waveneva ved Myopa rubida (Bigot 1887)

Hind femora entirely black; abdomen with entirely pale hairs

pariaotd, Mes beetle caer ie align Thecophora propinqua (Adam 1903)

Hind femora at least partly yellow; abdomen with at least some black hairs .. 13

Large species (total body length >6mm); hind femora almost entirely yellow

scale ideidiamnlesasenaat deme eae es Thecophora modesta (Williston 1883)

Smaller species (total body length <6mm); hind femora partly yellow and partly

A) ea AN CEO 6 iia ie cratd uate eae oli eS a 14

Less than one third of hind femora yellow; (this species is easily confused with

T. nigripes and T) occidensis) .............006 Thecophora luteipes (Camras 1945)

One third to three quarters:of hind femora yellow vo). ijsi.cscseeccsaeccaccsecsecnds 15

One third to one half of hind femora yellow; (this species is easily confused with

T. luteipes and T. occidensis) ..............66 Thecophora nigripes (Camras 1945)

More than one half of hind femora yellow (this species is easily confused with 7.

luteipesand TL migripes) 6165. 6i4soaises seks Thecophora occidensis (Walker 1849)

16.

Ey,

Aes

J. Entomol. Soc. Brit. Columbia 114, December 2017

First abdominal segment as broad as thorax; arista mid-dorsal; second

aristomere equal in length to first; scape quadrate; ocellar tubercle well-

developed; ocellar and postocellar bristles present; anterior margin of subcranial

cavity rounded; maxillary palpal length at least equal to prementum width;

basisternum narrow, divided posteriorly, with elongate and narrow posterolateral

extensions; more than two pairs of scutellar bristles usually present; vena spuria

absent; epandrium separate beyond cerci ............. Zodioninae ... Zodion ... 17

First abdominal segment narrow and thread-like; arista stylate and apical:

second aristomere usually expanded ventrally; scape at least twice as long as

wide; ocellar tubercle reduced or absent; ocellar and postocellar bristles absent;

anterior margin of subcranial cavity projecting forward at junction with medial

carina; maxillary palpi reduced or absent; basisternum broad, posterolateral

extensions short and blunt; zero or one pair of scutellar bristles; vena spuria

present; epandriumy fused Deyoud cere. siisc6. ai een Conopinae ... 22

‘Thorax dark avitiewi rein golden Sirines i a a eevee

BREE a ere CREA cs cn, oe Merete CID ne Zodion obliquefasciatum (Macquart ae

Thorax either entirely dark or lighter with darker spots or stripes ................

Small species (total body length <Smm); thorax pale grey or green with a

U1 CEE | ee Mie epee RE. at Ue EN Zodion americanum Wiedemann 1830

Larger species (total body length >5mm); thorax variable, but usually dark, —

OMCs Career ie rer ye a Se, QI a 19

Smaller species (total body length <6mm); almost entirely grey with some

AER CE FEE iin cwinne See Meme ar ark AS Se. as eR ae el 20

Larger species (total body length >6mm); grey to dark grey with some reddish

colouration in the abdomen; with or without darker stripes ....................04. Zi

Third tergite of female abdomen longer than all other tergites; (males may be

indistinguishable from Z. cinereiventre) ........ Zodion perlongum Coquillet 1902

Third tergite of female abdomen equal in length or shorter than at least one other

tergite; (males may be indistinguishable from Z. perlongum) .................0006

ee isc Oh Peas pak Wi dna ca netsigance Mabe ta cade Zodion cinereiventre Van Duzee 1927

Abdomen with extensive red colouration; (this species is easily confused with Z.

EMORY SOU LE BE BOE as. QR Zodion fulvifrons Say 1823

Abdomen without red colouration or with red limited to outer margins; (this

Species 1s.casily contused: wilt A fh ifrons iii eA i

Ocelli and ocellar tubercle absent; ventral half of proepisternum bare; prominent

row of setae on posterior surface of mesofemur absent; metafemur expanded

PORTIS NAA tlens erie ud st OT OAS. AGA 8 Physocephala ... 23

Three ocelli present; ocellar tubercle present; ventral half of prosepisternum

with setae and/or bristles; prominent row of setae on posterior surface of

mesofemur present; metafemur parallel-sided along entire length

NAN RO PEF aad tesa tedden bate cdiahe-natae aia yd Physoconops ... 25

Colouration dark to black throughout, especially on frontal markings

RPE PE SPOR Len pT men ee Me Caer ETO Physocephala furcillata (Williston 1882)

Colouration reddish throughout, especially on frontal markings ................. 24

Black markings on scutum limited to a single central stripe; gena uniformly dark

leesiad ile oo guubephey evapneatve iste CRd A Ces LB Physocephala burgessi (Williston 1882)

Black marking on scutum broad; forming either three stripes or else covering the

entire dorsal surface; gena with paler central spot (this character often difficult

tO Meee, HAA a hs A Physocephala texana (Williston 1882)

Frons, second abdominal tergite, and all of the scutum very dark to black

Rarer ORM TN CNia) soe Physoconops (Physoconops) obscuripennis (Williston 1882)

J. ENTOMOL. SOc. BRIT. COLUMBIA 114, DECEMBER 2017 43

~ Frons, second abdominal tergite, and at least part of the scutum reddish or light

browmaitidss, A.ccRaee ian Physoconops (Physoconops) fronto (Williston 1885)

Table 1

Species of Conopidae recorded in British Columbia, Yukon, and Alaska by Smith (1959) and

the present study.

British Columbia Yukon Alaska

Species GD CM SI SIM CI SBI NBM BP

Pisigie dc laced ee a

Physocephala burgessi 8G. 8.6 8.6... -G.08,G

Physocephala furcillata * G

Physocephala texana EI i

Physoconops fronto * G

Physoconops obscuripennis Bu |S

Dalmanniinae

Dalmannia blaisdelli S,G

Dalmannia picta * Geng

Dalmannia vitiosa * G

Myopinae

Myopa clausa G BG Ss

Myopa curticornis * G ei G G

Myopa longipilis 5,G ad

Myopa rubida S,G SG G G

Myopa vesiculosa Sir Gs § G

Myopa vicaria S,G Cory Gras G 6G G G

Myopa willistoni * G G

Thecophora luteipes S,G ene Ay 39 ee

Thecophora modesta S.G G $§G. G S

Thecophora nigripes G G S,G G

Thecophora occidensis 5 pees white © akin «iene 8 5 G G

Thecophora propinqua 8:G SG ar’ S

Zodioninae

Zodion americanum * G J

Zodion cinereiventre * Cte rr

Zodion fulvifrons ht 9 i tet

Zodion intermedium Chie EP Ri ee <eE

Zodion obliquefasciatum * G

pease PL Wed Tr 1, AOE SES, = ANN ee eR Re a eens ke NA Te sec AT. A i ae

- species recorded in British Columbia for the first time. S — species recorded in each region by

age 1959. G — Species recorded in each region by the present study. Ecoprovince abbreviations:

GD — Georgia Depression; CM — Coast and Mountains; SI — Southern Interior; SIM — Southern

Interior Mountains; CI — Central Interior; SBI — Southern Boreal Interior; NBM — Northern Boreal

Mountains; BP -- Boreal Plains.

Species Checklist

CONOPINAE

Physocephala burgessi (Williston 1882)

Specimens or records observed: CAS, CNC, DEBU, EMEC, RBCM, RSM, SEM,

WRME, WSU. BC: Alta Lake, Apex Mountain, Bamberton Provincial Park, Clinton,

Cobble Hill, Courtenay, Cranbrook, Crowsnest Pass, Errington, Fitzgerald, Flathead, Fort

Langley, Forward Harbour, Gang Ranch Junction, Goldstream, Hope, Jesmond, Kaslo,

Keremeos, Kishinena Creek, Kleena Kleene, Maple Bay, Mount Alava, Mount Cain,

Mount Kobau, Mount Seymour, Nanaimo, Nelson, Ocean Falls, Osoyoos, Pemberton,

Qualicum, Quesnel, Revelstoke, Robson, Saanich, Salmon Arm, Salvus, Savary Island,

Sayward, Seton Lake, Shawnigan, Sidney, Squamish, Stagleap Provincial Park,

44 J. Entomol. Soc. Brit. Columbia 114, December 2017

Strathcona Provincial Park, Terrace, Tulameen, Upper Carmanah Valley, Vancouver,

Vaseux Lake, Vernon, Victoria, Walhachin, Wellington, Whistler.

Distributional notes: Williston’s (1882) description is based on type specimens from

Colorado and California. Parsons (1948) records the range as Montana to New Mexico

and west to California. Camras and Hurd (1957) and Camras (1957, 1965) list the range

for this species as Alberta to Texas and West to the Pacific Ocean. Smith (1959) includes

this species in his list for British Columbia. Analyses of other specimens indicate that

within Canada, P. burgessi has only been detected in British Columbia and Alberta.

Flight period: June - August

Ecological associations: In California, P burgessi has been collected from Prunus sp.

(Rosaceae) and Ceanothus sp. (Rhamnaceae) (Bohart 1941). Camras and Hurd (1957)

report Bombus sonorus Say 1837 (Apidae) as a host. Males were collected from the

summits of Mount Kobau, Mount Cain, and Mount Finlayson.

Physocephala furcillata (Williston 1882)

Specimens or records observed: CNC, RBCM. BC: Chetwynd, Fort St. John,

Hudson’s Hope, Rolla.

Distributional notes: Williston (1882) describes the species from New Hampshire.

Parsons (1948) records it from Wisconsin to Atlantic Canada, south to New Jersey, but

also in Mexico and California. Camras and Hurd (1957) and Camras (1957, 1965) report |

this species as found from Atlantic Canada, south to Pennsylvania and West to Alberta,

but also possibly in California and Mexico. Analyses of other specimens indicate that P.

furcillata is present in every Canadian province except Newfoundland and Labrador.

Flight period: June

Ecological associations: One specimen observed from Manitoba was reared from

Bombus terricola Kirby 1837. MacFarlane and Pengelly (1975) reared this species from

Bombus vagans Smith 1854 in Ontario. Specimens were collected from Solidago sp.

(Asteraceae), Arctium sp. (Asteraceae), and Chamerion angustifolium (Onagraceae)

flowers. Mei et al. (2010) suggested that this species is a likely hilltopper based on

specimens collected in the Ottawa area.

Physocephala texana (Williston 1882)

Specimens or records observed: CNC, DEBU, RBCM, ROME, RSM, WRME. BC:

Ashcroft, Cascade, Castlegar, Chilcotin, Chopaka, Christina Lake, Clinton, Cranbrook,

Dog Lake, Edgewood, Fairview, Farwell Canyon, Flathead Valley, Gang Ranch Junction,

Inkaneep Provincial Park, Kamloops, Keremeos, Lillooet, Midway, Mount Kobau,

Nicola River, Oliver, Osoyoos, Penticton, Robson, Soda Creek, Summerland, Vaseux

Creek, Vaseux Lake, Vernon, Walhachin.

Distributional notes: Williston (1882) describes the species based on specimens from

California, Texas, and Kansas. Parsons (1948) documents the range of P texana as

California to Georgia, with an additional specimen from Quebec. Camras and Hurd

(1957) list it throughout the USA, but rare in the west; and Camras (1957, 1965) reports

it occurring throughout the USA, Canada, and into Mexico. Smith (1959) includes P.

texana in his list for British Columbia and other data indicate that it ranges east to

Quebec and Nova Scotia.

Flight period: June - September

Ecological associations: This is one of the few species confirmed as a parasitoid of

honey bees (Apis mellifera Linnaeus 1758 (Apidae)). It has been reared from commercial

bees in Wyoming and Washington (Van Duzee 1934, Riedel and Shimanuki 1966). In

California, it has been seen to attack, oviposit in, and emerge from Bembix occidentalis

buettenmuelleri Fox 1901 (Crabronidae) and B. comata Parker 1917(Bohart and

MacSwain 1939, 1940). It was reared from Nomia melanderi Cockerell 1906 (Halictidae)

in Idaho (Foote and Gittins 1961). Hobbs (1965, 1966) also reported this species as

“killing” queens of Bombus rufocinctus Cresson 1863 and B. fervidus (Fabricius 1798) in

J. ENTOMOL. SOC. BRIT. COLUMBIA 114, DECEMBER 2017 45

southern Alberta. It was also reared from B. bifarius Cresson 1878, B. californicus Smith

1854, B. flavifrons Cresson 1863, and B. occidentalis Greene 1858 in Alberta

(Otterstatter et al. 2002).In California, P texana frequents and mates on flowers of

Eriogonum sp. (Polygonaceae) and Heliotropium sp. (Boraginaceae) (Bohart and

MacSwain 1939). Freeman (1966) reports an association between this species and

flowers of Asclepias fascicularis (Apocynaceae), Achillea millefolium (Asteraceae),

Melilotus alba (Fabaceae), Mentha sp. (Lamiaceae), and Chrysothamnus sp.

(Asteraceae). It hilltops in Quebec (Mei et al. 2010).

Physoconops (Physoconops) fronto (Williston 1885)

Specimens or records observed: CAS. BC: Vernon.

Distributional notes: Williston (1885) describes this species from Kansas. Parsons

(1948) lists the range as Nebraska to Texas, west to California, with a single specimen

from Massachusetts. Camras and Hurd (1957) and Camras (1955, 1965) describe the

range as Massachusetts to Florida, west to California and Washington, south to Mexico.

Other specimens indicate that P fronto occurs in British Columbia, Alberta, and

Manitoba.

Flight period: August

Ecological associations: Bohart and MacSwain (1940) reared a specimen of Conops

argentifacies VanDuzee (a synonym of P. fronto) from Megachile (Xanthosaurus)

perihirta Cockerell 1898 (Megachilidae). Foote and Gittins (1961) reared it from a

nesting site of Nomia melanderi. Freeman (1966) lists the following plant associations

for P. fronto: Asclepias fascicularis, Chrysothamnus sp., Daucus carota (Apiaceae),

Melilotus alba, and Solidago sp.

Physoconops (Physoconops) obscuripennis (Williston 1882)

Specimens or records observed: CNC, RBCM, SEM. BC: Kamloops, Oliver,

Osoyoos, Penticton, Robson.

Distributional notes: Williston (1882) describe this species from South Carolina.

Parsons (1948), Camras and Hurd (1957), and Camras (1955, 1965) give its range as

Massachusetts to Florida, west to Alberta, British Columbia and Washington, likely in

California. Smith (1959) includes this species in his list for British Columbia. Other

Canadian specimens of P. obscuripennis are from British Columbia, Alberta, Manitoba,

and Ontario.

Flight period: June - July

Ecological associations: Freeman (1966) reports this species on flowers of Cirsium

arvense (Asteraceae), Melilotus alba, and Solidago sp.

DALMANNIINAE

Dalmannia blaisdelli Cresson 1919

Specimens or records observed: CNC, RBCM, SEM. BC: Kilpoola Lake, Old Hedley

Road, Oliver, Penticton, Vaseux Creek, Vernon.

Distributional notes: The original description by Cresson (1919) lists Colorado as the

type locality with paratypes from California. Bohart (1938) only reported specimens

from California. Camras and Hurd (1957) and Camras (1965) list the range as Colorado

and Wyoming, west to Oregon and California. Analyses of other Canadian specimens

indicate that D. blaisdelli has only been detected in British Columbia.

Flight period: May

Ecological associations: Bohart (1938) mentions that the species is associated with

heavily wooded areas.

Dalmannia picta Williston 1883

Specimens or records observed: CNC, SEM. BC: Oliver, Robson.

46 J. Entomol. Soc. Brit. Columbia 114, December 2017

Distributional notes: In the original description, Williston (1883) lists the type locality

as New Mexico. Bohart (1938) and Parsons (1948) record specimens from Arizona and

California. Camras and Hurd (1957) and Camras (1965) list the range for this species as

British Columbia to New Mexico, west to California. Smith (1959) includes this species

in his list for British Columbia. Analyses of other specimens indicate that it occurs

nowhere else in Canada.

Flight period: May - June

Ecological associations: Bohart (1938) notes that specimens in the Mojave Desert,

California were collected near large aggregations of Diandrena sp. (Andrenidae) bees.

Freeman (1966) mentions Brassica nigra (Brassicaceae) as a plant association.

Dalmannia vitiosa Coquillet 1892

Specimens or records observed: CNC. BC: Robson.

Distributional notes: Coquillet (1892) describes the species based on a specimen from

Los Angeles, California. Bohart (1938) gives it a wide range (California, Virginia,

Kansas) and Parsons (1948) lists specimens from New Hampshire to Virginia, plus

California, Kansas, Arizona, and Nevada. Camras and Hurd (1957) and Camras (1965)

list the distribution as patchy across North America from Atlantic to Pacific. Analyses of

other specimens indicate that D. vitiosa has been collected in all Canadian provinces

except Manitoba, Prince Edward Island, and Newfoundland and Labrador. |

Flight period: May - June

Ecological associations: Specimens were observed on Cornus sp. (Cornaceae)

blossoms in Alberta. This species might demonstrate hilltopping behaviour based on

observations from Ontario and Quebec (Mei et al. 2010).

MYOPINAE

Myopa clausa Loew 1866

Specimens or records observed: CNC, RBCM, SEM. BC: Agassiz, Aspen Grove,

Bowser, Chilcotin, Courtenay, Creston, Kamloops, Kelowna, Keremeos, Oliver,

Penticton, Quesnel, Robson, Saanich, Sorenson Lake, Summerland, Vancouver, Victoria,

Yale.

Distributional notes: Loew’s (1866) type specimen is from Maine. Williston (1885)

lists the range as New England. Banks (1916) records it only in the East. Parsons (1948),

as well, limits the range from Maine to North Carolina, and possibly from Iowa, Arizona,

Washington, Wyoming, and California. However, Camras and Hurd (1957) and Camras

(1953, 1965) give the distribution of MM clausa as Maine to Georgia, west to British

Columbia and California. Smith (1959) includes it in his list for British Columbia.

Analyses of other specimens indicate that M clausa occurs in all Canadian provinces

except Manitoba, Prince Edward Island, and Newfoundland and Labrador.

Flight period: April - June

Ecological associations: Specimens from Quebec have been collected from Viburnum

acerifolium (Adoxaceae) flowers. Mei et al. (2010) concluded that this species may be an

exclusive hilltopper based on specimens observed in the Ottawa region.

Myopa curticornis Kroéber 1916

Specimens or records observed: ,RBCM, SEM, UAM. AK: Fairbanks; BC:

Cranbrook, Hatzic, Penticton, Robson, Salmon Arm, Vancouver, Vaseux Lake, Vernon,

Wellington; YT: Ross River.

Distributional notes: Kréber’s (1916) type specimens are from Colorado and

California. Parsons (1948) mentions specimens from Washington, Oregon, California,

Utah, Colorado, and Maine. Camras and Hurd (1957) and Camras (1953, 1965) list the

range as Wyoming to Arizona, west to Washington and California. Analyses of other

specimens indicate that, in Canada, M. curticornis only lives in British Columbia and

Yukon.

J. ENTOMOL. SOC. BRIT. COLUMBIA 114, DECEMBER 2017 47

Flight period: April - June

Ecological associations: Specimens from Alaska have been collected from Prunus

padus and Salix alaxensis (Salicaceae) flowers.

Myopa longipilis Banks 1916

Specimens or records observed: CNC, RBCM, SEM. BC: Agassiz, Kamloops, Oliver,

Osoyoos, Penticton, Robson, Vancouver, Vernon.

Distributional notes: Banks’ (1916) original types are from Washington State. Parsons

(1948) mentions specimens from Oregon and California. Camras and Hurd (1957) and

Camras (1953, 1965) list the range as British Columbia to Utah, west to California.

Smith (1959) includes this species in his list for British Columbia. Analyses of other

specimens indicate that M. Jongipilis is known in Canada from only British Columbia

and Alberta.

Flight period: April - May

Ecological associations: Freeman (1966) reports this species from Prunus subcordata.

Myopa rubida (Bigot 1887)

Specimens or records observed: CAS, RBCM, SEM. BC: Highlands, Robson,

Saanich, Vernon, Victoria; YT: Stewart Crossing.

Distributional notes: Bigot’s (1887) types are from Colorado. Banks (1916) records

the species from Oregon and Washington. Camras and Hurd (1957) and Camras (1953,

1965) list the range as west of the Rocky Mountains. Smith (1959) includes this species

in his list for British Columbia. In Canada, M. rubida occurs in British Columbia,

Alberta, and Yukon.

Flight period: May - July

Ecological associations: MacSwain and Bohart (1947) successfully reared this species

from Andrena vierecki Cockerell 1904 (Andrenidae) in California. Smith (1959) reports

it from Capsella bursa-pastoris (Brassicaceae). Freeman (1966) lists additional plants

visited by this species: Brassica campestris, Prunus sp., and Ranunculus californicus

(Ranunculaceae). A male was collected from the summit of Lone Tree Hill (Highlands,

BC).

Myopa vesiculosa Say 1823

Specimens or records observed: CAS, RBCM, SEM, CNC. BC: Cranbrook, Grand

Forks, Grindrod, Kamloops, Osoyoos, Penticton, Robson, Salmon Arm, Vernon; YT:

Ross River, Stewart Crossing.

Distributional notes: Say (1823) describes the species based on specimens from

Pennsylvania. Williston (1885) lists the species only in the eastern United States. Banks

(1916) records a specimen from Nebraska. Parsons (1948) mentions specimens from

New Hampshire to Virginia, west to Washington, while Camras and Hurd (1957) and

Camras (1953, 1965) list the range for this species as Quebec to Florida, west to

Washington and California. Smith (1959) includes M. vesiculosa in his list for British

Columbia and analyses of other specimens indicate that it lives in the Yukon and all

Canadian provinces except Prince Edward Island and Newfoundland and Labrador.

Flight period: April - June

Ecological associations: Specimens from British Columbia were collected from

flowers of Sorbus sp. (Rosaceae). This species may be an occasional hilltopper in

Ontario and Quebec (Mei et al. 2010).

