J. ENTOMOL. SOC. BRIT. COLUMBIA 113, DECEMBER 2016 1

Journal of the,

Entomological Society of British

Volume 113 December 2016 ISSN#0071-0733

Directors of the Entomological Society of British Columbia 2016-2017....................45 2

Development and oviposition preference of Xenotemna pallorana Robinson (Lepidoptera:

Tortricidae) oncalfalta, aad tant ee TOR aie ios os Sense cc unsearooich Sire Gummewstarelaancuitae << 0b 3

Efficacy of SPLAT® Verb for protecting individual Pinus contorta, Pinus ponderosa, and

Pinus lambertiana from mortality attributed to Dendroctonus ponderosae

«se wh annapolis taa ohl a ce at 11

History of the balsam woolly adelgid, Adelges piceae (Ratzeburg), in British Columbia,

with notes On aTrecent range expansions ees 7a Sal eo can eee sa tenes ae ence nude pa:

Pollen preferences of two species of Andrena in British Columbia’s oak-savannah

SCOSV SECU, cs csccstes opens sesiea Wonca ore askin eagle Oe emir a ate ee trie el ie, cae scales 39

Relative efficacies of sticky yellow rectangles against three Rhagoletis fly species (Diptera:

Tephritidae) in Washington State and possible role of adhesives ....................0. ee eeee ees 49

SCIENTIFIC NOTES

Pacific Flatheaded Borer, Chrysobothris mali Horn (Coleoptera: Buprestidae), found

attacking apple saplings in the Southern Interior of British Columbia ........................ 71

NATURAL HISTORY AND OBSERVATIONS

First Canadian records for two invasive seed-feeding bugs, Arocatus melanocephalus

(Fabricius, 1798) and Raglius alboacuminatus (Goeze, 1778), and a range extension for a

third species, Rhyparochromus vulgaris(Schilling, 1829) (Hemiptera: Heteroptera)........ 74

Notes on insects recently introduced to Metro Vancouver and other newly recorded species

TROUT TORUS) © ne oe ta Seno cceuure sn a cane et ha ie cee 719

Rhyparochromus vulgaris (Schilling) (Hemiptera: Heteroptera: Rhyparochromidae): newly

GiSCOVEICH Ill [he Wtetior Ot Mriish COMIN. 5's cocci sconces esa repent cocater sacs sation ses en 90

SYMPOSIUM ABSTRACTS

Proceedings of the Pollination Science and Stewardship Symposium.......................04: 92

Symposium abstracts: Urban Insects — They Live Among Us ....................eceeeee ee 99

PROSCTNACOT ete Fe I oo ois dick nhs no 9 8 eee gap lle 0 ca aesacee Oka FAM ndaeeueen <4) 101

OBITUARY: Thelma Finlayson (29 June 1914 - 15 September 2016) ...................04.. 105

FR i eh TAR ab Meare Nica ilies bi nedsidsa «¥ eune WAS NS Caneaaian a lisse GE CM aa cura 107

J. ENTOMOL. Soc. BRIT. COLUMBIA 113, DECEMBER 2016 2

DIRECTORS OF THE ENTOMOLOGICAL SOCIETY

OF BRITISH COLUMBIA FOR 2016-2017

President:

Brian Van Hezewijk (president@entsocbc.ca)

Pacific Forestry Centre, Victoria

Ist Vice President:

Jenny Cory

Simon Fraser University, Burnaby

2nd Vice-President:

Lisa Poirier

University of Northern British Columbia

Past-President:

Steve Perlman

University of Victoria, Victoria

Treasurer:

Ward Strong (membership@entsocbce.ca)

Ministry Forests, Lands and Natural Resource Operations, Vernon

Secretary:

Tracy Hueppelsheuser (secretary@entsocbc.ca)

B.C. Ministry of Agriculture, Abbotsford

Directors:

Kathy Bleiker (second term), Tamara Richardson (first term)

Graduate Student Representative:

Joyce Leung

Regional Director of National Society: Editor, Boreus:

Bill Riel Gabriella Zilahi-Balogh

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

Service Canadian Food Inspection Agency

Web Editor: Editor, Emeritus:

Alex Chubaty (webmaster@entsocbc.ca) Peter & Elspeth Belton

Canadian Forest Service, Victoria Simon Fraser University

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

Staffan Lindgren, Ward Strong, Lisa

Poirier, Lee Humble, Bob Lalonde,

Lorraine Maclauchlan, Robert

McGregor, Steve Perlman

Editor-in-Chief: Dezene Huber

(journal@entsocbc.ca)