Myopa vicaria Walker 1849

Specimens or records observed: CAS, CNC, RBCM, SEM, UAM. AK: Fairbanks;

BC: Atlin, Chilcotin, Cranbrook, Kamloops, Lavington, Nelson, Oliver, Peace River,

Penticton, Robson, Vancouver, Vernon; YT: Rampart House.

48 J. Entomol. Soc. Brit. Columbia 114, December 2017

Distributional notes: The type for this species, described by Walker (1849), is from

Nova Scotia. Parsons (1948) gives the range as Nova Scotia to Virginia, west to Illinois,

plus specimens from Washington, Oregon, Wyoming, and Arizona, while Camras and

Hurd (1957) and Camras (1953, 1965) list it as Nova Scotia to Georgia, west to Alaska

and California. Smith (1959) includes this species in his list for British Columbia; it

occurs in the Yukon and all Canadian provinces except Prince Edward Island and

Newfoundland and Labrador.

Flight period: April - May

Ecological associations: This species has been collected from various species of

willow (Salix alaxensis, S. arbusculoides, S. planifolia, S. pulchra, S. scouleriana) in

Alaska and also Salix sp. in Alberta. In their study in the Ottawa region, Mei et al. (2010)

did not find this species on hilltops.

Myopa willistoni Banks 1916

Specimens or records observed: CNC, RBCM, SEM. BC: Caulfield, Summerland,

Vancouver, Vaseux Lake, Vernon.

Distributional notes: Williston (1885) originally describes this species as M.

pictipennis, which is a preoccupied name, from Arizona and California; Banks (1916)

provided the new species name and saw specimens from Oregon and California. Camras

and Hurd (1957) and Camras (1953, 1965) list the range as west of the Rocky Mountains,

south into Mexico. Analyses of other specimens indicate that M. willistoni has only been —

found in Canada in British Columbia.

Flight period: May

Ecological associations: None noted.

Thecophora luteipes (Camras 1945)

Specimens or records observed: CAS, DEBU, RBCM, SEM. BC: Hell’s Gate,

Penticton, Robson, Sparwood, Thetis Lake, Vernon, Westwick Lake.

Distributional notes: Camras (1945) describes this species based on specimens from

Colorado, Washington, Idaho, Utah, and California. Camras and Hurd (1957) and

Camras (1965) record the range as British Columbia to Colorado, west to California.

Smith (1959) includes this species in his list for British Columbia and examination of

other specimens shows that, in Canada, it only occurs in that province.

Flight period: June - September

Ecological associations: Freeman (1966) summarizes plant associations for this

species as: Crepis virens (Asteraceae), Daucus carota, Eriogonum elatum, and Trifolium

repens (Fabaceae).

Thecophora modesta (Williston 1883)

Specimens or records observed: CAS, CNC, RBCM, SEM. BC: Agassiz, Chase,

Clearwater, Comox, Creston, Hope Mountains, Kootenay Lake, Lillooet, Metchosin,

Midday Creek, Mount Kobau, Newgate, Okanagan, Oliver, Robson, Salmo, Vancouver,

Vernon, Victoria, Walhachin; YT: Dawson.

Distributional notes: Williston (1883) describes this species based on specimens from

California and Washington. Camras and Hurd (1957) and Camras (1945, 1965) record

the range of this species as Saskatchewan to New Mexico, west to the Pacific Ocean.

Smith (1959) includes this species in his list for British Columbia; it also occurs in Yukon

and Alberta.

Flight period: June - September

Ecological associations: Cole and Lovett (1921) report Halictus ligatus Say 1837

(Halictidae) as a host for this species in Oregon. Freeman (1966) includes: Anaphalis sp.

(Asteraceae), Brassica rapa, Cirsium sp., Solidago sp., and Trifolium hybridum as plant

associations. Individuals have been observed on the summits of Mount Tolmie and

Camas Hill (BC: Victoria region).

J. ENTOMOL. SOC. BRIT. COLUMBIA 114, DECEMBER 2017 49

Thecophora nigripes (Camras 1945)

Specimens or records observed: CNC, RBCM, SEM. BC: Australian, Burton,

Clinton, Merritt, Mount Kobau, Oliver, Penticton, Prince George, Saanich, Vancouver,

Westwick Lake.

Distributional notes: Camras (1945) describes this species based on a specimen from

Thunder Bay, Ontario, but mentions 132 paratypes from across Canada, USA, and

Guatemala. All subsequent works (Parsons 1948, Camras and Hurd 1957, Camras 1965)

record the range as Nova Scotia to Georgia, west to British Columbia and California,

south to Guatemala. Smith (1959) includes this species in his list for British Columbia. T.

nigripes has been found in all Canadian provinces except New Brunswick, Prince

Edward Island, and Newfoundland and Labrador.

Flight period: July - August

Ecological associations: Plant associations for this species (Freeman 1966) include

Chrysothamnus sp., Crepis virens, Chamerion augustifolium (Onagraceae), Prunus sp.,

and Solidago sp. Mei et al. (2010) are unsure if the species hilltops in the Ottawa area.

Thecophora occidensis (Walker 1849)

Specimens or records observed: BDUC, CAS, CNC, RBCM, SEM, UAM. AK:

Fairbanks, Matanuska; BC: Agassiz, Burton, Chilcotin, Cowichan, Cottonwood River,

Crowsnest, Flathead Valley, Galiano Island, Hedley, Hope Mountains, Kalamalka Lake,

Kootenay, Langford, Lytton, Mahoney Lake, Mount Kobau, Nanaimo, Nicola, Oliver,

Osoyoos, Penticton, Quesnel, Robson, Salmo, Sheep Lake, Soda Creek, Sparwood,

Strathcona Provincial Park, Thetis Lake, Vancouver, Vernon, Victoria, Walhachin,

Westwick Lake; YT: Carmacks, Dawson, Lone Tree Creek, Old Crow, Starr Creek,

Tagish, Whitehorse.

Distributional notes: Walker (1849) describes the species based on a specimen from

Ohio. Camras (1945), Parsons (1948), and Camras and Hurd (1957) describe the range of

Occemyia loraria, a synonym of T. occidensis, as throughout the USA and southern

Canada. Camras (1965) records the range of T. occidensis as Quebec to Georgia, west to

the Yukon and California, south to Mexico. Smith (1959) includes O. loraria Loew 1866

in his list for British Columbia. It also lives in Yukon, Northwest Territories, and all

Canadian provinces except New Brunswick and Prince Edward Island.

Flight period: June - September

Ecological associations: This species has been reared from Halictus confusus Smith

1853, H. ligatus, H. rubicundus (Christ 1791), Lasioglossum cinctipes (Provencher

1888), L. forbesii (Robertson 1890), L. imitatum (Smith 1853), L. laevissimum (Smith

1853), and L. lineatulum (Crawford 1906) in Ontario (Smith 1966, Knerer and Atwood

1967). In his list, Freeman (1966) lists plant associations for ZT. loraria as: Brassica

campestris, Chrysothamnus sp., Daucus carota, Hypericum perforatum (Hyperiaceae),

Melilotus sp., and Solidago sp. This species may or may not demonstrate hilltopping

behaviour in Ontario and Quebec (Mei et al. 2010).

Thecophora propinqua (Adams 1903)

Specimens or records observed: CAS, CNC, RBCM, SEM. BC: Cranbrook, Erickson,

Kamloops, Lytton, Midway, Mount Kobau, Osoyoos, Penticton, Robson, Saanich,

Saturna Island, Vernon.

Distributional notes: Adams (1903) does not provide a locality for the type. Parsons

(1948), Camras and Hurd (1957), and Camras (1965) record the range as Nova Scotia to

Alabama, west to British Columbia and California. Smith (1959) includes this species in

his list for British Columbia. Analyses of other specimens indicate that 7: propinqua

occurs in all provinces from British Columbia to Quebec.

Flight period: May - September

50 J. Entomol. Soc. Brit. Columbia 114, December 2017

Ecological associations: Specimens were collected on Mentha sp. from Vernon,

British Columbia. Freeman’s (1966) plant association list for this species includes:

Achillea millefolium, Amaranthus sp. (Amaranthaceae), Asclepias fascicularis, Brassica

nigra, Chrysothamnus sp., Cleome lutea (Cleomaceae), C. serrulata, Daucus carota, D.

pusillus, Eriogonum elatum, Grindelia sp. (Asteraceae), Medicago sativa (Fabaceae),

Melilotus alba, Phacelia sp. (Boraginaceae), Solidago sp., Solanum tuberosum

(Solanaceae), Triticum aestivum (Poaceae). Mei et al. (2010) does not record any

hilltopping in the Ottawa area. ,

ZODIONINAE

Zodion americanum Wiedemann 1830

Specimens or records observed: CNC, RBCM, SEM. BC: Burton, Creston, Dog

Creek, Mount Kobau, Robson, Salmon Arm.

Distributional notes: Wiedemann’s (1830) type specimen is from Uruguay. Camras,

(1944, 1965), Parsons (1948), and Camras and Hurd (1957) list the range for this species

as throughout Canada, USA, central America, and into South America and the Caribbean

Islands. Zodion americanum has been recorded in all provinces except New Brunswick

and Newfoundland and Labrador.

Flight period: June - September

Ecological associations: Freeman (1966) reports possible plant associates as Melilotus

alba and Solidago sp. Mei et al. (2010) does not give evidence for hilltopping behaviour ©

for this species in the Ottawa area.

Zodion cinereiventre Van Duzee 1927

Specimens or records observed: CAS, CNC, RBCM. BC: Fernie, Mahoney Lake,

Nicola, Osoyoos, Pavilion Lake, Penticton.

Distributional notes: The type specimen of Van Duzee (1927) is from California.

Parsons (1948), Camras (1944), and Camras and Hurd (1957) give the range as

throughout the USA, west of Illinois. Camras (1965) lists the range as Atlantic Canada to

North Carolina, west to British Columbia and California. Other specimens indicate that

Z. cinereiventre lives in all provinces from British Columbia to Ontario.

Flight period: June - August

Ecological associations: Freeman (1966) reports that possible plant hosts for this

species include Helenium tenuifolium (Asteraceae) and Senecio sp. (Asteraceae).

Zodion fulvifrons Say 1823

Specimens or records observed: CNC, DEBU, RBCM, SEM, USNM, WFBC. BC:

Bear Lake, Chilcotin, Cranbrook, Grand Forks, Hell’s Gate, Jesmond, Junction

Provincial Park, Kamloops, Kaslo, Kelowna, Lillooet, Lytton, Midway, Mount Kobau,

Nelson, Nicola, Okanagan Falls, Oliver, Osoyoos, Penticton, Quesnel, Robson, Rock —

Creek, Royal Oak, Salmon Arm, Savary Island, Summerland, Vancouver, Victoria,

Walhachin, Dog Creek.

Distributional notes: Say (1823) describes this species from Maryland and

Pennsylvania. Camras (1944), Parsons (1948), and Camras and Hurd (1957) list the

range as Atlantic Canada to Florida, west to Washington and California, south to Mexico.

Smith (1959) includes this species in his list for British Columbia and other specimens

show that Z. fulvifrons occurs in all Canadian provinces except Newfoundland and

Labrador.

Flight period: May - August

Ecological associations: Severin (1937) reared this species from honey bees (Apis

mellifera) from South Dakota. Foote and Gittins (1961) report it from flowers of

Asclepias sp., Aster sp. (Asteraceae), Brassica sp., Chaenactis sp. (Asteraceae),

Chrysothamnus sp., Eriogonum sp., and Trifolium repens in Idaho. In Alberta it has been

J. ENTOMOL. Soc. BRIT. COLUMBIA 114, DECEMBER 2017 5]

collected from Solidago sp. flowers. Mei et al. (2010) suggest that this species might

hilltop in the Ottawa area.

Zodion intermedium Banks 1916

Specimens or records observed: CAS, CNC, DEBU, SEM, RBCM. BC: Boswell,

Dog Creek, Enderby, Fort Steele, Hudson’s Hope, Kamloops, Kinbasket Reservoir,

Lillooet, Mount Kobau, Nicola, Oliver, Osoyoos, Penticton, Quesnel, Robson, Rock

Creek, Sorrento, Telegraph Creek, Terrace, Vernon, White Lake.

Distributional notes: Banks (1916) describes this species from Pennsylvania. Camras

(1944), Parsons (1948), and Camras and Hurd (1957) list the range as Atlantic Canada to

Florida, west to Washington and California, south to Mexico. Smith (1959) includes this

species in his list for British Columbia; Z. intermedium has been collected in all

provinces except Newfoundland and Labrador.

Flight period: May - August

Ecological associations: Freeman (1966) summarizes plant associations for this

species as: Chrysothamnus sp., Brassica rapa, Erigeron canadensis (Asteraceae), E.

linearis, Lupinus sp. (Fabaceae), and Solidago sp. Specimens were observed from

Achillea sp. in Alberta and a Potentilla (Rosaceae) meadow in British Columbia. Mei et

al. (2010) suggests that this species might be a hilltopper in the Ottawa area.

Zodion obliquefasciatum (Macquart 1846)

Specimens or records observed: CNC. BC: Penticton.

Distributional notes: Macquart’s (1846) type specimen is from Texas. Parsons (1948),

Camras (1965), and Camras and Hurd (1957) list the range as Wisconsin to Louisiana,

west to Alberta, Washington, and California, south to Mexico. Zodion obliquefasciatum

has been recorded from British Columbia to Manitoba.

Flight period: July - August

Ecological associations: Freeman (1966) summarizes plant associations for this

species as: Chrysothamnus sp., Baileya pleniradiata (Asteraceae), Veronica sp.

(Plantaginaceae), Centaurea repens (Asteraceae), Cirsium arvense, C. vulgare,

Eriogonum sp., Gaillardia pulchella (Asteraceae), Grindelia sp., Gutierrezia

microcephala (Asteraceae), Helianthus annuus (Asteraceae), H. petiolaris, Hemizonia

fasciulata (Asteraceae), Heterotheca subaxillarias (Asteraceae), Medicago sativa,

Melilotus alba, M. officinialis, Lupinus sp., Bahia absinthifolia (Asteraceae), Verbesina

enceliodes (Asteraceae), Sphaeralcea angustifolia (Malvaceae), Asclepias sp., Solidago

canadensis, and S. occidentalis.

Zodion perlongum Coquillet 1902

Specimens or records observed: CNC. BC: Lillooet, Royal Oak.

Distributional notes: Coquillet (1902) describes this species from Colorado

specimens. Camras (1944), Parsons (1948), and Camras and Hurd (1957) list the range as

Maine to North Carolina, west to California, south to Mexico. Specimens examined

indicate that Z. perlongum lives in British Columbia, Alberta, Saskatchewan, Ontario,

Quebec, and Nova Scotia.

Flight period: June - August

Ecological associations: Freeman (1966) reports this species from flowers of

Chrysothamnus sp.

Patterns of Distribution. Smith’s (1959) checklist is limited in its data — only 104

specimens from a single collection (SEM) — and he drew no conclusions regarding

provincial distributions of Conopidae. The present data, including many more records

from more sources, allows some conclusions to be drawn. Nevertheless, most of the

records and specimens examined are from a subset of locations within the region. In

British Columbia, the Georgia Depression, Southern Interior, Central Interior, and

52 J. Entomol. Soc. Brit. Columbia 114, December 2017

Southern Interior Mountains ecoprovinces are relatively well-collected (Table 1). Few

specimens or records were noted from north of 53°N or from the coastal regions,

including the western coast of Vancouver Island, the Gulf Islands, or Haida Gwaii. In

Yukon, specimens or records from both the Boreal Cordillera and Taiga Cordillera are

reported, but not in any other ecoprovinces. Specimens from Alaska are limited to the

Cook Inlet and Interior Bottomlands ecoprovinces.

Based on recorded conopid distributions in British Columbia, Yukon, and Alaska, a

few general geographical distribution patterns are evident. Some species can be best

described as widespread, occurring in many regions of the northwestern Nearctic as well

as across the continent. Such species include: Physocephala texana, Dalmannia vitiosa,

Myopa clausa, M. vesiculosa, M. vicaria, Thecophora nigripes, T. occidensis, T.

propinqua, Zodion americanum, Z. cinereiventre, Z. fulvifrons, Z. intermedium, and Z.

perlongum. Other species appear to be limited to west of the Rocky Mountains:

Physocephala burgessi, Dalmannia blaisdelli, D. picta, Myopa curticornis, M. longipilis,

M. rubida, M. willistoni, Thecophora luteipes, and T: modesta. A few species are

southern in distribution with only a limited incursion into British Columbia, mostly in

warm Southern Interior valleys: Physoconops fronto, P. obscuripennis, Zodion

obliquefasciatum. Physocephala furcillata occurs throughout Canada, but, in British

Columbia, only east of the Rockies in the Peace region. Conopid species apparently able

to tolerate conditions north of 60°N are Myopa curticornis, M. rubida, M. vesiculosa, M.

vicaria, Thecophora modesta, and T: occidensis, although further collecting in these

regions may add to this list. Present records are insufficient to determine if any conopid

species are truly cordilleran or coastal in distribution.

Ecological Associations. Host records are scarce. However, some generalizations

regarding ecological roles can be drawn. Hosts appear to be determined roughly along

generic lines within Conopidae. Large species, especially those of Physocephala and

Physoconops, parasitize larger bees and wasps as hosts such as Apidae (Apis, Bombus)

and Crabronidae (Bembix). Smaller conopids, including Dalmannia, Myopa, and

Thecophora, are possibly limited to smaller bees as hosts (Andrenidae, Halictidae). There

are not enough host records to estimate host patterns for Zodion species.

Dalmannia and Myopa appear to be the early emerging genera within Conopidae as

adult records are limited to April through June. For all other genera, adults emerge from

June to September. There evidently are no phenological differences among species within

a given genus, but more records are necessary to clarify this point.

Plant associations as recorded may be a by-product of conopid phenology. Most

species of Conopidae frequent many different families of plants. The only discernible

pattern appears to be the early emergence of Myopa coordinated with some early-

blooming plants including willows (Salicaceae). Of course, these plant associations do

not necessarily indicate that the flies are pollinating the plants visited.

Hilltopping is observed in all genera of Conopidae. Whether this behaviour is

obligate, facultative, or geographically determined in any species requires further

observations. Accessible hilltop locations may provide valuable data on this question.

ACKNOWLEDGEMENTS

Thanks are extended to collection managers, curators, and staff at all collections

providing specimens and records. Thanks as well to Gina Capretta, Claudia Copley,

Henry Choong, Kendrick Marr, Rob Cannings, Syd Cannings, Rob McGregor, Dezene

Huber, and two anonymous reviewers for providing input on the manuscript.

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56 J. Entomol. Soc. Brit. Columbia 114, December 2017

Supercolonies of the invasive ant, Myrmica rubra

(Hymenoptera: Formicidae) in British Columbia, Canada

KEN NAUMANN|, MARIO MONIZ DE SA?, ELEANOR LEWIS’,

and ROSHAN NORONHA*

ABSTRACT

Levels of intra-specific aggression between workers and mtDNA sequence

comparisons were used to demonstrate that the non-native, invasive ant, Myrmica

rubra L. has formed supercolonies in southwestern British Columbia. Ants from

most, but not all, infested areas act aggressively towards ants from other areas but

workers from widely separated locations within two of the largest areas show little

aggression towards each other. Comparisons of COX1 mtDNA nucleotide

sequences suggest that formation of different supercolonies may have followed

possible divergence after a single initial introduction to the province.

Key words: invasive, super-colony, aggression

INTRODUCTION

Worldwide, over 150 species of ants have been introduced into new environments

(McGlynn 1999) but a small number have become invasive, i.e., have reduced native ant

biodiversity (Holway et al. 2002). Naumann and Higgins (2015), Gargas et al. (2007),

and McPhee et al. (2012) have all reported that recently-established populations of

Myrmica rubra L. in northeastern North America and the Pacific Northwest have all the

characteristics of an invasive ant. In southwestern British Columbia M. rubra populations

have dramatically decreased the incidence and abundance of previously established ants

in three different plant communities: a well-drained riparian zone dominated by

cottonwood (Populus balsamifera subsp. trichocarpa (Torrey and Gray) Brayshaw;

Salicaceae) and Scotch broom (Cytisus scoparius (Linnaeus) Link; Fabaceae); a moister,

more shaded community, dominated by red alder (A/nus rubra Bongard; Betulaceae), and

two exotic blackberries, Himalayan blackberry (Rubus discolor Weihe and Nees;

Rosaceae), and evergreen blackberry (Rubus laciniatus Willdenow); and grassy fields

(Naumann and Higgins 2015). They also occur at unusually high densities compared to

previously established species. Myrmica rubra represented more than 99.99% of the

total ant fauna caught in the infested areas, and their capture numbers in the plant

communities ranged from 10 to 1300 times the total number of ai// ants collected in

corresponding M. rubra-free areas. The numbers of several other taxa of insects and

non-insect arthropods were also reduced where M. rubra was present (also reported by

Gargas et al. 2007).

Myrmica rubra is native to Northern Europe and western Asia and was first

documented in North America in Massachusetts in 1908 (reviewed in Groden et al.

2005). It has now been reported in all Canadian provinces east of Manitoba and in at

least six northeastern United States, and Washington state. Most of the reports are from

within the last ten years, suggesting that the North American populations are expanding

| Departments of Biology and Health Science, Langara College, 100 W 49 Ave., Vancouver, BC

V5Y2Z6; kennethnaumann@langara.ca

Department of Biology, Langara College

3 lewisele@gmail.com

46675 Waltham Ave., Burnaby, BC, V5H3V6; roshananoronha@gmail.com

J. ENTOMOL. SOC. BRIT. COLUMBIA 114, DECEMBER 2017 57

(Wetterer and Radchenko 2011). North American populations have not been observed to

produce flying females (Hicks 2012), so spread is suspected to occur via the transport of

garden products and by colony budding. The species likely established in southwestern

British Columbia over 20 years ago but went relatively unnoticed for several years

(Higgins 2013). Myrmica rubra comes to the attention of the public mostly because of a

painful sting and high densities. Stinging is an unusual feature among the ant species

listed from British Columbia (Naumann et al. 1999) and can make yard and garden work

difficult and cause distress for pets. There is also concern that these ants may be

interfering with the successful nesting of some birds (Higgins 2013). Robinson et al.