University of Northern British Columbia

Copy Editor: Monique Keiran Technical Editor: Jesse Rogerson

J. ENTOMOL. SOc. BRIT. COLUMBIA 113, DECEMBER 2016 5

Development and oviposition preference of Xenotemna

pallorana Robinson (Lepidoptera: Tortricidae) on alfalfa

and fruit tree foliage

C. A. NOBBS|, R. S. PFANNENSTIEL?, and J. F. BRUNNER?

ABSTRACT

Xenotemna pallorana Robinson (Lepidoptera: Tortricidae) is an alfalfa (Medicago

sativa L., Fabaceae) feeding leafroller that has been considered for incorporation into

apple (Malus domestica Borkh., Rosaceae) orchard ecosystems in Washington State,

U.S.A., as an alternative host for the leafroller parasitoid Colpoclypeus florus Walker

(Hymenoptera: Eulophidae). Xenotemna pallorana has been observed to feed on apple

foliage when populations deplete the foliage of alfalfa in a groundcover, but there have

been no studies to determine if foliage of fruit trees is suitable for full larval

development or would be attractive as oviposition sites. Leafroller larvae were fed

apple, cherry (Prunus avium L., Rosaceae), pear (Pyrus communis L., Rosaceae), and

alfalfa foliage, all of which proved suitable for development; although development

time and pupal weights varied among foliage types. Adult female X. pallorana

exposed to apple foliage under no-choice conditions oviposited on the upper side of

apple leaves. In a choice test between apple foliage and ground cover including alfalfa,

X. pallorana females preferentially selected alfalfa and other components of the

ground cover (98.04%) over the apple foliage (1.94%) for oviposition. Despite the

ability of X. pallorana to develop on fruit tree foliage, its distinct preference for

ovipositing on alfalfa suggests that it is unlikely to damage fruit. Therefore, X.

pallorana presents a low-risk opportunity to study enhancement of biological control

of leafrollers in orchards through ground cover management and host augmentation.

Key Words: Colpoclypeus florus, Xenotemna pallorana, ground cover, alternative

host

INTRODUCTION

Once considered secondary or minor pests, the leafrollers Choristoneura rosaceana

(Harris) (Lepidoptera: Tortricidae) and Pandemis pyrusana Kearfott (Lepidoptera:

Tortricidae) are now two of the most important pests in Washington pome fruit orchards

(Brunner and Beers 1990; Brunner 1994; Brunner 1996b; Brunner 1999). This is

especially true in orchards that use pheromone-based mating disruption for the control of

codling moth, Cydia pomonella L. (Lepidoptera: Tortricidae) (Gut and Brunner 1998).

The seasonal life histories of C. rosaceana and P. pyrusana are similar and fairly

synchronous, with both species having two generations per year in Washington State and

overwintering as small larvae (Beers et al. 1993). Control of leafrollers in orchards has

historically relied on broad-spectrum insecticides (Beers et al. 1993; Brunner 1999);

however, the development of insecticide resistance has made control difficult (Sial and

Brunner 2010a). Additionally, even the integration of insecticides with newer chemistries

might lead to resistance onset within 5 to 10 generations (Sial and Brunner 2010b) if not

carefully managed. The loss of conventional control products and continued evolution of

resistance to pesticides greatly increase the need for non-chemical alternatives to

suppress leafroller populations in orchards.

1 Dupont Crop Protection, 3213 Barge St., Yakima, WA 98902, USA.

2 Corresponding author: USDA-APHIS PPQ, 4700 River Rd, Riverdale, MD 20737 USA; (785)

236-9367, rpfannenstiel@email.com

3 Department of Entomology, Washington State University, Tree Fruit Research and Extension Center,

1100 Western Avenue, Wenatchee, WA 98801

4 J. ENTOMOL. Soc. BRIT. COLUMBIA 113, DECEMBER 2016

Biological control may play a significant role in an integrated approach for leafroller

control. One natural enemy that has shown promise against leafrollers in Washington

State is the gregarious ectoparasitoid Colpoclypeus florus Walker (Hymenoptera:

Eulophidae). This species was first discovered in Washington State in 1992, where it

parasitized about 80 percent of the summer generation P. pyrusana in an unsprayed apple

orchard (Brunner 1996a). In Europe, C. florus parasitizes more than 30 species of

tortricid larvae (Dijkstra 1986) and has been reported to parasitize large numbers of

leafrollers in Europe that feed on both apple and strawberries (Gruys and Vaal 1984).