(2013) estimated that the economic cost of this species in British Columbia could reach

$100 million/year if it spreads across its potential range in the province.

The formation of supercolonies may contribute to the ability of invasive ant species to

monopolize resources. An ant supercolony can be defined as a population of ants that

exists over a large, contiguous area and in which ants move freely between nests and

appear to show no aggression to conspecifics. (Haines and Haines 1978; Moffett 2012).

Ants within a supercolony are so numerous that is impossible for all members of the

colony to interact in their lifetime (Pedersen et al. 2006).

Holway et al. (2002) give a thorough review of the reported interactions between

invasive and native ants worldwide. Especially common are reports that invasive species

show greater efficiency at exploiting resources. Better resource utilization could be due

to a larger force of workers, i.e., more scouts and more foragers to recruit, and/or

physical aggression toward other species at a food item (Garnas et al. 2014).

Supercolonies are typically seen only in non-native populations and are typified by being

multiple-nested, multiple-queened, and lacking distinct behavioural boundaries among

physically separate nests. This sort of colony organization has allowed a small number of

non-native invasive species such as the Argentine ant, Linepethema humile (Mayr) (on all

continents except Antarctica); the little fire ant, Wasmannia auropunctata (Roger) (in

Africa, the Americas, and some Pacific islands); and the African big-headed ant,

Pheidole megacephala (Fabricius) (all continents except for Antarctica), to attain high

local abundances and consequently to dominate entire habitats (Holway et al. 2002). The

‘Large Supercolony’ of L. humile in California spans 1,000 km in distance (Moffett

2012). An apparent absence of intraspecific aggression within such supercolonies may

free up time and energy for other uses.

The purpose of this study was to gain a better understanding of how M. rubra has

come to dominate its new habitat in BC by determining if supercolony formation has

occurred. The number of behviourally and genetically distinct colonies may give insights

into whether there has been a single successful introduction or more than one.

METHODS AND MATERIALS

This study was carried out using ants from seven geographically distinct populations

of M. rubra in southwestern BC. Prior to this study, there was no indication of whether

those populations are the product of a single introduction or more than one. The

frequency of aggressive interactions between workers was used to determine whether

ants from the different areas, and from within them, treated each other as nestmates. The

degree of genetic similarity of workers from the same seven areas was estimated by

comparing nucleotide sequences of the mtDNA gene for cytochrome oxidase subunit I. It

was hoped that this would provide a molecular level confirmation of any patterns of

relatedness suggested by the behavioural data.

i) Sourcing and rearing the ants. Colonies of several hundred workers and at least

two queens were collected in the last week of May and first week of June 2014 from

nests within seven areas of infestation: Sea Island (Richmond), Fraser River Park

(Vancouver), Inter River Park (North Vancouver), University of BC (Point Grey),

Chilliwack, south Burnaby, and Oak Bay on Vancouver Island. It is unlikely that M.

58 J. Entomol. Soc. Brit. Columbia 114, December 2017

rubra has been established in each area for the same length of time. The source colonies

for this study were not identical in size, and not all workers were captured but the

assumption was made that this would not have an important influence on the behaviour

or individual workers. Each captured colony was maintained, in a laboratory, in a soil-

free, 34 x 23 x 8 cm lidded, plastic tub which contained an aluminum foil-covered, 13 x

9.5 x 5 cm plastic container that acted as the nest. The internal box contained multiple

folds of moist paper towel. Each colony was given a supply of water (a water-filled test

tube stopped with a cotton ball), 1:1 honey water mixture, apple slices, and recently

killed meal worms, and kept on a 12:12 h light-dark cycle.

ii) Inter-nest worker aggression between infestation areas. The level of aggression

in interactions between ant workers of the same species has often been used as a proxy

for levels of genetic difference — 1.e. as a method of determining nest mate recognition —

and many types of bioassays have been reported (Roulston et al. 2003). The recognition

system that ants use for identification with a colony and rejection of aliens is based on

shared cues, typically a colony-specific odour blend generated by queens or workers

(d’Ettorre and Lenoir 2010), although food and other environment factors can have an

influence (Liang and Silverman 2000). Our aim was to use the level of aggression

between workers from the seven different areas as a correlate of the degree of genetic

similarity. To minimize the confounding effects of foods and odours brought into the lab

colonies from their original environments, all colonies were maintained for at least one

week prior to being used for bioassays. It was assumed that several weeks in the lab

would not diminish the tendency for ants from different colonies to fight, which we

defined as ants locked together as they grasped each other with their mandibles.

Methods to measure the level of intraspecific aggression were similar to Roulsten et

al. (2003) and are summarized as follows. Sets of workers from each area of infestation

were matched with workers from a nest from each of the other areas. There were eight to

ten replicates (trials) for each pair. For each trial, five foragers from each of two colonies

were transferred to a fluon-coated 250 ml glass beaker which acted as a neutral arena.

The number of ants engaged in fights was recorded during five-second scan surveys

carried out once every minute for 10 minutes. For comparisons, we used the average (of

10 observations) percentage of ants involved in fights at one time across all colony pairs.

For half of the replicates, the first five ants into the arena came from one of the colonies

within each pair; for the other half, they came from the second colony. Controls

consisted of bioassays of two groups of five ants from the same colony.

The aggression bioassays were repeated a minimum of four weeks after capture, 1.e.,

during the second week of July, 2014. This was meant to test both that the initial one-

week latent period in the lab had been long enough to remove the effects of environment,

and that maintenance in the lab did not result in loss of aggression. This length of delay

was chosen because laboratory colony populations were beginning to decline at that time.

Colonies collected from Oak Bay were not included in this particular test because of

diminished worker numbers.

Within Infestations. The level of inter-nest aggression was also measured between

nests from within two of the largest areas of infestation, Sea Island in Richmond and

Fraser River Park in Vancouver. At Sea Island, four nests were collected at

approximately 500 m intervals along a 2 km transect line; at Fraser River Park, three

nests were collected approximately 300 m apart along a | km transect line. As before,

the nests were reared in the laboratory, as described above. Aggression bioassays were

carried out one week after the establishment of the nests; n = 6-10 for each pairing.

ii) The Genetic similarity of ants from different M. rubra populations.

Differences in mtDNA nucleotide sequences were measured as a way to determine if

there had been a single successful introduction of M. rubra into BC, or more than one. It

was also hoped that mtDNA differences could allow for discernment of ants from

different areas, i.e., different possible supercolonies. All mtDNA samples were collected

from workers from the nests used for the aggression bioassays.

J. ENTOMOL. SOc. BRIT. COLUMBIA 114, DECEMBER 2017 59

DNA was extracted from ants using a modified procedure of Schlipalius et al., 2001.

This procedure allowed for the use of whole insects combined with particular primers

that limit the possibility of contamination with microbial DNA. Individual frozen ants

were removed from storage at —80°C and immediately crushed in the bottom of a 1.5 ml

Eppendorf tube with an extraction buffer consisting of 30ul of boiling 5% Chelex in TE.

Each tube was then placed into a boiling water bath for 15 min and centrifuged at 13,000

rpm for 10 min. 20 pl of the supernatant was removed from each sample, and put into

storage at —20°C for later use as template DNA in PCR reactions.

The PCR primer pair LC1490 and HCO2198 (Folmer et al. 1994) were used for the

amplification of a 710 bp partial coding sequence of mitochondrial cytochrome oxidase

subunit I (COX 1). Primers were custom synthesized by INVITROGEN/ Life

Technologies ™. PCR was done using 2 ul of PCR buffer, 1 ul of 1 uM of primer

solution, | yl of Taq polymerase (Amplitaq, from Life Technologies), | ul of a 2 mM

dNTP, and | wl ant DNA, with dH20 added to a total volume of 20 pl. PCR was run on a

Techne Techgene thermal cycler. The program settings were: initial denaturation at 95°C

for 2 minutes; 30 cycles of the following: 30 seconds at 94°C, 45 seconds at 50°C, 2

minutes at 72°C, and a final extension for 5 minutes at 72°C. Successful amplification of

single 710 bp DNA from all ants was confirmed by agarose gel electrophoresis (data not

shown) and purified using a QIAquick PCR Purification Kit from QIAGEN. DNA was

sequenced using the Sanger method on an Applied Biosystems 3730 DNA analyzer at the

NAPS Unit at the University of British Columbia, Vancouver, BC.

Formica sinensis. Wheeler cytochrome oxidase subunit partial coding sequence

(Accession EU983580) was used as an outgroup to determine the order of descent among

DNA sequences. Phylogenetic analyses were done using MEGA7 (Kumar et al. 2016).

Sequences were imported into MEGA7 as fasta files and MUSCLE was used to generate

an alignment using the ALIGN CODONS option. Phylogenetic trees were generated

using the Maximum Likelihood Estimation (Tamura 1992; Felsenstein 1985). Initial trees

for the heuristic search were obtained automatically by applying Neighbor-Join and

BioNJ algortihms to a matrix of pairwise distances estimated using the Maximum

Composite Likelihood approach, and then selecting the topology with superior log

likelihood value. The analysis involved 17 nucleotide sequences. Codon positions

included were 1%*+2"4+3"4+Noncoding. All positions with less than 95% site coverage

were eliminated — 1.e., fewer than 5% alignment gaps, missing data, and ambiguous bases

were allowed at any position. There were a total of 438 positions in the final dataset.

RESULTS

Aggression Bioassays. With the exception of one pairing of localities, ants from

nests originating in different areas of southwestern BC showed high levels of worker-

worker aggression (Table 1). This included ants from Sea Island and Fraser River Park,

which are separated only by an arm of the Fraser River. The exception was a lack of

aggression observed when ants from Inter River Park in North Vancouver encountered

ants from Point Grey (University of British Columbia, Vancouver). The patterns of

fighting between workers from different localities did not change when tested again a

further four weeks after the nests were brought into the laboratory (Table 2).

There was comparatively little fighting between ants from nests within the Sea Island

or Fraser River Park M. rubra populations, even when nests were as much as 2 km apart

(Table 3).

Genetic Comparisons. Figure | shows that the nucleotide sequences of the COX]

subunits of the ants from the different outbreak areas fell into two groups. The North

Vancouver and Point Grey ants were within the same group. Different samples from

Fraser River Park in Vancouver fell within either group.

60 J. Entomol. Soc. Brit. Columbia 114, December 2017

Table 1 .

Mean percentage (+ SD) of ants from different pairs of colonies engaged in fights after one

week of laboratory rearing. The ants were from nests in seven different areas of southwestern

British Columbia. FR Park = Fraser River Park, Vancouver. Data with different superscripted

letters are significantly different (p < 0.05; ANOVA and LSD multiple comparison tests; F =

104.6; df = 20; P<0001. Same-nest comparison data were not included in the statistical

analysis).

Oak Bay Burnaby Chilliwack UBC |= NVan FR Park Sea Is

Sea Is TAG QP SIOZ" 72008) Pa 4426)" 6201 9)¢

FR Park SiG" SHAS B32) 84Gr. o80Cda)= ..0

N Van 78(12)*% 52(6)° 80(13)% 0(0) 0

UBC 92(27)"8 54(20)° 90(7) 0

Chilliwack 55(16)° = 83(6)2 0

Burnaby 7910)" 0

Oak Bay 0

Table 2

Mean (+ SD) percentage of ants from different pairs of colonies engaged in fights after six

weeks of laboratory rearing. The ants were from nests in six different areas of southwestern

British Columbia. FR Park = Fraser River Park, Vancouver; nests from a seventh locality

(Oak Bay) were not tested at the six week interval. Data with different superscripted letters

are significantly different (p < 0.05; ANOVA and LSD multiple comparison tests; F = 94.7; df

= 12; P<0.0001)). *Insufficient ants.

Burnaby Chilltwack UBC N Van FR Park Sea Is

Sea Is BHI) S321 S16) GAS 21S 0

FR Park Sie Op PO FIT ®

N Van ASQ BITSY & PME 0

UBC - ‘2 0

Chilliwack 63(21)¢ 0

Burnaby 0

DISCUSSION

Invasive populations of M. rubra have formed at least two large, multi-nest

supercolonies in BC, and it is reasonable that the same phenomenon has occurred in the

other distinct areas of infestation. The one on Sea Island is several kilometers across

and, as individual nests are often less than 5m apart, must contain thousands of nests and

millions of individual ants. This type of colony organization may be contributing to the

displacement of native ants and other epigaeic species that was reported by Naumann and

Higgins (2015).

J. ENTOMOL. SOC. BRIT. COLUMBIA 114, DECEMBER 2017 61

Based on aggression bioassays, most of the major outbreak areas of M. rubra in

southwestern BC represent different super-colonies, but workers from UBC and North

Vancouver interact as if they are nest mates, suggesting either a relatively recent common

origin, or that one site was the source of the founding population of the other.

Table 3

Mean (+ SD) percentage of ants from different pairs of colonies within the same outbreak

areas; SI = Seal Island; FP = Fraser River Park) that engaged in fights after one week of

laboratory rearing. The ants were from nests separated by approximately 500 m (SI) or 300 m

(FR) intervals along a transect line.

Sl S12 agEgIOn Sig FP! FP2 FP3

si4. | 27) 3(6) FP3

S33 | 3(6) 6) 0 FP2

eae Hs Pe FPI

si4 | 0

Levels of aggression between ants from different colonies are frequently used as a

proxy for levels of genetic difference (Roulsten et al. 2003). In this study, patterns of

aggression did not change markedly after a minimum of four weeks in the lab, suggesting

that it was not chemical cues associated with the original environments that led to

recognition of individuals from different locations, but rather colony-specific odor blends

generated by queens or workers (d’Ettorre and Lenoir 2010), and likely to be genetically

based.

We did not find enough molecular diversity in COXI to be able to distinguish

between different populations of M rubra in southwestern BC but the significant

separation into two groupings, with ants from Fraser River Park common to both, suggest

that an original introduction into BC may have occurred near there, and that divergence

of this subunit occurred later. The observation that ants from within the Fraser River

outbreak treat each other as nest mates argues against two genetically unique

introductions. The similarity of the COXI sequences of the non-aggressive ants from

UBC and North Vancouver provides further evidence that those two groups of ants are

particularly closely related. Hicks et al. (2012), also using mtDNA, reported evidence

that M. rubra populations on Newfoundland have come from at least four distinct

sources, including the UK and the Northeastern USA. We do not yet have enough data to

speculate on the possible source of the M. rubra populations in BC.

Possible mechanisms for the superior competitive abilities of invasive ant populations

include direct aggression, superior recruitment to resources, and higher activity levels.

Garnas et al. (2014) reported that M rubra shows both higher levels of recruitment and

aggression towards native ant species in Maine, USA; foragers consistently discover

baits faster and displace foragers from native species. Foragers from highly populous

supercolonies with many dispersed nests would have an advantage at discovering,

recruiting to, and exploiting food resources. For example, supercolony-forming L.

humile have been reported to be more numerous than other species in the same area, and

to be a superior interference competitor that displaces native species from contested

baits, often via direct physical aggression (Human and Gordon 1996). In addition, lack of

aggression between workers over large areas could leave more time and energy for

foraging. Linepithema humile for example, maintains higher colony activity levels,

62 J. Entomol. Soc. Brit. Columbia 114, December 2017

forages for longer periods each day, and recruits in greater numbers to food resources

than native species (Human and Gordon 1996).

Chilliwack

Oak Bay

Burnaby

Fraser River Park

Sea Isiand (A)

Sea Isiand (Si1)

Sea Isiand (S12)

Sea Island (S13)

Fraser River Park

UBC

North Vancouver

Formica CO1 (outgroup) 0.20

Figure 1. Molecular phylogenetic tree of the COXI subunit of mtDNA from M. rubra

workers from different outbreak areas of British Columbia. The tree with the highest log

likelihood (-1824.1574) is shown. The percentage of trees in which the associated taxa

clustered together is shown next to the branches. The tree is drawn to scale, with branch

lengths measured in the number of substitutions per site.

The proximate and ultimate causes of supercolony formation remain inconclusive.

According to Holway et al. (2002), the phenomenon is more common among ant species

that are non-native and have become invasive in their newly established environments.

They also tend to show relatively small size, omnivory, and a tendency towards multiple

queen nests. On the other hand, most of these species exhibit similar life histories in

their native ranges (Moffett 2012), and at least one other ant species, Liometopum

occidentale Emery, may form large (at least one km in diameter), habitat-dominating

supercolonies within its home range (Wang et al. 2010). Failure to form large colonies in

those areas may be due to constraints by other native species that are aggressive and

effective competitors. In other words, it is the release from those competitors in a new

region that allows for the formation of supercolonies (Moffett 2012). Supercolony

formation in M. rubra, as in other supercolony-forming species, may also be related to

the fact that virgin queens in North America do not carry out mating flights (Hicks 2012),

although they do in their home range. Instead, North American queens mate at or near

the nest and then travel a short distance, with a group of workers, to found a new nest.

Infestations thus expand relatively slowly via colony budding, and jump to new areas,

likely through human activities like the transport of infested nursery products. It is

possible that lack of contact with conspecifics from other colonies inhibits queen mating

flights or fails to stimulate them. If there are no intraspecific competitors in an adjacent

area, why risk a mating flight when territory that is likely to be suitable lies right next

door? Also, the success of incipient colonies is likely to be higher if the queen is not

alone, and if the number of founding workers is greater (reviewed in Holway et al. 2002).

Although it is now possible to add M. rubra to the list of invasive ant species that

share a suite of behavioural features such as supercolony formation, much work needs to

be done to resolve both the details of the M. rubra’s establishment in different areas of

North America, and the general mechanisms that lead to the formation of ant

supercolonies. Do some ants become ecologically dominant because they form

supercolonies or does the monopolization of resources by certain species lead to

supercolony formation (H6lldobler and Wilson 1977)?

J. ENTOMOL. SOc. BRIT. COLUMBIA 114, DECEMBER 2017 63

ACKNOWLEDGMENTS

Thanks to Emerald Naumann for assistance in collecting and maintaining colonies

and conducting bioassays; Sue Seward helped collect ants and format the manuscript;

and Rob J. Higgins provided advice and facilitated accessing study sites. Tyler House

assisted in maintaining the ant colonies in the lab and Kevin Craib gave statistical advice.

The following organizations and individuals gave permission to collect ants on lands that

they administer: the city of Vancouver, Sophie Dessureault; city of Richmond, Lesley

Douglas; the UBC Botanical Garden, Daniel Mosquin, Chris O’Rourke; Burnaby and

Region Community Garden Association, Eleni Haralias; the Canadian Wildlife Service

(for the Sea Island Conservation Area), Courteney Albert; and homeowners Nada

Traison, Julie-Ann Ichikawa, Tim Ebata, and Ben Van Drimmel. This work was funded

by a grant from the Langara College Scholarly Activity Committee.

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fire ant (Hymenoptera:Formicidae) in Newfoundland, Canada. Can Entomol. 146:457-464.

Higgins, R. 2013. European Fire Ant (Myrmica rubra) Project: Confirming Current Distribution in BC

and Development of Effective Control Methods Final Report. Prepared for the BC Inter-Ministry

Invasive Species Working Group. 26 pp.

Holldobler, B and E.O. Wilson. 1977. The number of queens; an important trait in ant evolution.

Naturwiss. 64:8-15.

Holway, D.A., L. Lach, A. V. Suarex, N. D. Tsusui and T. J. Case. 2002. The Causes and consequences

of ant invasions. Ann Rev Ecol Syst. 33: 181-233.

Human, K.G. and D. M. Gordon. 1996. Effects of Argentine ants on invertebrate biodiversity in northern

California. Conserv Biol. 11:1242-1248.

Kumar, S., G. Stecher, and K. Tamura. 2016. MEGA7: Molecular Evolutionary Genetics Analysis

version 7.0 for bigger datasets. Mol Biol Evol. 33:1870-1874.

Liang, D. and J. Silverman. 2000. “You are what you eat”: diet modifies cuticular hydrocarbons and

nestmate recognition in the Argentine ant, Linepithema humile. Naturwiss. 87:412-416.

McGlynn, T.P. 1999. The worldwide transfer of ants: geographical distribution and ecological invasions.

J Biogeogr. 26:535-548.

McPhee, K., J. R. Garnas, F. Drummond and E. Groden. 2012. Homopterans and an invasive red ant,

Myrmica rubra (L.), in Maine. Environ Entomol. 41:59-71.

Moffett, M. W. 2012. Supercolonies of billions in an invasive ant: What is a society? Behav. Ecol.

23:925-933.

Naumann, K. and R. J. Higgins. 2015. The European fire ant (Hymenoptera: Formicidae) as an invasive

species: impact on local ant species and other epigaeic arthropods. Can Entomol. 147: 592-601.

64 J. Entomol. Soc. Brit. Columbia 114, December 2017

Naumann, K., W.B. Preston and G. L. Ayre. 1999. An annotated checklist of the ants (Hymenoptera:

Formicidae) of British Columbia. J Entomol Soc BC. 96: 29-69.

Pedersen, J.S., M.J.B. Krieger, V. Vogel, T. Giraud, and L. Keller. 2006. Native supercolonies of

unrelated individuals in the invasive Argentine ant. Evolution. 60:782-791.

Robinson, D.C.E, D. Knowles, D. Kyobe and P. de la Cueva Bueno. 2013. Preliminary Damage

Estimates for Selected Invasive Fauna in B.C. Ecosystems Branch, British Columbia Ministry of

Environment, Victoria, B.C. 62 pp.

Roulston, T.H., G. Buczkowski and J. Silverman. 2003. Nestmate discrimination in ants: effect of

biosassay on aggressive behaviour. Insect Soc. 50:151-159.