However, it is absent or rare in spring, although it can be common in summer and early

fall, causing significant mortality of leafroller larvae (Gruys 1982). The absence of C.

florus in spring limits its impact as a biological control agent in Europe. The disparity in

the seasonality of parasitism is presumably caused by a lack of synchrony between the

phenology of C. florus and leafroller species present in European orchards (Gruys 1982).

Colpoclypeus florus attacks relatively large leafroller larvae (Gruys and Vaal 1984;

Dijkstra 1986); however, most tortricids found in orchard systems overwinter as early

larval instars that are not suitable hosts for C. florus in Europe (Gruys and Vaal 1984).

Female C. florus searching for suitable hosts in the fall within orchards cannot find them

and evidently leave the orchard environment in search of suitable overwintering hosts

(van Veen and Wijk 1987). In Europe, no leafrollers have been found that provide

overwintering opportunities in orchards for C. florus (Evenhuis and Vlug 1983).

However, if suitable hosts are placed within orchards, parasitism is high (Pfannenstiel e

al. 2012).

Parasitism of the summer leafroller generation in Washington by C. florus can be high

(>60%) (Brunner 1996a), but spring parasitism is generally low, and parasitism of the

summer generation is often insufficient to prevent fruit damage (Unruh et a/. 2001). As in

Europe, the leafroller pests in Washington orchards, C. rosaceana and P. pyrusana,

overwinter as instars that are typically too small to be suitable hosts for C. florus

(Pfannenstiel and Unruh 2003; Pfannenstiel et al. 2010). One possible tactic to augment

biological control of leafrollers would be to provide alternate overwintering hosts for C.

florus.

A leafroller being considered for this is Xenotemna pallorana Robinson (Lepidoptera:

Tortricidae). This leafroller is an alfalfa-feeding species that is in the same tribe

(Archipini) as both C. rosaceana and P. pyrusana, and may overwinter in an appropriate

stage for C. florus to attack in the fall. Xenotemna pallorana is common in alfalfa crops

in some parts of the Columbia Basin (CAN and RSP personal observation). Alfalfa

occurs in orchard groundcovers in Central Washington, but currently X. pallorana is not

commonly observed in orchards (RSP personal observation). Establishment of X.

pallorana in orchard groundcovers in the fall would allow C. florus to locate hosts for

overwintering within the orchard and, thus, be more abundant in orchards the following

spring. Xenotemna pallorana is bivoltine and highly polyphagous (Chapman and Lienk

1971), having been reported feeding on rose and birdsfoot trefoil in New York (Schott

1925; Neunzig and Gyrisco 1955), young white pine stands in Michigan (McDaniel

1936), strawberries in Ohio (Neiswander 1944), and seed alfalfa in Utah (Snow and

McClellan 1951).

Preliminary evaluations demonstrated that X. pallorana is a suitable host for C. florus

(CAN and RSP, unpublished data). When host foliage is depleted, X. pallorana will feed

on apple leaves on young trees where foliage is near the ground (Chapman and Lienk

1971). The occurrence of X. pallorana in Washington apple orchards is associated with

mature larvae using foliage for pupation sites (JFB personal observation).

Before X. pallorana can be recommended as a potential alternative host for C. florus

in orchard ground covers, it is necessary to demonstrate that it would not consistently

feed on and damage fruit trees. Our goal in this study is to determine whether xX.

pallorana feed and develop normally on foliage of three tree fruit species, whether adults

Oviposit on apple foliage under no-choice conditions, and finally whether X. pallorana

J. ENTOMOL. SOC. BRIT. COLUMBIA 113, DECEMBER 2016 5

prefers to oviposit on apple tree foliage when an alfalfa-dominated ground cover is

available.