Schlipalius, D.I., J. Waldron, B. J., Carrol, P. J. Collins, and P. R. Ebert. 2001. A DNA fingerprinting

procedure for ultra high-throughput genetic analysis of insects. Insect Mol Biol. 10(6), 579-585

Tamura, K. 1992. Estimation of the number of nucleotide sequences when there are strong transition-

transversion and G+ C-content biases. Mol Biol Evol. 9:678-687.

Wang, T.B., Patel, A., Vu, F. and P. Nonacs. 2010. Natural history observations on the velvety tree ant

(Liometopum occidentale): unicoloniality and mating flights. Sociobiology. 55:787-794.

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J. ENTOMOL. SOC. BRIT. COLUMBIA 114, DECEMBER 2017 65

SCIENTIFIC NOTE

First record of Aedes (Ochlerotatus) spencerii (Theobald)

(Diptera: Culicidae) in the Yukon

DANIEL A.H. PEACH!

Aedes spencerii (Theobald) (Diptera: Culicidae) is a small- to medium-sized

mosquito with characteristic alternating dark- and pale-scaled wing veins that is a

common inhabitant of grassland areas (Wood et al. 1979). It has two subspecies: Ae. s.

spencerii and Ae. s. idahoensis (Darsie and Ward 2005). Males and females have been

observed nectar feeding on catkins of willow (Salix sp.) (Knab 1907) and goldenrod

flowers (Solidago sp.) (Philip 1943), and females are known to take blood meals from

avian and mammalian hosts (Rempel et a/. 1946). It has very rarely been found carrying

western equine encephalitis (McLintock et al. 1970) and West Nile virus (Anderson ef al.

2015).

Ae. spencerii overwinters in the egg stage and is one of the first mosquito species to

emerge in the spring (Wood et al. 1979). Larvae are found in many habitats, including

pools of water formed by heavy rainfall, floodwater, or snow-melt (Belton 1983). Larvae

can develop rapidly at low temperatures, with pupae collected in mid-April near Ottawa,

Ontario, when the larvae of other Aedes spp. were only half grown (Wood ef al. 1979).

One or more generations will develop per year; however, suitable drying and subsequent

flooding of oviposition sites is necessary for the development of additional generations

beyond the first (Wood eft al. 1979). In some areas, females have emerged as late as

September when these conditions are met (Philip 1943).

The known distribution of Ae. spencerii ranges roughly from the Great Lakes to

central British Columbia, and from Colorado to Churchill, Manitoba. Scattered

populations also exist in Ottawa, Ontario, as well as in the states of New York, New

Jersey, and Oklahoma (Darsie and Ward 2005).

Two adult female mosquitoes attempting to bite the author were collected and placed

in ethanol on August 28, 2016, near Lake Creek campground in the Shakwak valley of

the southwest Yukon. The specimens were identified using the keys of Darsie and Ward

(2005) and Thielman and Hunter (2007) as Ae. s. spencerii (Fig. 1) and Aedes sticticus.

The collection site was mostly valley-bottom muskeg and riparian area with vegetation

present, including black spruce (Picea mariana), unidentified mosses, lingonberry

(Vaccinium vitis-idaea), willow (Salix sp.), and Labrador tea (Rhododendron

groenlandicum). Many snowshoe hares (Lepus americanus) and American red squirrels

(Tamiasciurus hudsonicus) were observed in the area. Growing nearby were patches of

fireweed (Chamerion angustifolium) and stands of aspen (Populus tremuloides).

Numerous small bodies of water were present in the vicinity, as was additional vegetation

that the author did not note at the time. The southwest Yukon is home to patches of

grassland, particularly in the Kluane Lake area (Laxton ef al. 1996; Conway and Danby

2014). This Ae. s. spencerii specimen may have originated from some such nearby patch

of grassland, or possibly from grassy patches along the margins of the nearby Alaska

Highway.

Mosquito collecting in the Yukon and the species recorded there were last reviewed

by Belton and Belton (1990). This is the first record of Ae. spencerii in the Yukon and

confirms the presence of Ae. sticticus, which was previously uncertain (Belton and

1 Department of Biological Sciences, Simon Fraser University, 8888 University drive, Burnaby, B.C.

V5A 186; dap3@sfu.ca

66 J. Entomol. Soc. Brit. Columbia 114, December 2017

Belton 1990). Both specimens have been deposited with the Beaty Biodiversity Museum

at the University of British Columbia.

Figure 1. Close-up of characteristic alternating dark- and light-scaled wing veination of

Ae. spencerii.

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10.1093/jmedent/7.4.446

Philip, C. B. 1943. Flowers as a suggested source of mosquitoes during encephalitis studies, and

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of Canada (Diptera: Culicidae). Agriculture Canada, Ottawa, ON. 279 pp.

68 J. Entomol. Soc. Brit. Columbia 114, December 2017

SCIENTIFIC NOTE

Cold requirements to facilitate mass emergence of spruce

beetle (Coleoptera: Curculionidae) adults in the laboratory

K. P. BLEIKER! and K. J. MEYERS

ABSTRACT— The spruce beetle, Dendroctonus rufipennis Kirby (Coleoptera:

Curculionidae, Scolytinae), is a native disturbance agent of spruce (Picea spp.) forests

in North America. Based on field observations, it is widely accepted that new adults

must overwinter regardless of the length of the life cycle. We tested the effect of

different lengths of time at 4° C on spruce beetle emergence. Our objective was to

determine a protocol for rearing spruce beetle to facilitate laboratory-based research.

We found that for spruce beetles from north-central Alberta and southern British

Columbia, 70 d at 4° C led to rapid mass emergence of adults. Adults also emerged in

the absence of a cold period, but over an extended period of time.

Key words: spruce beetle, Dendroctonus rufipennis, bark beetle, rearing, emergence,

diapause, overwintering |

The spruce beetle Dendroctonus rufipennis Kirby (Coleoptera: Curculionidae,

Scolytinae) is a native disturbance agent of spruce (Picea spp.) forests in North America.

Large-diameter weakened and injured trees, fresh-cut stumps, cull logs, windthrow, and

drought-stressed trees are the preferred hosts (Dyer and Taylor 1971; Safranyik 2011;

Hart et al. 2013). Populations may build up in these hosts and spill over into mature

healthy standing trees once the preferred hosts have been depleted (Safranyik ef al. 1983;

Safranyik 2011). Controlled rearing of spruce beetle in the laboratory may facilitate

experiments aimed at understanding factors affecting the population dynamics and

control of this economically important insect.

Bark beetles are easily reared in logs in the laboratory; however, some species require

a cold period to complete their life cycle (Ryan 1959). A two-year life cycle is common

throughout much of spruce beetle’s range; under warmer conditions, the life cycle may

be completed in one year and, in areas with cool, wet summers, the life cycle may take as

long as three years (Massey and Wygant 1954; Knight 1961; Berg et al. 2006; Werner et

al. 2006). Adult beetles emerge from overwintering in the natal host to attack new trees

in the spring, usually in late May or June, although attacks can occur throughout the

summer. Females bore into the inner bark, where they are joined by a male, mate,

construct egg galleries, and lay eggs. The majority of the life cycle is completed under

the bark, with larvae mining the inner bark before pupating and eclosing to new adults.

Cool temperatures trigger what has been described as a facultative larval diapause in the

two- and three-year life cycles, although the conditions that trigger the diapause and

instar sensitive to the cue might vary geographically (Dyer and Hall 1977; Hansen ef al.

2011). Based on numerous field observations, new teneral adults always overwinter once,

regardless of the length of the life cycle, and it is widely accepted that new adults must

overwinter to become sexually mature (Massey and Wygant 1954). Although it has not

been demonstrated experimentally, it is now widely accepted that spruce beetle has an

obligatory adult diapause (e.g., Raffa et al. 2015). New teneral adults overwinter in

windthrow or standing trees where they developed; however, a proportion of beetles in

standing trees may emerge in the fall, move to the base of the same tree, and bore under

the bark where they overwinter in groups insulated from cold winter temperatures and

1 Natural Resources Canada, Canadian Forest Service 506 West Burnside Road, Victoria, British

Columbia, Canada V8Z 1MS5; (250) 298-2365, katherine.bleiker@canada.ca

J. ENTOMOL. SOC. BRIT. COLUMBIA 114, DECEMBER 2017 69

woodpecker predation by the snow pack (Massey and Wygant 1954; Knight 1961; Fayt et

al. 2005).

In this paper, we report the effect of different lengths of time at 4° C on spruce beetle

emergence. Our objective was to determine a protocol for rearing spruce beetle in the

laboratory that would lead to mass emergence of new adults. We selected 4° C as our

treatment temperature, because it met the adult cold diapause requirements of Douglas-fir

beetle D. pseudotsugae Hopkins (Ryan 1959) and it could be easily achieved with a

regular refrigerator. We used two distant populations in the trial: Grande Prairie (GP) in

north-central Alberta (N54.860800, W118.713567; 649 m) and Placer Creek (PL) in

southern British Columbia (N49.15332, W120.47453; 650 m).

Spruce bolts (35-cm-long logs) were cut from a recently infested tree at each site and

transported to the laboratory at the Pacific Forestry Centre, in Victoria, British Columbia.

Five bolts were cut at GP in late-May 2013 and four bolts were cut at PL in mid-June

2013. The GP tree was likely pure white spruce (Picea glauca (Moench) Voss) and the

PL tree was likely pure Engelmann spruce (P. engelmannii Parry ex Engelm.), although

introgression between the species beyond their reported ranges is possible (see Maroja ef

al. 2007 and references therein). Parent beetles were constructing galleries and laying

eggs at the time the material was collected. The bolts were placed vertically in emergence

cages and held in a room at approximately 22° C with a photoperiod of 16 light:8 dark

until 29 August 2013, when the presence of fully darkened teneral adults was confirmed

by removing several small pieces of bark. As developing larvae were reared at 22° C, the

facultative larval diapause was not triggered. At this time, one bolt from each population

was left at 22° C and the other bolts were placed in a walk-in cold room at 4° C. The cold

room was dark, except when someone transferred material in or out of it. One bolt from

each population was removed after 15 (GP only), 30, 50 and 70 days in the cold room

and placed back at 22° C. Henceforth in this report, we use the population code followed

by the number of days at 4° C to refer to the different treatments. For example, GP30 and

GPO refer to insects from Grande Prairie, which were held at 4° C for 30 d and 0 d,

respectively, while PL30 and PLO refer to insects from Placer Creek receiving those

treatments.

Emerging beetles were collected at least three times per week. Average daily

emergence was calculated by dividing the number of beetles collected by the number of

days since the last collection. After at least 10 days of zero beetles emerging from a bolt,

the bark was removed and any teneral adults remaining under the bark were counted and

recorded as alive or dead.

The majority of beetles in all treatments emerged. Eighty-nine percent or more of the

beetles emerged, with two exceptions: PLO, 28% of the total number of beetles failed to

emerge and were found dead under the bark; and GP50, 18% of the total number of

beetles failed to emerge and most of these beetles were found alive under the bark (Table

1). For rapid mass emergence, the best cold treatment was 70 d for both beetle

populations (Figure 1). Beetles subjected to 70 d of cold had a notable increase in

emergence within 10 d after the cold treatment was terminated, and the vast majority of

beetles emerged rapidly within a 10-d period (Figure 1). Beetles in the 50-d treatment

also emerged soon after being removed from the cold room. However, after

approximately 65% and 80% of the beetles had emerged from PLS0 and GP50,

respectively, emergence plateaued for almost two weeks (Figure 1). The remaining

beetles emerged from PLSO but, in the case of GP5O, the bolt was peeled after 10 d of no

emergence and 17% of the total number of beetles were still under the bark and were

alive (Table 1); these beetles may also have emerged given more time. Our results are

similar to what Ryan (1959) reported for Douglas-fir beetle, which also overwinters as a

teneral adult: 90 d of cold treatment was more effective than 50 d for triggering

emergence. He also tested a number of cold temperatures from 0.5 to 12.5° C, and found

that 6.6° C was the optimum cold temperature, although emergence following 90 d at 6.6

and 4° C were similar.

70 J. Entomol. Soc. Brit. Columbia 114, December 2017

Table 1

Percentage of spruce beetles emerging from bolts (short logs) after 0, 15, 30, 50 or 70 d at 4°

C. Insects were fully darkened teneral adults when cold treatments were initiated. After at

least 10 d of no emergence, the bark was removed and the number of live and dead beetles

remaining under the bark was recorded. Bolts were cut at two sites: Grande Prairie (GP), in

north—central Alberta, and Placer Lake (PL), in southern British Columbia.

ey eee ne Peg wy ene ar

i 129 94 6 0

30 Zia 94 6 0

50 328 §2 ] 17

70 169 99 ] 0

30 259 95 5 0

50 281 98 y 0

70 282 89 11 0

Emergence tended to be slower from bolts receiving 30 d or less of cold treatment

(Figure 1). Emergence from both GP30 and PL30 did not notably increase until after 21 d

at 22° C, and emergence was also delayed after the cold treatment was terminated for

GP15, although once it started it was relatively rapid (Figure 1). In the absence of a cold

treatment, beetles emerged over an extended period of time (over 100 d) and it took

longer for beetles to start emerging from GPO than from PLO. The new adults may have

been at different levels of maturation, as the populations may have received slightly

different degree day accumulations based on temperatures in the field and when the trees

were infested and cut. In addition, there may be geographic variation in the

developmental rates of bark beetles (Bentz et a/. 2001) or the proportion entering

diapause (McKee and Aukema 2015). We also cannot determine if beetles were emerging

to overwinter at the base of the tree or to disperse to attack a new host tree. Ryan (1959)

determined that Douglas-fir beetles in diapause had underdeveloped reproductive organs.

We did use a number of the new adults emerging from GP70 and PL70 in another

experiment. These beetles entered fresh bolts and successfully reproduced, indicating

they were sexually mature upon emergence; however, we did not use beetles from the

other cold treatments, so their level of sexual maturity remains unknown.

Our results indicate that a cold period promotes the rapid mass emergence of new

spruce beetle adults. In the absence of a cold period, 72% of PL beetles and 96% of GP

beetles still emerged, although they emerged slowly over approximately 100 d. We have

demonstrated that 70 days of cold treatment at 4° C is sufficient to trigger mass

emergence of adult spruce beetles. This treatment can be used to rear insects in the

laboratory, thereby facilitating experimental research.

J. ENTOMOL. SOC. BRIT. COLUMBIA 114, DECEMBER 2017 71

1.0 ;

0.9 |

0.8 |

0.7 |

| a) Grande Prairie

0.6 4

0.5 |

0.4 |

0.3 |

Cumulative Emergence

0.2 |

0.1 |

0.0 [a= PA URS vate Pee ner eee ee

120 140 160 180 200

b) Placer Creek

Sag

oF)

& Days at 4°C

$ rae

& —15

& —— 30

O ll

‘lieahinlsed

unin -wanir “sear alleges ne re

Day

Figure 1. Cumulative emergence of all spruce beetles in bolts (short logs) after 0, 15, 30, 50

or 70 d exposure to 4° C before being held at 22° C (day 0). Bolts were cut at two sites: a)

Grande Prairie (GP), in north-central Alberta, and b) Placer Lake (PL), in southern British

Columbia.

ACKNOWLEDGEMENTS

We thank C. Leverett for collecting and processing emerging beetles, G. Smith, P.

Borrett and K. Hogg for help collecting infested wood in the field. We also thank L.

v2 J. Entomol. Soc. Brit. Columbia 114, December 2017

Safranyik and V. Nealis for commenting on an earlier version of the manuscript, as well

as B. Van Hezewijk for helpful discussion. We appreciate the improvements made to the

manuscript by two anonymous reviewers. This study was supported by Natural

Resources Canada.

REFERENCES

Bentz, B. J., J. A. Logan, and J. C. Vandygriff. 2001. Latitudinal variation in Dendroctonus ponderosae

(Coleoptera: Scolytidae) development time and adult size. The Canadian Entomologist 133:375—387.

Berg, E. E., J. D. Henry, C. L. Fastie, A. D. De Volder, and S. M. Matsuoka. 2006. Spruce beetle

outbreaks on the Kenai Peninsula, Alaska, and Kluane National Park and Reserve, Yukon Territory:

relationship to summer temperatures and regional differences in disturbance regimes. Forest Ecology

and Management 227:219-—232. 10.1016/j.foreco.2006.02.038

Dyer, E. D. A., and P. M. Hall. 1977. Factors affecting larval diapause in Dendroctonus rufipennis

(Coleoptera: Scolytidae). The Canadian Entomologist 109:1485-1490. 10.4039/Ent1091485-11

Dyer, E. D. A., and D. W. Taylor. 1971. Spruce beetle brood production in logging slash and windthrown

trees in British Columbia. Canadian Forest Service, Pacific Forest Research Centre, Victoria, BC.

Information Report BC-X-62. http://citeseerx.ist.psu.edu/viewdoc/download?

doi=10.1.1.832.7596&rep=rep | &type=pdf

Fayt, P., M. M. Machmer, and C. Steeger. 2005. Regulation of spruce bark beetles by yeas fa —a

eee review. Forest Ecology and Management 206: 1-14.

Hansen, E. M., B. J. Bentz, J. A. Powell, D. R. Gray, and J. C. Vandygriff. 2011. Prepupal diapause and

instar IV development rates of the spruce beetle, spruce beetle, Dendroctonus rufipennis (Coleoptera:

Curculionidae, Scolytinae). Journal of Insect Physiology 57:1347-1357. 10.1016/).jinsphys.

2011.06.011 :

Hart, S. J., T. T. Veblen, K. S. Eisenhart, D. Jarvis, and D. Kulakowski. 2013. Drought induces spruce

beetle (Dendroctonus rufipennis) outbreaks across northwestern Colorado. Ecology 95:930—939.

10.1890/13-0230.1

Knight, F. B. 1961. Variations in the life history of the Engelmann spruce beetle. Annals of the

Entomological Society of America 54:209—214. 10.1093/aesa/54.2.209

Maroja, L. S., S. M. Bogdanowicz, K. F. Wallin, K. F. Raffa, and R. G. Harrison. 2007. Phylogeography

of spruce beetles (Dendroctonus rufipennis Kirby) (Curculionidae: Scolytinae) in North America.

Molecular Ecology 12:2560—2573. 10.1111/.1365-294X.2007.03320.x

Massey, C. L., and N. D. Wygant. 1954. Biology and control of the Engelmann spruce beetle in

Colorado. US Department of Agriculture Circular, 944. 44 pp. https://archive.org/details/

biologycontrolof944mass

McKee, F. R., and B. H. Aukema. 2015. Successful reproduction by the eastern larch beetle (Coleoptera:

Curculionidae) in the absence of an overwintering period. The Canadian Entomologist 147:602—610.

10.4039/tce.2014.81

Raffa, K. F., B. H. Aukema, B. J. Bentz, A. L. Carroll, J. A. Hicke, and T. E. Kolb. 2015. Responses of

tree-killing bark beetles to a changing climate. Pages 1730201 in C. Bjérkman and P. Niemela, eds.

Climate change and insect pests. CABI, Oxfordshire, U.K. 292 pp. 10.1079/9781780643786.0173

Ryan, R. B. 1959. Termination of diapause in the Douglas-fir beetle, Dendroctonus pseudotsugae

Hopkins (Coleoptera: Scolytidae), as an aid to continuous laboratory rearing. The Canadian

Entomologist 91:520—525.

Safranyik, L. 2011. Development and survival of the spruce beetle, Dendroctonus rufipennis, in stumps

and windthrow. Natural Resources Canada, Canadian Forest Service, Pacific Forestry Centre,

Victoria, BC. Information Report BC-X-430. 21 pp.

Safranyik, L., D. M. Shrimpton, and H. S. Whitney. 1983. The role of host-pest interactions in the

population dynamics of Dendroctonus rufipennis (Kirby) (Coleoptera: Scolytidae). Pages 197-212 in

A.S. Isaev, ed. Role of host-pest interactions in the population dynamics of forest pests. Krasnoyarsk,

USSR.

Werner, R. A.; E. H. Holsten, S. M. Matsuoka, and R. E. Burnside. 2006. Spruce beetles and forest

ecosystems in south-central Alaska: A review of 30 years of research. Forest Ecology and

Management 227:195-206. 10.1016/j.foreco.2006.02.050

J. ENTOMOL. SOC. BRIT. COLUMBIA 114, DECEMBER 2017 ho

SCIENTIFIC NOTE

Production of epicormic buds by Douglas-fir in central

British Columbia, Canada, following defoliation by western

spruce budworm (Lepidoptera: Tortricidae)

LISA M. POIRIER!

Western spruce budworm, Choristoneura freemani Razowski (= C. occidentalis

Freeman), is an important defoliator of Douglas-fir, Pseudotsuga menziesii (Mirb.)

Franco, in southern British Columbia (Maclauchlan ef a/. 2006). Defoliation by larvae

can result in reduced tree growth, top-kill and occasional tree mortality (Alfaro ef al.

1982). Larvae feed on the new foliage of all ages of trees; mortality is most common in

immature and suppressed understorey trees (Maclauchlan and Brooks 2009), but

repeated, severe defoliation can kill larger trees.

Flushing of buds late in the season is a proposed mechanism by which conifers can

compensate for defoliation (Piene 1989), and increased late bud production may explain

why some species or individuals experience greater survival and faster recovery

following defoliation (Piene and Eveleigh 1996). The terminology used in the literature

for these late-flushing buds varies, but Meier et al. (2012) recommend calling them

epicormic buds, while sequential buds are those formed during shoot elongation.

The most recent western spruce budworm outbreak in British Columbia extended

farther north than had been observed previously. Near the northern edge of that outbreak,

my observations suggested that defoliated Douglas-fir had fewer flushed epicormic buds

at the end of the summer than might have been expected farther south. At higher

latitudes, a short growing season and early frosts may reduce the ability of trees to

compensate for defoliation by producing epicormic buds.

In 2010, two-year-old P. menziesii var. glauca seedlings were obtained from Pacific

Regeneration Technologies, Inc. in Red Rock, B.C. Seedlings were planted individually

in conical pots of all-purpose potting mix on May 17, when the earliest sequential buds

on local Douglas-fir trees began to swell. Third- to fifth-instar western spruce budworm

larvae were collected from two sites north of Williams Lake, B.C. (52.266° N, 122.285°

W and 52.471° N, 122.434° W) on June 16. Larvae were kept on live foliage in large

plastic bags and transported to the University of Northern B.C. (UNBC) in Prince

George, B.C. (53.893° N, 122.816° W), then transferred to the experimental seedlings

within 48 hr. Each seedling had 18—22 sequential buds at approximately the same stage

of development as those of Douglas-fir on campus.

Five treatments (n = 24 each) were applied on the same day as follows.

1. Control: No further manipulation of seedlings.

2. Control+mesh: A white, polyester-netting (BioQuip Products Inc.) cylinder was

secured with elastic bands over each seedling.

3. Scissors: All sequential buds were removed at the base with fine scissors.

4. Scissorst+mesh: Sequential buds were removed with scissors, and a netting cylinder

was secured over each seedling.

5. Larvae+mesh. A netting cylinder was secured over each seedling, and 10 larvae

were added. The larvae on each seedling ranged from approximately third to

fifth instar, representing the range and distribution of larvae collected at field

sites.

' Ecosystem Science and Management Program, University of Northern British Columbia, 3333

University Way, Prince George, B.C., Canada V2N 4Z9; (250) 960-6124, lisa.poirier@unbc.ca

74 J. Entomol. Soc. Brit. Columbia 114, December 2017

A sixth treatment, larvae on seedlings without mesh, was not possible due to the risks

of releasing insects in an area where they are not currently found. Seedlings were

randomized within racks and placed outside in the compound of the Enhanced Forestry

Laboratory at UNBC. They were watered daily, and pupae were removed twice weekly.

Once all larvae had pupated or died, the mesh bags were removed, and flushing

epicormic buds were counted for each seedling. Due to heteroscedasticity and small

sample size, treatments were compared using a Kruskal—Wallis rank sum test and a

Nemenyi post hoc test with chi-squared approximation for independent samples

(PMCMR v4.1 package; Pohlert 2014) within the R 3.4.0 statistical programming

language (R Development Core Team 2016).

Control seedlings, both with and without mesh bags, flushed no epicormic buds (Fig.

1). Destruction of sequential buds, by either scissors (P < 0.001 in all cases) or western

spruce budworm larvae (P = 0.029), significantly increased numbers of epicormic buds

over the control seedlings (Fig. 1). Seedlings defoliated by larvae had significantly fewer

epicormic buds than those defoliated with scissors (P = 0.034 for scissors, and P = 0.026

for scissors+mesh). In most cases, epicormic buds on seedlings with larvae showed

feeding damage.

son init nN

mo ra ©

Number of Epicormic Buds per Seedling

Contro! Control+ Mesh Larvae + Mesh Scissors Scissors + Mesh

Treatment

Figure 1. Numbers of epicormic buds per two-year-old Douglas-fir seedling. Control

seedlings were not defoliated; other treatments had expanding spring buds removed with

scissors or by feeding of third- to fifth-instar western spruce budworm larvae. Treatments

including “Mesh” had seedlings contained in white, polyester-netting cylinders. Boxplots

portray the median (midbar in the box), the 25" and 75" percentiles (box), lowest and highest

points within 1.5x the inter-quartile range (lower and upper vertical lines), and outliers (small

circles). Large circles show the mean values.

Seedlings defoliated by any means responded with production of new foliage late in

the summer. Seedlings with larvae caged on them had significantly fewer flushing

epicormic buds than did seedlings that had been defoliated with scissors. Defoliation by

insects can have different impacts than defoliation using scissors (Piene and Little 1990);

however, in the current experiment, the apparent reduction in epicormic buds in the

J. ENTOMOL. SOC. BRIT. COLUMBIA 114, DECEMBER 2017 fe

budworm-defoliated seedlings may have been due to the continued feeding activity of the

larvae. It was not possible to distinguish between sequential and epicormic buds once

they had been eaten, as individual buds were not tracked in detail in this experiment.

The larval collection sites north of Williams Lake, B.C., experienced 1,539—1,688

mean annual growing degree days, and 174—180 mean frost-free days from 1961-1990

(http://www.climatewna.com/ accessed 2017). Sites near Monte Creek, B.C., where

outbreaks have occurred more commonly, have experienced 1,910 mean annual growing

degree days and 207 mean frost-free days from 1961-1990 (http://www.climatewna.com/

accessed 2017). Both the number of growing degree days and the number of frost-free

days can vary substantially with location and year; in general, a shorter growing season

can be expected in the north and at higher elevations, with a greater risk of early fall

frosts, than in the south at lower elevations.

These results suggest that northern trees may experience greater growth losses and

potentially higher mortality during western spruce budworm outbreaks than might be

anticipated further south. Synchrony between larvae and their host trees is a key

component of the population dynamics of this insect; both the beginning and end of the

phenological window are important to survival and fecundity (Nealis 2012). The

availability of high nutritional—quality buds during later instars could improve the

survival and fecundity of larvae at the end of the phenological window (Régniére and

Nealis 2016). Depending on location and local weather conditions, however, the short

growing season in the north could also result in higher insect mortality in some years.

Further work is needed to investigate the interaction between western spruce

budworm larvae and epicormic buds at northern latitudes and at higher elevations.

Experiments that track individual buds, compare the effects of natural larval feeding to

bud removal later in the season, and examine the impacts on mature trees would all

improve understanding of defoliator impacts on northern stands. If field populations of

mature trees carry less new foliage late in the summer in the north than they do in the

south, the impact of a western spruce budworm outbreak could be more severe in the

northern part of the insect’s range.

REFERENCES

Alfaro, R. 1, G. A. Van Sickle, A. J. Thomson, and E. Wegwitz. 1982. Tree mortality and radial growth

losses caused by the western spruce budworm in a Douglas-fir stand in British Columbia. Canadian

Journal of Forest Research 12:780-787.

Maclauchlan, L. E., and J. E. Brooks. 2009. Influence of past forestry practices on western spruce

budworm defoliation and associated impacts in southern British Columbia. B.C. Journal of

Ecosystem Management 10:37—-49.

Maclauchlan, L. E., J. E. Brooks, and J. C. Hodge. 2006. Analysis of historic western spruce budworm

defoliation in south central British Columbia. Forest Ecology Management 226:351-—356.

Meier, A. R., M. R. Saunders, and C. H. Michler. 2012. Epicormic buds in trees: A review of bud

establishment, development and dormancy release. Tree Physiology 32(5):505—509.

Nealis, V. G. 2012. The phenological window for western spruce budworm: Seasonal decline in resource

quality. Agricultural and Forest Entomology. 14:340-347.

Piene, H. 1989. Spruce budworm defoliation and growth loss in young balsam fir: recovery of growth in

spaced stands. Canadian Journal of Forest Research 19:1616—1624.

Piene, H., and E. S. Eveleigh. 1996. Spruce budworm defoliation in young balsam fir: The “green” tree

phenomenon. The Canadian Entomologist 128:1101--1107.

Piene, H., and C. H. A. Little. 1990. Spruce budworm defoliation and growth loss in young balsam fir:

artificial defoliation of potted trees. Canadian Journal of Forest Research 20:902—909.

Pohlert, T. 2014. The Pairwise Multiple Comparison of Mean Ranks package (PMCMR). R Package.

http://CRAN.R-project.org/package=PMCMR.

R Development Core Team. 2016. R: A Language and Environment for Statistical Computing. R Found.

Stat. Comput. Vienna Austria. 0: {ISBN} 3-900051-07-0.

76 J. Entomol. Soc. Brit. Columbia 114, December 2017

Régniere, J., and V. G. Nealis. 2016. Two sides of a coin: host-plant synchrony fitness trade-offs in the

population dynamics of the western spruce budworm. Insect Science 1-10. DOT:

10.1111/1744-7917.12407.

J. ENTOMOL. SOC. BRIT. COLUMBIA 114, DECEMBER 2017 77

NATURAL HISTORY AND OBSERVATIONS

Archilestes californicus McLachlan (Odonata: Zygoptera:

Lestidae):a damselfly new to Canada

ROBERT A. CANNINGS! & RUSSELL V. PYM?

Archilestes californicus McLachlan (California Spreadwing) is a large damselfly

native to western North America, ranging from Washington and Idaho south to New

Mexico, Arizona and California and, in Mexico, to Sonora and Baja California Sur

(Paulson 2011; Westfall and May 2006). This note records the species for the first time

in Canada—from three sites in the southern Okanagan Valley, British Columbia (BC;

Figure 1).

Russell Pym saw several males and females at a small, shallow, artificial pond at the

end of an artificial stream near the entrance to the Liquidity Winery at 4720 Allendale

Road, Okanagan Falls, BC (49.32553°N, 119.54993°W). He observed them from 13:00

to 14:00 PDT on 26 September 2016; one male was photographed (Figure 2). From

16:30 to 17:00 PDT the same day, he recorded a female in knee-high grass, three to four

metres from the shore of a dugout pond across the road from Walnut Beach Resort, 4200

Lakeshore Drive, Osoyoos, BC (49.01825°N, 119.43580°W). Cattail (Typha latifolia)

and willows (Salix spp.) lined the pond margins.

At the north end of Vaseux Lake the next day, 27 September 2016, Russell

photographed a lone male (Figure 3) perched on cattails in a mixed willow swamp and

cattail marsh (13:00 to 14:30 PDT). The site was along the boardwalk to the bird blind at

49.30348°N, 119.53696°W.

Archilestes (Stream Spreadwings) is a New World genus of eight species; two are

North American, the others live from Mexico to Argentina (Paulson 2009). These

damselflies are larger than the related Lestes (Pond Spreadwings) species, which are

common and more familiar to Canadian observers.

Archilestes californicus is a large spreadwing (42—60 mm long) with eyes and labrum

blue in males. The thorax is metallic brown dorsally, white laterally on the

metepisternum and metepimeron, with a brown stripe on the metapleural suture dividing

the white areas. The resulting white stripes are good field marks. The pterostigmas are

white or tan. The abdomen is brown dorsally, slightly metallic and often with a green

tinge; segments 9-10 are pruinose white in males (Figures 2 & 3). Paraprocts are short

and parallel, visible from above. Females are coloured as males, but lack pruinosity; the

eyes are dull blue to brown; the ovipositor reaches the tip of segment 10.

The flight season is late; in Washington, adults fly from July to November (Paulson

2009). Manolis (2003) records that the breeding season in California is mainly in

September and October; this is probably the case in much of its range. Archilestes

californicus lives along small, slow, often intermittent streams and associated ponds.

River backwaters are also inhabited. Paulson (2009, 2013) notes that larvae often swim

in open water like little minnows and he believes that waters lacking fish are important to

this species. Adults mate and lay eggs where willows and alders line the shore. When not

breeding, they often leave the water, flying out into open woodland, fields, and sagebrush

grassland (Manolis 2003; Paulson 2009).

Individuals fly out from their perches in waterside shrubs to catch prey and return

quickly. When disturbed, they dart into dense vegetation (Kennedy 1915; Manolis 2003).

| Royal British Columbia Museum, 675 Belleville Street, Victoria, BC, Canada V8W 9W2;

rcannings@royalbcmuseum.bce.ca

2 1027 Scottswood Lane, Victoria, BC, Canada V8Y 2V1; rpym@shaw.ca

78 J. Entomol. Soc. Brit. Columbia 114, December 2017

Males often perch conspicuously on dead twigs, spreading their wings, defending small

territories (Paulson 2009). Pairs oviposit in tandem in willow or alder branches about 0.5

—1.0 cm thick, often up to 3 m above the water. The female inserts a group of six eggs

into the cambium, then backs down the stem briefly and repeats the process, laying up to

180 eggs per session. The eggs apparently overwinter before hatching (Kennedy 1915;

Manolis 2003; Paulson 2009).

Kennedy (1915) found Archilestes californicus abundant at Satus Creek and at other

locations in the Yakima Valley of south—central Washington in 1913 and noted that these

were the only records north of California at the time. He also corrected an earlier record

of A. grandis from Yakima that, as Paulson (1970) clarified, should be referred to A.

californicus. Since then, the species has been recorded at many localities in Oregon and

Washington. In Washington, west of the Cascade Mountains, first county records in the

OdonataCentral website (Abbott 2006-2016) roughly indicate a northern movement:

Clark County, 1997; Thurston County, 2009; King County, 2011. Jim Johnson (Abbott

2006-2016; pers. comm.) finds it commonly in Clark County near the mouth of the

Columbia River. Dennis Paulson (pers. comm.) says that “it really is moving north. It’s

common in parts of Seattle now, definitely consolidating its range in this state.” The

assumption that this is a natural range extension is complicated by the possibility that

some of the wetlands involved in Thurston and King counties are artificially constructed

wetlands surrounded by planted willows, in which the eggs of Archilestes may have been

introduced (Paulson 2013).

East of the Cascades, where the species was first reported in Washington, it is

recorded in Adams, Benton, Douglas, Grant, Kittitas, Okanogan, Whitman and Yakima

counties. There is a broad corridor of counties, from the Columbia River north to the

Canadian border, in which the species has been found. Most relevant to the new

Canadian records, Jim Johnson found it in the southwest corner of Okanogan County in a

pond along Black Creek Canyon (48.07006°N, 120.01917°W) on 1 September 2002

(Abbott 2006—2016). This is 114 km southwest of the Osoyoos, BC, site.

Based on this history, it is not surprising that A. californicus has finally appeared in

the Okanagan Valley in Canada. The Osoyoos locality is only 2 km north of the United

States border. The Vaseux Lake site is 37.2 km north of the Osoyoos site and 2.5 km

south of the Okanagan Falls locality. The number of sites reported and the significant

distances between them suggest that Archilestes probably lives at additional locations in

the area and may have been overlooked in the Canadian part of the valley for several

years, or at least long enough for it to expand northward more than 40 km from the

United States. Further observations will clarify the status of the damselfly in British

Columbia and Canada.

Archilestes grandis (Rambur) (Great Spreadwing) is the only other North American

species in the genus. It comes no closer to British Columbia than northern California,

ranging from California east to New England and extreme southwestern Ontario, and

south to Venezuela (Paulson 2009, 2011). It is larger than A. californicus; mature males

are darker overall and have more extensive green highlights. The pterostigmas are dark.

The pale lateral thoracic stripe is longer and usually yellow rather than white. The

paraprocts are divergent under the cerci, and are difficult to see from above (Paulson

2009).

Archilestes grandis has also been on the move—and for a long time. It was first

recorded and described from the southwestern states, but its spread to the east and

northeast is well documented: it was recorded in Ohio as early as 1931 and, in Canada, at

Windsor in 2002 (Pratt and Paiero 2004). Paulson (2011) postulates that this impressive

range expansion might have been assisted by the damselfly’s tolerance of poor-quality

streams, which makes it a successful competitor.

J. ENTOMOL. SOC. BRIT. COLUMBIA 114, DECEMBER 2017 79

Figure 1. Localities of Archilestes californicus in the southern Okanagan Valley, BC,

September 2016. See text for details. Yellow horizontal line represents the Canada—United

States boundary (49° N). Scale line = 5 km.

80 J. Entomol. Soc. Brit. Columbia 114, December 2017

Figure Zz: Male Archilestes californicus photographed by Russell Pym at Liquidity Winery,

Okanagan Falls, BC, 26 September 2016.

J. ENTOMOL. SOC. BRIT. COLUMBIA 114, DECEMBER 2017 81

Figure 3. Male Archilestes californicus photographed by Russell Pym at the north end of

Vaseux Lake, BC, 27 September 2016.

ACKNOWLEDGEMENTS

Thanks to Jim Johnson and Dennis Paulson, who supplied information on the present

status of Archilestes californicus in Washington. Syd Cannings and Dennis Paulson

commented on an early draft of the paper.

REFERENCES

Abbott, J. C. 2006-2016. OdonataCentral: An online resource for the distribution and identification of

Odonata. Available at http://www.odonatacentral.org. (Accessed 6 November 2016).

Kennedy, C. H. 1915. Notes on the life history and ecology of the dragonflies of Washington and

Oregon. Proceedings of the United States National Museum 49:259-345.

Manolis, T. 2003. Dragonflies and damselflies of California. California Natural History Guides.

University of California Press, Berkeley CA.

Paulson, D. R. 1970. A list of the Odonata of Washington with additions to and deletions from the state

list. The Pan-Pacific Entomologist 46:194—198.

Paulson, D. R. 2009. Dragonflies and damselflies of the West. Princeton University Press, Princeton NJ.

Paulson, D. R. 2011. Dragonflies and damselflies of the East. Princeton University Press, Princeton NJ.

82 J. Entomol. Soc. Brit. Columbia 114, December 2017

Paulson, D. R. 2013. An apparently introduced population of California Spreadwings (Archilestes

californicus). Argia 25(3):14—15.

Pratt, P. D., and S. M. Paiero. 2004. Archilestes grandis (Rambur) (Odonata: Lestidae), new to Canada.

Pages 11-12 in P. M. Catling, C. D. Jones and P. Pratt, eds., Ontario Odonata, vol. 4. Toronto

Entomologists’ Association, Toronto, Ontario, Canada.

Westfall, M. J., Jr, and M. L. May. 2006. Damselflies of North America. Scientific Publishers,

Gainesville, FL.

J. ENTOMOL. SOC. BRIT. COLUMBIA 114, DECEMBER 2017 83

NATURAL HISTORY AND OBSERVATIONS

Evidence of established brown marmorated stink bug

populations in British Columbia, Canada

PAUL K. ABRAM!, TRACY HUEPPELSHEUSER?’, SUSANNA

ACHEAMPONG?’, PEGGY CLARKE!, HUME DOUGLAS‘, AND

TARA D. GARIEPY>

ABSTRACT— We report four new detections of invasive agricultural pest

Halyomorpha halys (Stal) (Hempitera: Pentatomidae), the brown marmorated stink

bug, in the Lower Mainland and Okanagan Valley regions of British Columbia (BC),

Canada, in 2015 and 2016. These finds include two confirmed breeding populations,

as well as homeowner collections at the same residence in two consecutive years.

Preliminary comparisons of mitochondrial DNA haplotypes from these collections

suggest that H. halys populations in BC are the result of movement and spread of

existing populations in North America, likely from the Pacific Northwest USA.

The brown marmorated stink bug, Halyomorpha halys (Stal) (Hemiptera:

Pentatomidae), native to Asia, is a globally invasive pest with a broad host-plant range.

This species often causes economic losses to tree fruit, berries, vegetables, and

ornamental plants (reviewed in Rice et a/. 2014). The stink bug is also a nuisance pest for

homeowners when it seeks indoor overwintering sites in the fall and winter months.

Since the detection of new invasive populations, beginning more than 20 years ago, H.

halys has become broadly established in Europe and North America (reviewed in Haye ef

al. 2015), including most of the continental USA. (see http://www.stopbmsb.org/where-

is-bmsb/state-by-state/) and the Canadian provinces of Ontario (Fogain and Graff 2011;

Gariepy et al. 2014a) and Quebec (Jacques Brodeur, personal communication). In British

Columbia (BC), H. halys has been intercepted in shipments from Japan, Korea, and

China several times since 1993 (Fogain and Graff 2011; Gariepy ef al. 2014b), but

breeding populations were not detected.

From 2015 to 2016, we detected a total of 487 H. halys at four different sites in the

Lower Mainland and Okanagan Valley regions of BC (Table 1). Evidence of H. halys

reproduction (eggs and/or nymphs with adults) was found on host plants at one site in the

Lower Mainland (Chilliwack Mountain) and another in the Okanagan Valley. At the

Chilliwack (Rosedale) site, where the first detection was made in 2015, an increased

number of stink bugs returned to the same residence in the fall of 2016, indicating that a

breeding population is established in this area. An H. halys nymph and two adults were

found by two different residents in the Kitsilano neighborhood of Vancouver, BC.

Possible routes of brown marmorated stink bug invasion into BC include natural

spread of already-established populations in the Pacific Northwest of the USA (Oregon,

' Agriculture and Agri-Food Canada, Agassiz Research and Development Centre, 6947 Highway 7, P.O.

Box 1000, Agassiz, BC VOM 1A0; (604) 796-6075, paul.abram@canada.ca

2 British Columbia Ministry of Agriculture, Plant Health Unit, 1767 Angus Campbell Road, Abbotsford,

BC V3G 2M3

> British Columbia Ministry of Agriculture, Plant Health Unit, 1690 Powick Road, Kelowna, BC V1X

7G5

4 Agriculture and Agri-Food Canada, Ottawa Research and Development Centre, 960 Carling Avenue,

Ottawa, ON KIA 0C6

> Agriculture and Agri-Food Canada, London Research and Development Centre, 1391 Sandford Street,

London, ON N5V 4T3

J. Entomol. Soc. Brit. Columbia 114, December 2017

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J. ENTOMOL. SOc. BRIT. COLUMBIA 114, DECEMBER 2017 85

Washington), as well as accidental human-mediated translocation from Asia, Europe, or

other Canadian or American localities. To gather preliminary information on possible

invasion routes for this species in BC, we used COI haplotyping, which has previously

been used to identify possible sources of H. halys invasions in Canada, the USA, and

Europe (e.g., Gariepy et al. 2014b; Xu et al. 2014; Gariepy et al. 2015). We amplified

and sequenced a 658-base pair (bp) region of the mitochondrial Cytochrome C oxidase

subunit 1 (COI) gene from several specimens collected at each site (Table 1) to identify

COI haplotypes (see Gariepy et al. 2014b for primers and methodology). Previous

studies concluded that eastern North American H. halys populations originated from a

single source population in the Beijing area of China. In contrast, European populations

are derived from several Asian source populations, including China, Korea, and other

currently unidentified locations (Gariepy et al. 2015). Populations from the western USA

(California, Oregon, Washington) include multiple haplotypes, some of which differ from

those found in eastern North America (Haye eft al. 2015). Our genetic analysis of field-

collected specimens from BC supports these findings, demonstrating that multiple

haplotypes also occur in western Canada. In total, three COI haplotypes were detected

among the specimens collected in BC: Hl, H3, and a currently undescribed COI

haplotype (Hn) (Table 1). COI haplotypes H1 and H3 were described by Gariepy ef al.