MATERIALS AND METHODS

A laboratory colony of X. pallorana was initiated from individuals collected from

alfalfa in the Columbia Basin near Quincy, WA, in June 1996. Xenotemna pallorana

larvae were reared in groups (8) within 118-ml plastic portion cups (Solo®), on a

modified pinto bean-based diet (Shorey and Hale 1965) at 24 + 2 °C with a 16:8 (L:D)

photoperiod. Pupae were collected and placed into a cylindrical wire mesh oviposition

cage 20 cm tall and 10 cm in diameter. Waxed paper was placed into oviposition cages,

and X. pallorana was allowed to eclose, mate and oviposit on the waxed paper. Exposure

to natural light greatly increased oviposition, so oviposition cages were positioned near

windows in the rearing rooms. Egg masses were cut from the waxed paper, washed in a

5% sodium hypochlorite solution, and then rinsed with distilled water. Eggs were

carefully peeled from the waxed paper and placed into petri dishes (Falcon 5009, 50 x 9

mm), with a 1-cm cube of artificial diet. Neonates were transferred in groups of eight to

104-ml portion cups, with the bottoms filled approximately 1 cm deep with artificial diet

for larval development.

Development on fruit tree hosts. Forty newly hatched X. pallorana larvae were

placed individually into Petri dishes (Falcon 5009, 50 x 9 mm) containing an

approximate 4-cm? portion of a mature leaf from a growing shoot (4 to 10 leaves from

shoot apex) of apple (Malus domestica Borkhausen var. Red Spur Delicious), pear (Pyrus

communus L. var. Bartlet), or cherry (Prunus avium L. var. Bing). For alfalfa, the apical 4

to 5 leaves of a non-blooming shoot were collected and placed in the same arena type as

fruit tree foliage. Our goal here was simple; to see if the larvae would complete

development to adult on the different foliage types. The use of cut foliage, which can be

lower in quality, provides a conservative estimate of suitability and allowed the study to

be conducted under controlled conditions. Conducting this study on growing trees in the

orchards would have exposed larvae to significant environmental variation and mortality

that might have masked the effects of foliage type and prevented estimation of plant-

based survival rates.

Fruit tree foliage was obtained from multiple trees for each type within unsprayed

orchards at the Washington State University Tree Fruit Research and Extension Center

(WSU-TFREC) in Wenatchee, WA. Alfalfa foliage was obtained from plants transplanted

from the field to 3.8-1 pots in the greenhouse containing 3-1 of Sunshine potting mix #1

(Sungro Horticulture, Agawam, MA, USA) with 30 ml of long-release fertilizer granules

spread over the surface (Osmocote, Everris NA). Xenotemna pallorana \arvae were

placed in a controlled environment at 24 + 2 °C, 70 + 10 % RH, and 16:8 (L:D)

photoperiod. Larvae were examined daily, and mortality was recorded. Food was

changed as needed, when it had been consumed, or if a decline in apparent quality was

observed, but no less than twice per week. Pupating larvae were set aside and weighed

within 48 h of pupal formation but after they had completed melanization. Chi-square

analysis was used to determine if food source affected survivorship of X. pallorana. The

sex of newly emerged adults was recorded. Larval and pupal development time and pupal

weights for each sex reared on each foliage type were analyzed using one-way analysis

of variance (ANOVA; Super ANOVA general linear model program; Abacus Concepts,

Berkeley, CA). Mean separations were done with the Fisher protected least significant

difference test (LSD a = 0.05).

Oviposition Experiments. Experiments were conducted to determine whether X.

pallorana would oviposit on apple foliage in choice and no-choice situations. For the no-

choice studies, 12 sleeve cages made of cloth and wire screen (window screen tubes,

approximately 50 cm long and 25 cm in diameter) were placed over unsprayed apple (c.v.

Red Spur) foliage in the field. Newly emerged male and female X. pallorana adults

6 J. ENTOMOL. SOC. BRIT. COLUMBIA 113, DECEMBER 2016

(<12h old, two of each) obtained from the laboratory colony were released into each

cage. After a period of seven days, each cage was removed, all egg masses were

collected, and their locations were recorded.

A second experiment was conducted to determine if X. pallorana females prefer to

oviposit on foliage of a fruit tree host, apple, or their typical field host, alfalfa, as part of

a groundcover. Four nylon organdy mesh cages (1.22 m x 1.22 m x 1.22 m) were

suspended from a frame of plastic irrigation pipe (PVC, 2 cm) and placed over patches of

alfalfa plants in a fescue (Festuca spp; Poaceae) dominated groundcover. A single potted

apple tree (c.v. “Oregon Spur’, ~ 1 m tall, with at least 30 fully developed leaves) was

placed in each cage. Alfalfa typically comprised <50% of the ground cover foliage.