(2014); H1 is the predominant haplotype in eastern North America and some areas of

Europe, whereas H3 is known from several regions in Europe, predominantly in

Switzerland (Gariepy eft al. 2015). Concurrent research in the western USA has employed

different gene regions for haplotype analysis (COII and 12S), but comparison of

representative datasets demonstrates that COI H1 and H3 haplotypes are already known

from Washington and Oregon (Marie Claude Bon and Kim Hoelmer, personal

communications). The third COI haplotype (Hn) has not previously been described from

samples collected in Asia, Canada, or Europe (Gariepy et al. 2014b; Gariepy et al. 2015).

This haplotype may occur in the Pacific Northwest of the USA; however, additional

DNA sequencing will be necessary to determine how it corresponds to the haplotypes for

the COII and 12S genes that have been analysed for H. halys specimens collected in

Washington, Oregon, and California (Marie Claude Bon and Kim Hoelmer, personal

communication).

Continuing public outreach will be important for tracking the spread of H. halys

populations in BC and for gathering specimens to examine population genetics and detect

invasion pathways. For example, a citizen discovered the Chilliwack Mountain

population following a BC Ministry of Agriculture newspaper advertisement, and the

Kitsilano reports were a result of residents seeing news articles reporting on the brown

marmorated stink bug. Additionally, just before the submission of this note, two

additional citizen reports of individual finds in the Lower Mainland (Langley, BC) and

the Interior (Kelowna, BC) were received (Hueppelsheuser and Acheampong,

unpublished data). To continue to track the spread and establishment of the brown

marmorated stink bug in BC, a citizen science approach is planned, to be complemented

by pheromone trapping and grid-based beating sheet sampling of known host plants.

ACKNOWLEDGEMENTS

Representative voucher specimens are deposited in the Canadian National Collection

of Insects, Arachnids, and Nematodes (Ottawa, Canada; CNC655203, CNC655204,

CNC655205). We thank Michael Schwartz for confirming initial identifications. We are

also thankful to Caitlin Miller, Emma Walker, Karlee Friesen, Sam Nigg, Tatiana

Percival, and Gabriella Zilahi-Balogh for field assistance, and Allison Bruin for technical

assistance. Thanks to Marie Claude Bon and Kim Hoelmer (USDA - ARS) for

86 J. Entomol. Soc. Brit. Columbia 114, December 2017

discussions and insight regarding H. halys population genetics in the USA, and all BC

homeowners who reported H. halys finds.

REFERENCES

Fogain, R., and S. Graff. 2011. First records of the invasive pest, Halyomorpha halys (Hemiptera:

Pentatomidae), in Ontario and Quebec. J. Entomol. Soc. Ontario 142:45—48. |

Gariepy, T. D., H. Fraser, and C. D. Scott-Dupree. 2014a. Brown marmorated stink bug (Hemiptera:

Pentatomidae) in Canada: recent establishment, occurrence, and pest status in southern Ontario. Can.

Entomol. 146:579-—582.

Gariepy, T. D., T. Haye, H. Fraser, and J. Zhang. 2014b. Occurrence, genetic diversity, and potential

pathways of entry of Halyomorpha halys in newly invaded areas of Canada and Switzerland. J. Pest

Sci. 87:17-28.

Gariepy, T. D., Bruin A., T. Haye, P. Milonas, and G. Vetek. 2015. Movement and spread of an invasive

pest: occurrence and genetic diversity of new populations of Halyomorpha halys in Europe. J. Pest

Sci. 88:451—460.

Haye, T., T. D. Gariepy, K. Hoelmer, J. P. Rossi, J. C. Streito, X. Tassus, and N. Desneux. 2015. Range

expansion of the invasive brown marmorated stinkbug, Halyomorpha halys: an increasing threat to

field, fruit and vegetable crops worldwide. J. Pest Sci. 88:665—673.

Rice, K. B., C. J. Bergh, E. J. Bergmann, D. J. Biddinger, C. Dieckhoff, G. Dively, H. Fraser, T. D.

Gariepy, G. Hamilton, T. Haye, A. Herbert, K. A. Hoelmer, C. R. Hooks, A. Jones, G. Krawezyk, T.

Kuhar, H. Martinson, W. Mitchell, A. L. Nielsen, D. G. Pfeiffer, M. Raupp, C. Rodrigues-Saona, P.

Shearer, P. Shrewsbury, P. D. Venugopal, J. Whalen, N. G. Wiman, T. C. Leskey, and J. F. Tooker.

2014. Biology, ecology, and management of brown marmorated stink bug (Hemiptera:

Pentatomidae). J. Integr. Pest Management 5:A1—A13.

Stop BMSB — Where is BMSB -— State by State. Specialty Crop Research Initiative, USDA. URL: http://

-1S- -by- [accessed 28 November 2016].

Xu, J., D. M. Fonseca, G. C. Hamilton, K. A. Hoelmer, and A. L. Nielsen. 2014. Tracing the origin of US

brown marmorated stink bugs, Halyomorpha halys. Biol. Invas. 16:153—166

J. ENTOMOL. SOC. BRIT. COLUMBIA 114, DECEMBER 2017 87

NATURAL HISTORY AND OBSERVATIONS

An unusual specimen of the subgenus Lasioglossum Curtis

from British Columbia, Canada (Hymenoptera, Halictidae)

CORY S. SHEFFIELD! and JENNIFER HERON

The genus Lasioglossum Curtis s. /. (Halictidae) represents one of the largest and

most taxonomically difficult groups of bees (Michener 2007; Gibbs 2010a), with at least

1,750 described species (Gibbs ef a/. 2012). In North America north of Mexico, six

subgenera are currently recognized: Dialictus Robertson, Evylaeus Robertson,

Hemihalictus Cockerell, Lasioglossum s. str., Sphecodogastra Ashmead, and_ the

introduced Leuchalictus Warncke. Globally, all subgenera have been placed within one of

two Lasioglossum “series” (Michener 2007; Gibbs ef a/. 2012). The Lasioglossum series

contains species in which only one (1.e., the third) submarginal crossvein (1.e., lrs-m) is

weakened, and is represented in North America by the subgenera Lasioglossum s. str. and

Leuchalictus. All other subgenera in North America normally have two weakened

submarginal crossveins and have been placed in the Hemihalictus series. However, weak

venation is not always perceptible in males and some female specimens of Lasioglossum

s. 1, and Ebmer (1969), Michener (2007) and Gibbs ef a/. (2013) stress the difficulty in

placing these problematic specimens within subgenera and even within series using these

diagnostic characters.

The problems with wing venation in Lasioglossum are not limited to these difficult

cases, as recently summarized by Gibbs (2010b). Hemihalictus, as originally defined by

Cockerell (1897), was monotypic, the species L. /ustrans (Cockerell) differing from all

other non-metallic Hemihalictus series Lasioglossum in having only two submarginal

cells; Cockerell (1897) used this character to separate his H. /ustrans from all other

Halictus Latreille. However, Gibbs (2010b) and Gibbs ef al. (2013) showed that a small

proportion of specimens of L. /ustrans have three submarginal cells. Similarly, Dialictus

was originally defined (Robertson 1902a) as monotypic and included one metallic

species with two submarginal cells, L. anomalum (Robertson); Chloralictus was used to

distinguish metallic species with three submarginal cells with two weakened submarginal

crossveins (Robertson 1902b). Gibbs (2010b) indicated that L. anomalum is also known

to have individuals with three submarginal cells. Therefore, wing venation alone is not

always reliable for species- or subgenus-level identification in Lasioglossum s. 1. In fact,

Stephen eft al. (1969) felt that these differences were so minimal that they considered

many of the taxa currently recognized as subgenera of Lasioglossum to be subgenera of

Halictus.

Both Hemihalictus and Dialictus are now defined much more broadly than these

historic usages. Based on phylogenetic data, Gibbs ef a/. (2013) placed many species of

Evylaeus (1.e., the non-metallic carinaless species, or non-metallic Dialictus) into the

subgenus Hemihalictus; thus, the subgenus is no longer considered monotypic (as per

Michener 2007). Mitchell (1960) was the first to define Dialictus (at genus level) as all

metallic Halictinae with two submarginal cells or two weakened submarginal crossveins.

Michener et al. (1994) were among the first to consider these as subgenera of

Lasioglossum. To this date, two submarginal-celled forms are known from only the

1 Royal Saskatchewan Museum, 2340 Albert Street, Regina, Saskatchewan, Canada S4P 2V7;

cory.sheffield@gov.sk.ca

* B.C. Ministry of Environment, Species Conservation Science Unit, University of British Columbia,

315-2202 Main Mall, Vancouver, British Columbia, Canada V6T 1Z1

88 J. Entomol. Soc. Brit. Columbia 114, December 2017

Hemihalictus series of Lasioglossum, specifically in the subgenera Hemihalictus and

Dialictus.

Our objective here is to describe an aberrant specimen of Lasioglossum s. str. from

British Columbia, Canada; it therefore represents the first documented case of a two-

submarginal celled individual within the Lasioglossum series. As part of ongoing work

on bee diversity and taxonomy in Canada, specimens were collected throughout the

Western Interior Basin Ecozone (=Southern Interior Ecoprovince) of British Columbia.

This area is considered the most bee species—rich in the country, with more than 50% of

Canada’s bee species known from this relatively small area (5.7 million ha), over 1/3 of

which have not been recorded elsewhere in the country (Sheffield et a/. 2014; Heron and

Sheffield 2016). Among the specimens collected from this area between 2009 and 2016,

one male specimen of Lasioglossum collected on Mt. Kobau, South Okanagan

Grasslands Protected Area, near Osoyoos [49.106, —119.651, 08 Aug 2014, Col. C.

Sheffield] was morphologically unique in being a non-metallic Lasioglossum s. 1. with

two submarginal cells (Figure 1), thus superficially resembling L. /ustrans of eastern

North America, although with antenna resembling Lasioglossum s. str. (McGinley 1986).

Since the “two submarginal cell” condition of L. Justrans and L. anomalum is not always

consistent (Gibbs 2010b) and because no Halictinae with two-submarginal cells in the

forewing have been previously recorded from western North America (Stephen ef al.

1969), including western Canada (Gibbs 2010b), DNA barcoding was used to compare

sequences from the specimen (BOLD Sample ID CCDB-20945 F01) to other specimens

from western Canada (following methods of Sheffield et al. 2009, 2017). Despite

examining thousands of specimens from the Western Interior Basin, including from the

Mt. Kobau area of British Columbia, no additional specimens of Lasioglossum with two

submarginal cells could be found. The specimen in question (Figure 1) was identified as

a member of the subgenus Lasioglossum s. str. due to the characteristic basal antennal

structure (Figure 2; proportion of length of flagellomere 1 to 2), and tentatively as L.

(Lasioglossum) sisymbrii (Cockerell) largely due to the characteristic pale, translucent

tegulae (Figure 1) and by dissection and examination of the genitalia (Figure 3),

including sternum 8 (Figure 4) (McGinley 1986); this identification was supported

through comparison of COI sequences, matching identically with material in BOLD

identified as this species (see Sheffield et al. 2017). Although both sexes of this species

typically have a complete basal hair band on tergum 1 (McGinley 1986), this was

probably worn on our specimen. Our specimen also differed in having rather pale tarsi

(Figure 1)}—not concolorous with the tibiae (McGinley 1986).

A puzzling feature of our male specimen is the 12-segmented antennae (1.e., 10

flagellomeres; Figure 2)—the normal condition for female bees. Males of most bees have

13-segmented antennae, although males of Cherogas (Halictidae: Augochlorini) and a

few other genera are 12-segmented (Michener 2007; Engel 2010). One possibility is that

the specimen is a gynandromorph, although this condition has not been reported

previously for this species (see Wcislo et al. 2004; Michez et al. 2009; Hinojosa-Diaz et

al. 2012). If the specimen is indeed a gynandromorph, this condition seems restricted

only to the antennae, and the specimen otherwise resembles a male. Wcislo et a/. (2004),

Michez et al. (2009), and Hinojosa-Diaz et al. (2012) have listed bee species known as

gynandromorhps, the most recent study indicating that 1t has only been observed in 21

species of Halictidae, eight of which are Lasioglossum s. 1. The other possibility is that

this condition represents a development anomaly with this specimen.

This specimen of L. sisymbrii becomes the first account of a member of the

Lasioglossum series having two submarginal cells, and perhaps the first reported case of

gynandromorphy in a North American Lasioglossum. As with the other published works

cited above, we feel that documenting such anomalous specimens is important to account

for the variation that exists within species and, as discussed above, variable wing

venation has had an important impact on nomenclatural history for sweat bees in the past.

J. ENTOMOL. SOC. BRIT. COLUMBIA 114, DECEMBER 2017 89

Figure 1. Lateral habitus of male Lasioglossum sisymbrii (Cockerell), with two submarginal

cells in each forewing (numbered for the left forewing).

90 J. Entomol. Soc. Brit. Columbia 114, December 2017

Figure 2. Face of male Lasioglossum sisymbrii (Cockerell) showing the 12-segmented

antennae (i.e., 10 flagellomeres; the normal condition in males is 11 flagellomeres).

Figure 3. Genitalia Lasioglossum sisymbrii (Cockerell) — A) dorsal, B) ventral, and C) lateral

views.

J. ENTOMOL. SOC. BRIT. COLUMBIA 114, DECEMBER 2017 91

Figure 4. Sternum 8 of Lasioglossum sisymbrii (Cockerell).

ACKNOWLEDGEMENTS

Laurence Packer, York University, provided helpful comments on an early draft of the

manuscript, and two anonymous reviewers for constructive suggestions. Dave Fraser and

Orville Dyer provided local information. Orville Dyer, Dawn Marks, Cara Dawson, Sara

Bunge, Mark Weston and Brenda Costanzo assisted with surveys. Funding for surveys

provided by B.C. Ministry of Environment, B.C. Ministry of Forests, Lands and Natural

Resource Operations and federal Habitat Stewardship Program — Prevention Stream. The

specimen has been deposited in the Royal Saskatchewan Museum.

REFERENCES

Cockerell, T. D. A. 1897. On the generic position of some bees hitherto referred to Panurgus and

Calliopsis. The Canadian Entomologist 29:287—290.

Ebmer, A. W. 1969. Die Bienen des Genus Halictus Latr. s./. im Grossraum von Linz (Hymenoptera,

Apidae). Systematik, Biogeographie, Okologie und Biologie mit Beriicksichtigung aller bisher aus

Mitteleuropa bekannten Arten. Teil I. Naturkundliches Jahrbuch der Stadt Linz 15:133-—183. [in

German]

Engel, M. S. 2010. Revision of the bee genus Chilerogella (Hymenoptera, Halictidae), Part II: South

American Species and Generic Diagnosis. ZooKeys 47:1—100.

Gibbs, J., S. Brady, K. Kanda, and B. N. Danforth. 2012. Phylogeny of halictine bees supports a shared

origin of eusociality for Halictus and Laslogionsan (Apoidea: Anmhophita: Halictidae). Molecular

Phylogenetics and Evolution 65:926—939. http:// ij |

Gibbs, J., L. Packer, S. Dumesh, and B. N. Danforth. 2013. Revision and reclassification of

Lasioglossum (Evylaeus), L. (Hemihalictus) and L. (Sphecodogastra) in eastern North America

(Hymenoptera: Apoidea: Halictidae). Zootaxa 3672:1—117.

Gibbs, J. 2010a. Revision of the metallic species of Lasioglossum (Dialictus) in Canada (Hymenoptera,

Halictidae, Halictini). Zootaxa 2591:1—382.

Gibbs, J. 2010b. Atypical wing venation in Dialictus and Hemihalictus and its implications for

subgeneric classification of Lasioglossum. Psyche (605390):1—6. http://dx.doLorg/

10.1155/2010/605390

Heron, J. M., and C. S. Sheffield. 2016. First record of the Lasioglossum (Dialictus) petrellum species

group in Canada. Journal of the Entomological Society of British Columbia 112:88—91.

Hinojosa—Diaz, I. A., V. H. Gonzalez, R. Ayala, J. Mérida, P. Sagot, and M. S. Engel. 2012. New orchid

and leaf-cutter bee gynandromorphs, with an updated review (Hymenoptera, Apoidea).

Zoosystematics and Evolution 88:205—214.

92 J. Entomol. Soc. Brit. Columbia 114, December 2017

McGinley, R. J. 1986. Studies of Halictinae (Apoidea: Halictidae), I: Revision of New World

Lasioglossum Curtis. Smithsonian Contributions to Zoology 429:1-294.

Michener, C. D. 2007. The Bees of the World. Second Edition. Johns Hopkins University Press,

Baltimore, MD.

Michener, C. D., R. J. McGinley, and B. N. Danforth. 1994. The Bee Genera of North and Central

America (Hymenoptera: Apoidea). Smithsonian Institution Press, Washington, D.C.

Michez, D., P. Rasmont, M. Terzo, and N. J. Vereecken. 2009. A synthesis of gynandromorphy among

wild bees (Hymenoptera: Apoidea), with an annotated description of several new cases. Annales de

la Société entomologique de France 45:365-375.

Mitchell, T. B. 1960. Bees of the Eastern United States: Volume I. North Carolina Agricultural

Experimental Station Technical Bulletin 141:1-538.

Robertson, C. 1902a. Some new or little-known bees—II. The Canadian Entomologist 34:48-49.

Robertson, C. 1902b. Synopsis of Halictinae. The Canadian Entomologist 34:243—250.

Sheffield, C. S., J. Heron, J. Gibbs, T. M. Onuferko, R. Oram, L. Best, N. deSilva, S. Dumesh, A. Pindar,

and G. Rowe. 2017. Contribution of DNA barcoding to the study of the bees (Hymenoptera:

Apoidea) of Canada: progress to date. The Canadian Entomologist https://doi.org/10.4039/tce.

2017.49.

Sheffield, C. S., S. D. Frier, and S. Dumesh. 2014. The bees (Hymenoptera: Apoidea, Apiformes) of the

Prairies Ecozone, with comparisons to other grasslands of Canada. Arthropods of Canadian

Grasslands 4:427—-467.

Sheffield, C. S., P. D. N. Hebert, P. G. Kevan, and L. Packer. 2009. DNA barcoding a regional bee

(Hymenoptera: Apoidea) fauna and its potential for ecological studies. Molecular Ecology Resources

9(s1):196—207.

Stephen, W. P., G. E. Bohart, and P. F. Torchio. 1969. The Biology and External Morphology of Bees

with a Synopsis of the Genera of Northwestern America. Agricultural Experiment Station, Oregon

State University, Corvallis, OR.

Weislo, W. T., V. H. Gonzalez, and L. Arneson. 2004. A review of deviant phenotypes in bees in relation

to brood parasitism, and a gynandromorph of Megalopta genalis (Hymenoptera: Halictidae). Journal

of Natural History 38:1443—1457.

J. ENTOMOL. SOc. BRIT. COLUMBIA 114, DECEMBER 2017 93

NATURAL HISTORY AND OBSERVATIONS

First identifications of aphid and diamondback moth

populations on wasabi in British Columbia

JESSE L. MACDONALD!”, ERIC MAW’, and PEGGY CLARKE*

ABSTRACT

Wasabi is a highly valued crop in the Pacific Northwest where commercial

production is increasing. To date, little attention has been paid to its invertebrate

pests. Two wasabi polyhouses in Agassiz, BC, were monitored for insect pests for

15 months. Pemphigus populitransversus Riley (poplar petiole gall aphid)

recurred annually in winter months on roots throughout the polyhouses. Lipaphis

pseudobrassicae Davis (turnip aphid) infested the leaves of a large number of

plants. Myzus persicae Sulzer (green peach aphid) and Macrosiphum euphorbiae

Thomas (potato aphid) were noted in very low numbers. Plutella xylostella

Linnaeus (diamondback moth) caused shot-hole damage of the leaves. Further

investigation into the role of insects as vectors and their role in pathogen

pathways on this unique crop is needed.

Key words: wasabi, Pemphigus populitransversus, Lipaphis pseudobrassicae,

Plutella xylostella

INTRODUCTION

Wasabi (Wasabia japonica (Miq.) Matsumura) (Brassicaceae) is native to Japan,

where it grows in shaded stream environments (Adachi 1987). It is currently cultivated in

Asia, Australasia, and North America for its valuable rhizome, which is used as a freshly-

ground condiment eaten with traditional Japanese meals (Hodge 1974; Chadwick et al.

1993). It can fetch US$150-300/kg on the international market. Although the rhizome 1s

the primary plant part for culinary use, the leaves can also be used to flavour soups or

salads (Chadwick ef al. 1993). In B.C., there is an estimated 5-10 acres of commercial

wasabi in production using hydroponic or similar systems in polyethylene tunnels

(polyhouses) or traditional glass greenhouses. Plants are typically grown in river rock

substrate, as plants grown in soil are thought to produce an inferior quality rhizome

(Chadwick 1993; Sultana et al. 2003). It takes 12 — 18 months before plants are of

marketable quality, and due to the humid growing environment and propagation from

axillary shoots, disease issues are the most common reason for crop loss in British

Columbia (Rodriguez & Punja 2009; Punja et al. 2017; MacDonald & Punja 2017). To

date, little research has been directed toward arthropod pests, and all reports are

anecdotal. We report the first occurrence of insect pests on wasabi in North America at a

research planting in Agassiz, British Columbia.

| Agriculture & Agri-Food Canada, Pest Management Centre, 4200 Highway 97, Summerland, BC VOH

1Z0, Canada, Jesse. MacDonald@agr.gc.ca

* Simon Fraser University, Department of Biological Sciences, 8888 University Drive, Burnaby, BC V5A

1S6, Canada

3 Agriculture & Agri-Food Canada, Canadian National Collection of Insects, Arachnids and Nematodes,

CEF, 960 Carling Ave, Ottawa, ON K1A 0C6, Eric. Maw@agr.gc.ca

4 Agriculture & Agri-Food Canada, Agassiz Research & Development Centre, 6947 Highway 7, Agassiz,

BC VOM 1A0, Canada

94 J. Entomol. Soc. Brit. Columbia 114, December 2017

MATERIALS AND METHODS

Two polyhouses, each planted with 500 tissue-cultured (cv. Daruma) and 500

auxiliary-shoot propagated (cv. Mazuma) wasabi plantlets, were established in January

2015 at the Agriculture and Agri-Food Canada (AAFC) Agassiz Research and

Development Centre in Agassiz, BC. Prior to transplant into the polyhouses, the plants

were maintained in a production greenhouse for one month. A commercial nutrient

growing system was used, with overhead misters fertigating at regular intervals or when

triggered by a photosensor. Plants were grown in ~20 cm of 2-3 cm diameter river-rock.

From April to October, 70% shade cloth covered the polyhouses to reduce direct

exposure to UV radiation. Weekly or bi-weekly inspections were conducted by trained

staff to identify pests and for treatment recommendations. Aphid populations were

treated with imidacloprid. Two releases of Diadegma insulare Cresson parasitoids were

conducted weekly to manage feeding diamondback moth larvae, followed by treatment

with flubendiamide.

Leaves, petioles, and roots were visually inspected at random throughout each

planting. Approximately five infested plants with representative pest populations from

either leaves and petioles or roots were selected for each sample. A soft paintbrush was

used to gently brush specimens off of plant material into vials of 95% ethanol (EtOH) for

identification. Aphids were identified by morphological determination or by sequencing

mitochondrial cytochrome C oxidase, subunit 1 (“DNA barcoding”). Adult diamondback

moths were identified by morphology under a binocular microscope. Assessments were

carried out until harvest, after 15 months.

RESULTS AND DISCUSSION

Poplar petiole gall aphids. In January 2015, W. japonica ‘Daruma’ plantlets grown

in soil-less plugs were uprooted for transplant into polyhouses and a heavy root aphid

infestation was noted. Aphids were present throughout the crop and each root system,

and a characteristic white waxy secretion was visible (Fig. 1A). No alates were present.

The following January to March (2016) on the same crop of Daruma and a neighbouring

crop of Mazuma, identical root aphid populations were again found. In both cases

treatment with imidacloprid appeared to provide control. All populations were identified

as Pemphigus populitransversus Riley by sequence matching to specimens collected

from galls on Populus deltoides (Fig. 1A, B).

Root aphids have been implicated as pests of wasabi historically (Miles & Chadwick

2008; Chadwick et al. 1993) but identified only once, in New Zealand, as P. bursarius

Linnaeus (Douglas & Follett 1992); that population was difficult to control. Pemphigus

populitransversus is known to alternate hosts between roots of various Brassicaceae,

sometimes as a significant pest (for example Chen ef al. 2009), persisting by

parthenogenesis, and a sexual stage on Populus spp. trees, where they overwinter as eggs

and form galls on the petioles of the leaves the following spring (Jones & Gillette 1918).

Aphid damage to roots and rhizomes may be an important pathway for pathogens such as

Pectobacterium carotovorum subsp. carotovorum, which has been found to cause

vascular blackening of the rhizome after entry through small wounds (Rodriguez & Punja

2009). This is the first published report of P populitransversus in B.C. that the authors

are aware of, although there are specimens of unidentified Pemphigus species from

wasabi collected in Aldergrove and Langley in 1997 and 1998 in the Canadian National

Collection of Insects, Arachnids and Nematodes.

Turnip aphids. In spring of 2016, aphids were found predominantly on leaves of

one-quarter of the affected polyhouse and identified as Lipaphis pseudobrassicae Davis

(n = 62 specimens). Macrosiphum euphorbiae Thomas (n = 2) and Myzus persicae Sulzer

(n = 1) were also present in the sample.

J. ENTOMOL. SOC. BRIT. COLUMBIA 114, DECEMBER 2017 95

Figure 1. A) Parthenogenic P. populitransversus population with waxy exudate in the root

mass of a W. japonica plant grown in a plug-tray. B) Apterous P populitransversus with

proboscis in W. japonica root. C) L. pseudobrassicae colony consisting of different instars on

a W. japonica leaf. D) M. euphorbiae aptera with proboscis in a W. japonica leaf. Photos by

by the Minister of Agriculture and Agri-Food 2016.

The most serious issue associated with aphids on wasabi is their ability to transmit

viruses (Douglas & Follett 1992). Wasabi is susceptible to tobacco mosaic virus (TMV),

turnip mosaic virus (TuMV), and cucumber mosaic virus (CMV) (Chadwick et al. 1993;

Wilson 1998), and although problematic elsewhere, no viruses have been identified on

wasabi in B.C. Should these diseases be reported locally, L. pseudobrassicae should be

assessed as a potential vector of TuMV and CMV (Chan et a/. 1991).

Diamondback moth. In June 2015, a heavy infestation of diamondback moth,

Plutella xylostella Linnaeus, and associated ‘shothole’ damage on leaves was found.

Adults were prevalent and flew as plants were disturbed.

Diamondback moth is the most destructive pest of Brassica crops worldwide. It has

been reported on wasabi crops in Japan (Hodge 1974; Adachi 1987; Chadwick ef al.

1993; Miles & Chadwick 2008). Due to successful management of the infestation with

flubendiamide, it is unclear whether D. insulare releases were effective. Interestingly, a

single mobile parasitoid adult was photographed almost 10 months later in a

96 J. Entomol. Soc. Brit. Columbia 114, December 2017

neighbouring polyhouse which had no previous biocontrol releases, suggesting the

population persisted.

This first survey of insect pests of commercial wasabi production suggests that there

is considerable potential for economic damage. Currently, no insecticides are registered

in the United States for use on wasabi and only one synthetic insecticide is registered in

Canada (permethrin). Although there are a number of biopesticides available in Canada,

these may not be sufficient if the aphids are vectors for viruses. Investigations into the

relationship between aphids and pathogens (such as P. carotovorum), or as vectors of

viruses, may generate interest in an integrated management approach, as well as the

registration of additional control products for resistance management for use in

commercial wasabi crops.

ACKNOWLEDGEMENTS

We thank staff at the Agriculture and Agri-Food Canada Research & Development

Centre in Agassiz: James Nicholson and Seth Nussbaum for maintenance and surveying

of research plots, and Markus Clodius and Dave Gillespie for advice. We also thank Tom

Lowery and Howard Thistlewood at Summerland Research & Development Centre for

providing comments on the manuscript.

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Chadwick CI, Lumpkin TA, Elberson LR. 1993. The Botany, Uses and Production of Wasabia japonica

(Mig.) (Cruciferae) Matsum. Econ Bot 47(2): 113-135. DOI:10.1007/BF02862015

Chan CK, Forbes AR, Raworth DA. 1991. Aphid-transmitted viruses and their vectors of the world. Agri

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Chen N, Liu T-X, Sétamou M, French JV, Louzada ES. 2009. Molecular identification and population

dynamics of two species of Pemphigus (Homoptera: Pemphidae) on cabbage. Insect Science 16:

115-124, DOI:10.1111/4.1744-7917.2009.00262.x

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BF02861977

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MacDonald JL, Punja ZK. 2017. Occurence of botrytis leaf blight, anthracnose leaf spot, and white

blister rust on Wasabia japonica in British Columbia. Can J Plant Pathol. 39(1): 60-71. DOI:

10.1080/07060661.2017.1304021

Miles C, Chadwick C. 2008. Growing Wasabi in the Pacific Northwest. Farming the Northwest. WSU.

PNW0605: 1-12.

Punja ZK, Chandanie WA, Chen X, Rodriguez G. 2017. Phoma leaf spot of wasabi (Wasabia japonica)

caused by Leptosphaeria biglobosa. Plant Pathol. 66(3): 480-489. DOI:10.1111/ppa.12589

Rodriguez G, Punja ZK. 2009. Vascular blackening of wasabi rhizomes caused by Pectobacterium

carotovorum subsp. carotovorum. Eur J Plant Pathol. 124(3): 483-493. DOI:10.1007/

$10658-009-9435-1

Sultana T, Porter NG, Savage GP, McNeil DL. 2003. Comparison of Isothiocyanate Yield from Wasabi

Rhizome Tissues Grown in Soil or Water. J Agri Food Chem. 51(12): 3586-3591. DOI:10.1021/

jf021116c

Wilson CR. 1998. First report of cucumber mosaic cucumovirus on Wasabi in Australia. Plant Dis. 82(5):

590. DOI:10.1094/PDIS.1998.82.5.590A

J. ENTOMOL. SOC. BRIT. COLUMBIA 114, DECEMBER 2017 97

Proceedings of the Pollination: Science and Stewardship

Symposium

University of British Columbia - Okanagan Campus, Kelowna,

British Columbia, March 20, 2017

INTRODUCTION

Jennifer Heron! and Cory S. Sheffield’

Assessing the threats to and the conservation status of pollinators is emerging as one

of the greatest challenges facing conservation practitioners today. The diversity of

pollinator taxa and their cumulative contributions to natural ecosystem health and human

well-being involve complex, albeit often poorly understood, relationships. Increased

concern about the plight of pollinators has resulted in increased funding for education

and research on these topics, which is strengthening science-based policy and increased

public awareness. A symposium was held on March 20, 2017, at the University of British

Columbia, Okanagan Campus, in Kelowna, on pollinator science and stewardship. The

symposium brought together eight speakers who discussed topics related to pollinator

conservation, providing examples and case studies of conservation assessment, public

engagement, pollinator policy, and ideas regarding how to address the challenges that are

faced by pollinators and pollinator-stewardship practitioners. The symposium also

facilitated connections that enable lands managers, owners, stewards and conservation

practitioners to take this information and apply it to their own conservation practices.

Support for the symposium was provided by the federal Habitat Stewardship Program

for the Prevention of Species At Risk, the B.C. Ministry of Environment and Climate

Change Strategy, the Royal Saskatchewan Museum, the British Columbia Conservation

Foundation and the Entomological Society of British Columbia.

The buzz on Yukon bees

Syd Cannings, Environment and Climate Change Canada, Canadian Wildlife Service,

Whitehorse, YT Y1A 5X7

Amid the growing concern for the fate of bees, | have begun several studies on bees

in northern British Columbia and the Yukon, collaborating with Paul Williams at the

Natural History Museum (UK) and Cory Sheffield (Royal Saskatchewan Museum). Over

the past six years, these studies have revolved around focused collecting with nets and

traps in the various ecosystems of the north.

In general, bumblebee species that have declined dramatically in the south (e.g., the

Western Bumblebee [Bombus occidentalis occidentalis] and Yellow-banded Bumblebee

[Bombus terricola]) are still common in the southern Yukon. However, the Gypsy

Cuckoo Bumblebee (Bombus bohemicus; assessed Endangered in Canada by the

Committee on the Status of Endangered Wildlife in Canada [COSEWIC]) seems to be

much sparser than it was in the 1980s. We have found this species in two localities:

Stewart Crossing in 2014 and Kluane in 2016. These are the only detections of this

species in North America in the past five years.

Mitochondrial DNA analysis of some of the bumblebees we’ve collected has revealed

a new species of subarctic bumblebee in the subgenus Alpinobombus, now named

Bombus kluanensis. Even though we now have a better idea of the status of most

northern bees, we do not have good data on ongoing trends. To tackle that issue, we are

! B.C, Ministry of Environment and Climate Change Strategy, Species Conservation Science Unit,

Vancouver, B.C., V3R 1E1

2 Royal Saskatchewan Museum, Regina, SK, S4P 2V7

98 J. Entomol. Soc. Brit. Columbia 114, December 2017

planning to institute a repeatable monitoring plan for bumblebees in the Yukon and

northern B.C., modeled after the North American Breeding Bird Survey.

Honeybees and honeybee health

Rob W. Currie, University of Manitoba, Faculty of Agricultural and Food Sciences,

Department of Entomology, Winnipeg, MB R3T 2N2

Honey bees (Apis melanifera) have been experiencing high levels of colony loss on a

regular basis over the past decade. While speculation originally centered on the idea that

there was a single mysterious cause, we now know that multiple stressors are interacting,

sometimes in unpredictable ways to cause problems for this critically important crop

pollinator. Exciting progress is being made on high- and low tech-solutions to help

mitigate these losses, and some of these research innovations include using molecular

and proteomic markers, as well as conventional approaches to breed bees for resistance

to parasites and pathogens. Managing viruses through more effective management of

their primary vector, the varroa mite, and using RNAi to control viruses also can be

effective in helping beekeepers mitigate losses from some of the more critical stressors in

the system.

COSEWIC and the General Status of Species in Canada

David F. Fraser, British Columbia Ministry of Environment and Climate Change

Strategy, Species Conservation Science Unit, Victoria, B.C. V8W 1M&

The status of species at risk in Canada is assessed by the Committee on the Status of

Endangered Wildlife in Canada (COSEWIC). COSEWIC recommends species in the at-

risk category to the federal Minister of Environment and Climate Change for listing

under the federal Species At Risk Act (SARA). To date, there have been 976 wildlife

species assessed by COSEWIC, and 521 of these are listed on Schedule | of SARA. The

Program on the General Status of Species in Canada provides overview of the status of

biodiversity in Canada every five years and bears no legal implication. In 2013, this

program moved to using the same assessment system as used by NatureServe and the

B.C. Conservation Data Centre. The latest General Status report, covering the 2000-2015

timespan, assessed 29,848 species. The COSEWIC assessment process requires extensive

time and resources, and prioritizing which species to recommend for assessment is a

challenging task. Results from the General Status are one of the inputs that helps guide

the determination of which species are priorities. In addition, other factors, such as the

percent of the species range in Canada, the species global status, and the pattern of

decline, is used by COSEWIC to modify the priority score a species is given. A thorough

understanding of both the General Status and COSEWIC processes is important for

prioritizing species recommended for status assessment.

Butterfly conservation in Canada: threats and challenges

Jennifer M. Heron, British Columbia Ministry of Environment and Climate Change

Strategy , Species Conservation Science Unit, Vancouver, B.C. V3R 1E1

Butterflies are a well-known and well-studied group of pollinators. Approximately

275 butterfly species are known to occur in Canada, although only 21 have been assessed

nationally by the Committee on the Status of Endangered Wildlife in Canada

(COSEWIC). There are numerous challenges to assessing the lesser known and poorly

documented butterfly species, particularly when the host plant(s) are unknown, the

threats are unclear, and the species’ habitat and associated plant communities are

undescribed. Museum collections are important sources for historical information and, at

one time, butterfly specimens were more widely collected and deposited at museums.

However, historical collection data are often biased, not databased, or incorrectly

identified. In the past few decades, butterfly surveys have moved away from specimen

collection and museum deposition, and instead focus on visual surveys or photographic

evidence—a method that is good for conserving populations but has other drawbacks.

J. ENTOMOL. SOC. BRIT. COLUMBIA 114, DECEMBER 2017 99

Using examples from the south Okanagan, this talk will provide an overview of the

challenges to assessing butterflies, how candidate species are recommended for

COSEWIC assessment, challenges to assessing the lesser known species, and ways

conservation practitioners can include butterflies in land management decisions and

planning.

Border Free Bees: artists linking science and communities for pollinator

conservation

Nancy Holmes and Fionncara MacEoin, The University of British Columbia, Okanagan

Campus, Faculty of Creative and Critical Studies, Kelowna, B.C. V1V 1V7

Border Free Bees (BFB) is a Social Sciences and Humanities Research Council

(federal) funded provincial initiative in which artists lead community engagement

projects to enhance awareness and inspire action around pollinator conservation. Along

with several projects in the Lower Mainland, BFB has two major projects underway in

Kelowna—tThe Public Art Pollinator Pasture and the Kelowna Nectar Trail. Designed to

address the decline in native habitat, BFB is an ambitious and creative pollinator-focused

arts-based research initiative, headed by Dr. Cameron Cartiere, Associate Professor at

Emily Carr University of Art + Design (ECUAD) and Nancy Holmes, Associate

Professor in Creative Studies at The University of British Columbia, Okanagan (UBCO).

The research project’s mission includes raising awareness of the plight of wild

pollinators, particularly bees, and transforming underused urban sites into aesthetically

pleasing and scientifically viable habitats. Border Free Bees uses public art and design

methodologies to empower communities to actively engage in these restoration initiatives

and equips individuals and communities with the knowledge and tools to take

stewardship of such public projects.

From personal to planetary: making an impact on pollination at different scale

Hien T. Ngo, IPBES Secretariat, Intergovernmental Platform on Biodiversity and

Ecosystem Services (IPBES), UN Campus, Platz der Vereinten Nationen 1, D-53113,

Bonn, Germany

Research and initiatives that focus on pollination have impacts on different scales.

With the International Pollinator Initiative (UN-FAO), local researchers worked with

small-scale farmers using a common method to examine pollination deficits. This was

repeated in multiple countries in multiple regions around the world, resulting in a scaled-

up key finding regarding the role of wild pollinators in agroecosystems. Recently, the

Intergovernmental Platform on Biodiversity and Ecosystem Services (IPBES) completed

their Summary for Policymakers and assessment report on Pollinators, Pollination and

Food Production. These key findings, which included policy options, were adopted

(Decision XIII/15) at the Convention on Biological Diversity Thirteenth Conference of

the Parties (COP-13). Furthermore, these key findings have already had an impact on

many national pollinator strategies and were the basis of a new multinational initiative,

the Coalition of the Willing on Pollinators.

Pesticides and pollinators: evidence, controversy and policy

Nigel Raine, University of Guelph, School of Environmental Sciences, Rebanks Family

Chair in Pollinator Conservation, Guelph, ON NIG 2W1

Recent concern over global pollinator declines has led to considerable research on

pesticide impacts. Here, we report on a series of recent studies that examine the extent to

which field-realistic insecticide exposure can lead to significant sublethal impacts on

individual bumblebee behaviour (e.g., reduced queen colony founding success and

impaired worker learning and foraging), colony function (e.g., effects on growth rates

and forager recruitment), and the critical ecosystem services they provide to crops and

wild plants. Taken together, these effects could have widespread implications for the

stability of wild pollinator populations, sustainable production of pollinator-limited

100 J. Entomol. Soc. Brit. Columbia 114, December 2017

crops, and maintaining wild-plant biodiversity. Considering these studies that report

insecticide impacts on non-Apis (honey) bees into the wider context, particularly

alongside divergent results from honey bee field trials, has important potential

ramifications for pesticide-use policies.

Integrated wild pollinator management: putting wild bees to work for crop and

wildflower pollination

Cory S. Sheffield, Royal Saskatchewan Museum, Regina, SK S4P 2V7

Bees, unlike many other groups of pollinating insects, are Central Place Foragers,

foraging for floral resources in areas surrounding their nest, the radius being

approximately equal to the maximum flight distance of the individual species (larger bees

typically flying further). For a nesting bee, being restricted to this area has implications

for both pollination and conservation, because this landscape must provide ample nectar

and pollen and, for some species, nesting materials; areas lacking all the requirements

will be abandoned and, over the long term, will lose bee populations. Canada has more

than 850 wild bee species, and a large proportion of these are generalist pollen users and

visit (thus pollinate) many of our crops. Many of these same species also visit non-crop

plants, so provide valuable ecological services to the natural and semi-natural

communities surrounding crop lands. Central Place Foraging, body size, flight range, and

floral resource availability all have to be considered when considering the use of wild

bees for crop pollination and in maintaining populations for pollination in non-crop

habitats. These factors, along with life history characteristics of bees, will be discussed in

the context of pollination and management of wild bees.

Pollinator Partnership Canada

Lora Morandin, Pollinator Partnership Canada, Victoria, B.C.

Pollinator Partnership Canada (P2C) 1s the first international expansion of Pollinator

Partnership (P2), which is the largest non-profit organization dedicated solely to the

preservation of pollinators and their ecosystems. Pollinator Partnership and P2C work to

conserve pollinators through research, policy, outreach and education, collaboration, and

habitat creation. Pollinators are directly responsible for providing approximately one-

third of the food we eat and are essential to natural ecosystems. Yet, both managed and

wild pollinators are facing numerous pressures and population declines due to habitat

loss, pest and diseases, invasive species, climate change, and exposure to pesticides. In

Canada, P2C has created a national planting guide for honey bee forage in association

with the Bee Health Roundtable, created 14 new ecoregional pollinator planting guides

with native plant lists, and reviewed Canadian bee habitat programs. We are beginning

new programs to promote monarch conservation through research and habitat creation

and are launching local networks to facilitate education and collaborative action.

J. ENTOMOL. SOC. BRIT. COLUMBIA 114, DECEMBER 2017 101

Presentation Abstracts

Entomological Society of British Columbia

Annual General Meeting

Student Union Building, University of the Fraser Valley,

Abbotsford, BC

October 13, 2017

Yeast enhances the attraction of yellowjackets to dried fruit and fruit powder

Tamara Babcock!, R. Gries!, L. Palmero!, G. Gries!, J. Borden’, A. Mattiacci*, M.

Masciocchi?, J. Corley’, ‘Simon Fraser University, *Scotts Canada, 7>GEPI-INTA

There is need for the development of a better trap bait which can effectively trap

pestiferous yellowjacket species. We field tested dried fruit and fruit powder baits with

and without yeast, and found that the addition of yeast improved the attractiveness of

fruit baits to yellowjackets by up to 50-fold.

Toxin diversity and specificity in a Drosophila defensive symbiosis

Matt Ballinger and Steve Perlman, Department of Biology, University of Victoria

Ribosome-inactivating proteins (RIPs) have been implicated in Spiroplasma symbiont-

mediated defense of Drosophila against parasitic nematodes. We test the activity of these

toxins against parasitoid wasps, implicating them in protection against endo- but

not ectoparasitoids, and examine the role of diverse natural enemies of insects in driving

the evolution of bacterial toxin repertoires.