Although the leaf area was not directly measured for each foliage type, alfalfa foliage

made up a smaller proportion of the potential oviposition substrates than the apple and

fescue foliage (CAN and RSP personal observation). Ten newly emerged colony-reared

X. pallorana, (1:1 sex ratio) were released into each cage and allowed to mate and

Oviposit for seven days. Seven days after the moths were released, cages were removed,

and egg masses were collected from foliage within each cage (n=12). The proportion of

egg masses deposited on apple foliage vs. alfalfa and other groundcover plants was

compared to the null hypothesis of no preference (50:50) using Chi-square analysis.

RESULTS AND DISCUSSION

A similar number of X. pallorana survived to adulthood when reared on the various

foliage types (7? = 3.42, df = 3, P = 0.3313). Survival of X. pallorana on the factitious

hosts was 65% on cherry, 72.5% on pear, and 75% on apple foliage. Survival on alfalfa

was the lowest overall, at 56%; most likely, this is an artifact related to the quality of the

alfalfa foliage declining more quickly than the apple foliage in the bioassay arena. The

host material that larvae were reared on significantly affected larval developmental time

for both males and females (/’ = 16.155; df = 3; P < 0.0001 and F = 7.647; df = 3; P=

0.0003, respectively), whereas pupation time was not different (Table 1). There was also

a significant effect of host material on pupal weight for both males and females (F =

57.477; df = 3; P < 0.0001 and F' = 43.864; df = 3; P < 0.0001, respectively; Table 2).

Xenotemna pallorana was able to complete its life cycle on the foliage of apple, cherry,

and pear, in addition to alfalfa. Larval development was fastest on alfalfa. Pupal weights

were consistently higher for larvae reared on apple and cherry foliage than on alfalfa and

pear. Adults from the heaviest pupae (reared on apple and cherry foliage) had unusually

large abdomens in proportion to their wing size and had difficulty flying (CAN and RSP,

personal observation). It would have been difficult for these adults to disperse and either

mate or deposit egg masses. It may be that the larvae developing on apple and cherry

develop to such a large size through a supernumerary instar, which would account for

both the longer development time and the larger pupal size. Adults that developed on

alfalfa and pear foliage had physical proportions similar to those of individuals collected

from the field or reared on artificial diet.

In no-choice trials, X. pallorana readily oviposited on apple foliage. Egg masses were

found on apple foliage in 8 of 12 sleeve cages, with an average of 2.08 egg masses per

cage. These egg masses successfully hatched, and the larvae were observed to begin

feeding on the leaves; however, further development was not monitored. Egg masses

were found only on the upper surface of apple leaves. Therefore, without a choice, X.

pallorana will accept apple foliage for oviposition. Because X. pallorana oviposits on

wax paper in the laboratory and has been observed to lay egg masses on other smooth

material such as glass or plastic, oviposition on apple foliage when confined was not

surprising.

J. ENTOMOL. SOC. BRIT. COLUMBIA 113, DECEMBER 2016 7

Table 1

Development time of X. pallorana reared on different hosts.

Larval developmental time Pupation time

Host Sex n (mean days + SEM) (mean days + SEM)

Cherry Male 13 35.6 + l.la 11.9+0.2a

Pear 21 32.0 + 0.9b 11.8+0.2a

Apple 1] 34.4 + 1.2ab 11.94 0.3a

Alfalfa 14 25.3.2 1 b6 11.9 +0.2a

Cherry Female 13 40.8+ l.la 11.1:+0.2a

Pear 8 36.5 + 1.4b 10.9+0.3a

Apple 19 35.8+0.9b 10.8+0.2a

Alfalfa 9 33,144.36 102 2 038

Means within a column for the same sex followed by the same letter are not significantly

different (P > 0.05); Fisher’s protected LSD test.

Table 2

Pupal weight of X. pallorana reared on different hosts.

Pupal weight (g)

Host Sex n (mean + SEM)

Cherry Male 13 0.055 + 0.002b

Pear 22 0.038 + 0.002d

Apple 1] 0.073 + 0.002a

Alfalfa 14 0.047 + 0.002c

Cherry Female 13 0.058 + 0.004b

Pear f 0.044 + 0.005c

Apple 19 0.094 + 0.003a

Alfalfa 9 0.042 + 0.005c

Means within a column for the same sex followed by the same letter are not significantly

different (P > 0.05); Fisher’s protected LSD test.