Population state-dependent invasion potential of the mountain pine beetle in

Alberta

Jordan Lewis Burke, Richard Hamelin, Allan Carroll, Dept. of Forest and Conservation

Sciences, Faculty of Forestry, University of British Columbia

The mountain pine beetle [MPB] has invaded novel pine habitat in Alberta, and has

transitioned from the native species, lodgepole pine, into the novel boreal species, jack

pine. While evidence currently supports the prediction that MPB will have an advantage

in jack pine during epidemic population phases, the fate of low-density endemic

populations is unclear. Here, I present two studies conducted over the last two years,

which surveyed trees typically selected by endemic MPB for competitor dynamics in the

field, and compared MPB symbiont growth characteristics under a range of conditions

and compared these to an antagonistic competitor in the lab. Results demonstrate a clear

disadvantage for endemic MPB in jack pine compared to lodgepole. MPB, and

potentially other eruptive bark beetles, are likely to exhibit population state-dependent

invasion potential, where epidemic behavior may lead to success while endemic

behaviors may lead to failure to establish in novel systems with no coevolutionary

history. These results support continued effort by forest managers in Alberta to prevent

MPB populations from breaching epidemic thresholds, as their populations at low-

density are unlikely to be stable, and unable to persist long-term.

Creating a DNA biomarker to identify dengue refractory and susceptible Aedes

aegypti

Heather Coatsworth, Clara Ocampo, Carl Lowenberger, Simon Fraser University

Dengue viruses transmitted by Aedes aegypti infect 50-100 million people each year. In

Colombia, 30% of feral Ae. aegypti are dengue refractory. We conducted a genome-wide

association study comparing susceptible and refractory mosquitoes. Variants were

102 J. Entomol. Soc. Brit. Columbia 114, December 2017

investigated for possible relevance to the phenotypes. Secondary validation of these

variants is currently underway.

Cutworm Killer: an Okanagan Beauveria bassiana isolate shows promise for

climbing cutworm control in vineyards

Naomi DeLury and Tom Lowery, AAFC-SuRDC

Climbing cutworms (Lepidoptera: Noctuidae) are a major pest of grapevines in the

Okanagan and Similkameen valleys of BC, attacking grape buds early in the spring when

temperatures are low. We compare the efficacy of a local field-collected isolate of

Beauveria and commercial strains against local and introduced cutworm species.

Mixed pathogen interactions: how does host nutrition modulate disease?

Pauline S. Deschodt, Olivia J. H. Walker, Alana K. Breitkreutz and Jenny S. Cory,

Department of Biological Sciences, Simon Fraser University

Individual hosts are commonly challenged by multiple pathogen species. Yet, studies on

insect-pathogen interactions mainly focus on interactions between a single host and a

single pathogen. Two (or more) pathogens co-infecting a host may compete directly

(interference) or indirectly, for resources or via the host immune system. These

competitive interactions could increase or decrease host mortality, or result in no change,

as well as alter the transmission of disease within the population. In insects, increased

dietary protein can increase survival, to pathogens such as baculoviruses and bacteria,

even when nutrition is altered post-infection. However, the role of nutrition in mixed

pathogen infections is not known, but is likely to relate to the relative cost of resistance

to different pathogen groups. Using the cabbage looper, Trichoplusia ni, its

nucleopolyhedrovirus (TnSNPV) and the entomopathogenic fungus, Beauveria bassiana,

we asked whether host nutrition could alter the outcome of a mixed infection. We

challenged T. ni larvae with either a single pathogen species or two simultaneously; then

reared the larvae on an artificial diet differing in levels of two major macronutrients,

protein and digestible carbohydrate (quality) or the total amount of these two

macronutrients (quantity). The results suggest that the virus and fungus respond

differently to host nutritional intake, especially on different ratios of protein and

carbohydrate. As expected, poor quantity diet exacerbates the negative effect of pathogen

on host survival. Moreover, in co-infection, the effect of diet composition on host

mortality is greater at lower pathogen doses. These results indicate that diet could be an

important modulator of mixed infections.

Within-individual repeatability of behavioural activity levels of the parasitoid

Pachycrepoideus vindemmiae |

Wendy Fleming, University of Victoria

A model system for tracking parasitoid behavioural activity levels and connecting them

to biological control performance was developed using the pupal parasitoid

Pachycrepoideus vindemmiae. Three aspects of activity levels were studied: 1) circadian

patterns; ii) links to sex and body size; and 111) within-individual repeatability

(“personality’’).

How to Train your Parasitoid (in Sawdust)

Jessica Y.W. Leung! and Paul K. Abram’, 'Simon Fraser University, ?Agriculture & Agri-

Food Canada

In a proof-of-concept study, we show that the parasitoid Pachycrepoideus vindemmiae, a

candidate biological control for the berry pest Drosophila suzukii, can be retained for

longer in a realistic substrate where hosts are usually present (sawdust mulch) when it

has been "trained" to associate the substrate with D. suzukii pupae.

J. ENTOMOL. SOC. BRIT. COLUMBIA 114, DECEMBER 2017 103

Synthetic Aphid Honeydew Volatiles Attract Mosquitoes (Diptera: Culicidae)

Dan A.H. Peach, N. Young, R. Gries, S. Kumar, G. Gries, Simon Fraser University

Adult mosquitoes exploit a variety of plant sugar sources. Plant-derived semiochemicals

guide mosquitoes to inflorescences and fruit, but the cues that attract mosquitoes to other

sources remain largely speculative. Drawing on literature reports of aphid honeydew

volatiles, we tested the attraction of synthetic honeydew volatile blends to Aedes aegypti

mosquitoes.

Flash in the pan or long term threat? MPB in novel pine habitats

Stan Pokorny, University of British Columbia

No abstract provided.

Manipulating Vector Competence in the Yellow Fever Mosquito, Aedes aegytpi

Lea Sanchez Milde, Heather Coatsworth, and Carl Lowenberger, Simon Fraser

University

Aedes aegypti is the principal vector of dengue viruses. We are using CRISPR-Cas9

technology to knock out specific mosquito genes to generate dengue-refractory

mosquitoes. We then will evaluate fitness and vector competence of the knockout lines to

determine their suitability for use in dengue reduction programs.

Effect of nutrition status on the lifespan and reproductive output of the click beetle

Agriotes obscurus

Kari Zurowski'!, Jenny Cory', Jessi Ly', Danielle White’, Todd Kabaluk’, Alida Janmaat’,

!Simon Fraser University, ?University of the Fraser Valley, *Agriculture and Agri-Food

Canada

Adult A. obscurus were paired and provided with an apple slice (fed) or no apple

(starved) to determine the effect of nutrition on reproduction. Egg numbers and

Oviposition were recorded. Starved females laid fewer eggs for a shorter period than fed

females, suggesting nutrition is important for A. obscurus reproduction.

Trade-offs between reproduction and disease resistance in the click beetle Agriotes

obscurus

Kari Zurowski'!, Jenny Cory', Jessi Ly', Danielle White’, Todd Kabaluk?, Alida Janmaat’,

!Simon Fraser University, *University of the Fraser Valley, *Agriculture and Agri-Food

Canada

Adult A. obscurus were challenged with a high concentration, a low concentration, or a

control of M. brunneum and their reproduction was monitored. Egg numbers and

Oviposition were recorded. Females challenged with the pathogen laid fewer eggs for a

shorter amount of time than unchallenged insects, suggesting lifespan restricted

fecundity.

Effect of duration and location of pheromone trap placement in field margins on

population estimates of two click beetle species

Wim Van Herk, Agriculture & Agri-Food Canada

Pheromone traps can be used to approximate the population size of pest click beetle

species in areas where their larvae (wireworms) cause extensive damage to field crops

(e.g. in PEI). If trap catches are used for making decisions in an IPM program for

wireworms, it is important to know under what conditions trap catches are representative

of the beetle populations present. The main pest click beetle species in Canada disperse

primarily by walking, and hence it is likely that keeping a pheromone trap in a permanent

location (e.g. in field margins) causes the population immediately around it to be

depleted. Hence depending on how long they are maintained in a fixed location in the

field, traps may underestimate actual populations. In this talk we demonstrate that this

occurs, is affected by weather, and that it varies with species.

104 J. Entomol. Soc. Brit. Columbia 114, December 2017

A trait-based approach to predicting spread rates of invasive forest insects

Brian Van Hezewijk & Lara Van Akker, Natural Resources Canada, Pacific Forestry

Centre

Being able to predict how fast an invasive species will spread is crucial information for

the management of novel alien species. Based on an historical database that documents

the invasion of Canada’s forests by 329 species of arthropods, we developed a statistical

model that incorporates biological traits as well as geographic variables to predict the

asymptotic rate of spread of new invaders.

J. ENTOMOL. SOc. BRIT. COLUMBIA 114, DECEMBER 2017 105

Symposium Abstracts:

Biological Control — A Safe Approach to Pest Management

Entomological Society of British Columbia

Annual General Meeting

Kwantlen Polytechnic University, Langley, BC, October 14, 2017

Biological control in cannabis

Amanda Brown, Biobest Canada Ltd.

I presented an overview of the beneficial insects that are currently used in indoor

Cannabis production in Canada and the USA. Biological control programs are widely

used because of the limited number of pesticides that can be used for insect pest control

in order to comply with Health Canada and state regulations. The main pests in this crop

are thrips, spider mites, fungus gnats — and occasionally russet mites, broad mites, and

root aphids. Many biocontrol agents — including predatory mites, soil predators, minute

pirate bugs, nematodes and parasitoids — are used for successful control of these pests.

Insect pathogens as biological control agents

Jenny S. Cory, Department of Biological Sciences, Simon Fraser University, Burnaby,

British Columbia

Insect pathogens have been studied for over 100 years, initially because of their

negative effects on insects of economic interest, such as silkworms, and more recently as

agents for pest management. The most studied insect pathogens are the

entomopathogenic fungi, bacteria belonging to the Bacillus species, baculoviruses and

entomopathogenic nematodes. All of these pathogen groups occur naturally in insect

populations and many can cause wide-scale epizootics in their hosts. For example, when

the locally common western tent caterpillar, a cyclic species, periodically reaches very

high numbers, they will invariably die of a baculovirus infection. The insect pathogens

used for pest management have narrow host ranges; all are restricted to insects, most

only infect a few species and many are host specific. The most commercially successful

microbial insecticide is the bacterium Bacillus thuringiensis, but representatives of all the

major pathogen groups are available commercially. Insect pathogens tend to be used like

chemical insecticides in that they are applied to high density pest populations. They are

commonly used in forestry and for greenhouse crops, and they have also been widely

used in crops such as maize, soybean and cotton and have been developed for fruit crops

such as apples and pears. Their main advantages are that they are not toxic and have a

narrow host range, and thus do not cause environmental damage or have health effects on

humans. They could potentially be used more widely with the development of more

novel pest management strategies which use their ecology, for example, their capacity to

be dispersed and re-cycle naturally in pest populations.

Invasive insects: Is biological control an option?

Tracy Hueppelsheuser, British Columbia Ministry of Agriculture

Invasive species impact North American ecosystems both managed and unmanaged

in significant ways. The Centre for Invasive Species Research in California states that

invasive species cost California $3 billion/year, every 60 days a new invasive enters

California, and 6 new invasive species establish each year in California. Many new

species to BC come up from initial introductions in California. Additionally, An Invasive

Alien Species Strategy for Canada (2004) states that invasive alien species are the second

most significant threat to biodiversity, after habitat loss. Canada has a long history of

non-native introductions. The earliest record is Codling Moth (Cydia pomonella) in

Ontario in 1635. The Canadian Food Inspection Agency states that invasive introductions

106 J. Entomol. Soc. Brit. Columbia 114, December 2017

are on the increase due to increasing volume of trade, access to international markets,

tourism and other travel, and decreasing transportation time.

The population of a new species in a new niche or location follows a sigmoidal curve,

with the first few years being sub-detection, followed by some years of rapid increase in

numbers which is usually when the new species is detected, and finally after some years

will reach the carrying capacity of the new environment. Biological control agents,

either naturally occurring or introduced, can play a role in decreasing the carrying

capacity of the new environment, and ideally keeping the new species below acceptable

levels where it doesn’t cause significant damage. New or invader species may be more or

less prone to control with biocontrol agents in the new environment, but most will fall

somewhere in the middle.

Some examples of major new invaders to North America which are having an impact

on agriculture crop production as well as urban landscapes are Spotted Wing Drosophila

(Drosophila suzukii), Brown Marmorated Stink Bug (Halyomorpha halys), other stink

bugs and leafhoppers. In all these cases, biocontrol probably has the best fit as part as a

multi-faceted systems approach to overall crop and ecosystem management. For

example, many ideas are being explored or currently contributing to D. suzukii

management, including more precise insecticide use, cultural and mechanical methods,

exploitation of insect behaviour, in addition to exploring impacts and utility of

entomopathogens, predators, and parasites on this new pest to berries and stone fruit. In

the case of D. suzukii and H. halys, biological control with native parasitoids is low,

generally less than 2%. Reasons include the fact that parasitoids aer not used to

searching for hosts in the new niche that D. suzukii utilizes (ripe fruit vs decaying fruit),

and that D. suzukii is especially good at encapsulating parasitoids, preventing them from

developing. In the case of H. halys, native parasitoids do recognise the egg masses as

suitable and oviposit in them, but unfortunately, the progeny cannot develop. Though it is

a long and arduous process, biologically and from a regulatory perspective, there are

significant efforts by researchers in the USA and Canada to screen and test suitable

specialist parasitoids from the countries of pest origin in Asia. Suitable candidates will

confer much higher levels of parasitism, and enable classical or innundative release as a

practical component of pest management.

Understanding insect oviposition behaviour and its influence on purple loosestrife

biocontrol success

Alida F: Janmaat, Biology Department, University of the Fraser Valley

Patterns of oviposition can be used to elucidate the role of biotic and abiotic factors in

the oviposition decisions made by insect biocontrol agents. Findings from a longterm

study on the oviposition patterns of Galerucella calamariensis, \eaf-feeding beetles

released to control purple loosestrife, were presented. These findings coupled with

laboratory experiments suggest that female oviposition choices made on the level of an

individual plant may provide explanations for variation in biocontrol success observed

across sites. Furthermore, the relationships observed suggest that cannibalism may play

an under-appreciated role in the persistence of biocontrol insects in the field.

What is biological control and why do we need it?

Judith H Myers, Biodiversity Research Centre, University of British Columbia,

Vancouver, BC

Biological control broadly defined is any non-chemical control. Generally biological

control involves the use of natural enemies to control pest species. Five types of

biological control were discussed at the symposium: 1. natural control, 2. augmentative

control based on the release of natural enemies that have been collected or reared for this

purpose, 3. conservation control in which habitat is preserved to maintain populations of

natural enemies in the vicinity of agricultural fields, 4. microbial control using fungus,

bacteria, virus or nematodes, and 5. classical biological control of exotic insects and

J. ENTOMOL. SOC. BRIT. COLUMBIA 114, DECEMBER 2017 107

weeds through the release of natural enemies from the area where the pest is native.

Classical biological programs are expensive and are taken on when there is evidence that

the problem 1s of sufficient economic or ecological cost to warrant the expense and there

is wide support for the program. Although host testing precedes releases of biological

control agents, concerns about nontarget impacts have resulted in fewer programs being

initiated in the last 20 years. Classical biological control programs have been successful

in British Columbia for the weeds hounds tongue, tansy ragwort, diffuse knapweed, St.

John’swort, and Dalmatian toadflax and for the winter moth. Currently programs for the

release of biological control agents on knotweed and spotted wing drosophila are

underway. It is important that classical biological control remains in the toolbox for

dealing with exotic pests in the future.

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NOTICE TO CONTRIBUTORS

The Journal of the Entomological Society of British Columbia is published online as submissions

are accepted. The JESBC provides immediate open access to its content on the principle that

making research freely available to the public supports a greater global exchange of

knowledge. Manuscripts dealing with all facets of the study of arthropods will be considered for

publication. Submissions may be from regions beyond British Columbia and the surrounding

jurisdictions provided that content is applicable or of interest to a regional audience. Review and

forum articles are encouraged. Authors need not be members of the Society. Manuscripts are peer-

reviewed, a process that takes about six weeks.

Submissions. The JESBC accepts only electronic submissions via the journal homepage: http://

journal.entsocbe.ca/. Style and format guidelines, open access fees, and other information for

authors can be found at the journal homepage.

Scientific notes. Scientific notes are an acceptable format for short reports. They must be two

journal pages maximum, about four manuscript pages. Scientific Notes do not use traditional

section headings, and the term "Scientific Note" precedes the title. A short abstract may be included

if desired. Notes are peer- reviewed in the same manner as regular submissions.

Natural History & Observations. Natural history observations are short — typically about two

pages maximum — pieces outlining entomological observations such as (but not exclusively) insect

outbreaks, population collapses, observations in unexpected locations, novel behaviors, etc. The

purpose of these papers will be to record potentially important phenomena — that may otherwise be

overlooked — for the use of future researchers. Papers of this sort were common in many of the

earlier years of the journal, and we feel that they still have a place, particularly as we move further

into the Anthropocene. Natural History and Observations pieces will be subject to peer review.

While not necessarily required, pieces that supply photographic, video, audio, two-witness,

voucher specimens, and/or other rigorous evidence of the observation will be more likely to be

accepted. Photographs will be included in the journal. Video, audio, or other such files should be

deposited in a DOI-based repository (such as figshare, Dryad, or an institutional repository) with

the DOI noted in the written piece. Voucher specimens should be deposited in recognized musuems

with a notation to that effect in the text. The JESBC encourages entomologists and other naturalists

working in British Columbia and the surrounding jurisdictions to consider submitting their

observations as part of the long term record of our regional entomological natural history.

Review and forum articles — Please submit ideas for review or forum articles for consideration to

the editor at journal@entsocbc.ca. Reviews should provide comprehensive, referenced coverage of

current and emerging scientific thought on entomological subjects. Forum articles of about 1000

words in length should provide opinions, backed by fact, on topics of interest to entomologists and

to the general public. Both review and forum submissions are only published in the journal

following full and rigorous peer review.

Electronic reprints. As an open access journal, article reprints from all issues of the journal can be

downloaded as PDFs from the journal homepage.

Back issues. Electronic back issues are available online free-of-charge.

Membership in the Society is open to anyone with an interest in entomology. Dues are $20 per

year for domestic memberships, $10 for students, and $25 for international memberships. Members

receive electronic copies of Boreus (the newsletter of the Society), and when published, occasional

papers. Membership and membership renewals can be purchased at: http://entsocbc.ca/

membership/.

WLU A

3 9088 01976 7789

Journal of the

Entomological Society of British Columbia

Volume 114 December 2017 ISSN#0071-0733

Directors of the Entomological Society of British Columbia 2016-2017........ Gn eee Z

Grape leaf rust mite, Calepitrimerus vitis (Acari: Eriophyidae), a new pest of grapes in

Beastie CTU soa serosa ae ys eas eset Pt Bs Beaded ph viakirte id whats See 3

Insect taxa named for the Rev. John H. Keen, early naturalist on the Queen Charlotte

islands ane al-Vietlakaila,JoritisieCOnMmMDiat nuke es Oo Sa es OR 15

Western balsam bark beetle, Dryocoetes confusus Swaine (Coleoptera: Curculionidae:

Scolytinae), in situ development and seasonal flight periodicity in southern British

OUT CIG ARE ICUs (SRAM et i A CREAT vicif LAs Dalgien 9 00 Cat SOR eA mR Om A a eRe ER a2

An updated and annotated checklist of the thick-headed flies (Diptera: Conopidae) of

rishi Orumpia, the Yukon, and Alaska se cc sk i ens ow ees 38

Supercolonies of the invasive ant, Myrmica rubra (Hymenoptera: Formicidae) in British

COminila, Cain At ae el ee ahs a ia ued nee 56

SCIENTIFIC NOTES

First record of Aedes (Ochlerotatus) spencerii (Theobald) (Diptera: Culicidae) in the Yukon

ee eee ee ee Oe eh nae Lhe we cu iran eae Wanda Eo eaanic cals aul 65

Cold requirements to facilitate mass emergence of spruce beetle (Coleoptera:

Crbecutiomagdac adults ite Ta Or anny oie iene. consol do dive acs op uiwidnw Wee ead ais ae 68

Production of epicormic buds by Douglas-fir in central British Columbia, Canada,

following defoliation by western spruce budworm (Lepidoptera: Tortricidae) .............. 73

NATURAL HISTORY AND OBSERVATIONS

Archilestes californicus McLachlan (Odonata: Zygoptera: Lestidae):a damselfly new to

i eh teat tattle ies ie EN oles We si nletaos colo aarclolan vi baabiay tan lwaboinan Sp Rane Ti

Evidence of established brown marmorated stink bug populations in British Columbia,

NG a ria rrtey slucccte ope Rae Pats oc beks cap ie cueidl al cs ncgaains Sala aoluja Waselh oe Re sett aang 83

An unusual specimen of the subgenus Lasioglossum Curtis from British Columbia, Canada

(Hey ENA gr AIO TBO) ore fe lk ces ve es an Sic saan aes mre eta 87

First identifications of aphid and diamondback moth populations on wasabi in British

TL i EES Ones EE ORT SRS GN) TS Sed eo Ane er Sea Re eRe 93

SYMPOSIUM ABSTRACTS

Proceedings of the Pollination: Science and Stewardship Symposium.....................045 D/

Pieper As irae: ESIC AG ops eas asw on. cie seieiatbusiaitertucomcan!atesstaivtclon sieltulo wn ae woos 101

Symposium abstracts: Biological Control -A Safe Approach to Pest Management ...105

PE MO LOIN PTR eG secs stciabtagstyicksccuic'l-5 owls po) oS da oniisies Senda gee sobs Inside Back Cover