In choice trials, X. pallorana were presented with apple trees and alfalfa in a fescue-

dominated ground cover. A total of 51 egg masses were collected (Table 3), with an

average of 4.25 egg masses per cage (n = 12). The number of egg masses found on

alfalfa and other ground cover plants was significantly higher than on apple, with slightly

over 98% of the egg masses (50) found on alfalfa and other ground cover plants (74.5%

on alfalfa and 23.5% on other ground cover plants; y? = 31.5, df = 1, P << 0.0001) when

compared to the null hypothesis of no preference. Only one egg mass was found on apple

foliage—less than 2% of the total collected.

8 J. ENTOMOL. SOc. BRIT. COLUMBIA 113, DECEMBER 2016

Table 3

Oviposition choice of X. pallorana females between apple foliage and an alfalfa-dominated

cover crop.

Foliage type Total # egg masses _— Percent of total Average # per cage

collected | |

Apple ] 2.0 0.08

Other 12 2 Fae 1.00

Alfalfa 38 74.5 nF

Therefore, when provided a choice, X. pallorana females predominantly deposited

their eggs on alfalfa and other groundcover plants over apple foliage. Other than alfalfa,

ground cover plants on which egg masses were found included blades of fescue

(Poaceae); dandelion (Zaraxacum officinale Weber, Asteraceae); and field bindweed

(Convolvulus arvensis L, Convolvulaceae). The single egg mass found on apple, 2% of

the total egg masses, may have been the result of the location of foliage in close

proximity to the ground cover. Xenotemna pallorana will lay eggs on apple foliage under

no-choice situations. The species can be locally abundant in alfalfa fields in the

Columbia Basin, but are not observed frequently in orchards (CAN and RSP, personal

observation). Oviposition preference for alfalfa most likely explains why X. pallorana,

although occurring in fruit growing areas of Washington and commonly observed in

alfalfa fields, is not commonly observed feeding on apple or other fruit trees.

Additionally, the density of alfalfa in eastern Washington apple orchards varies

considerably, and most orchards use pesticides for insect control, making observations of

X. pallorana less likely.

In conclusion, X. pallorana can complete development on apple, cherry, and pear

foliage, as well as it can on alfalfa, although developmental time is longer in most cases.

Although females may oviposit almost exclusively on a limited range of hosts, they may

develop as fast and as well on non-preferred hosts (Thompson 1988). There is no

aversion of females to oviposition on apple foliage when not given a choice, but under

natural conditions it is clear that they preferentially select ground cover habitats for

oviposition sites. It may be that X. pallorana doesn’t need to deposit eggs exclusively on

hosts such as alfalfa as long as suitable hosts are nearby, which may explain the

frequency of oviposition on non-alfalfa groundcover plants. Larvae may search for and

disperse to alfalfa in groundcover to some extent following hatch. Although this research

shows that at least three orchard crops are suitable hosts for X. pallorana, we believe the

strong oviposition preference for alfalfa would preclude any pest potential by this

species. The X. pallorana—alfalfa combination may offer a low-risk model for the

orchard environment to test the hypothesis that cover-crop management could be used to

enhance leafroller biological control.

ACKNOWLEDGMENTS

We thank Kathy Pierre for technical support in rearing X. pallorana. Dr. Thomas

Unruh and Dr. Dana Nayduch (both USDA-ARS) kindly reviewed an earlier draft of this

manuscript. This research was sponsored by the Washington Tree Fruit Research

Commission.

J. ENTOMOL. SOc. BRIT. COLUMBIA 113, DECEMBER 2016 9

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J. ENTOMOL. SOc. BRIT. COLUMBIA 113, DECEMBER 2016 11

Efficacy of SPLAT® Verb for protecting individual Pinus

contorta, Pinus ponderosa, and Pinus lambertiana from

mortality attributed to Dendroctonus ponderosae

CHRISTOPHER J. FETTIG', BRYTTEN E. STEED’, BEVERLY M.

BULAON?, LEIF A. MORTENSON!, ROBERT A. PROGAR’,

CLIFFORD A. BRADLEY?, A. STEVEN MUNSON®, and AGENOR

MAFRA-NETO’

ABSTRACT

Verbenone (4,6,6-trimethylbicyclo[3.1.1]hept-3-en-2-one) is an antiaggregant of the

mountain pine beetle, Dendroctonus ponderosae Hopkins (Coleoptera: Curculionidae,

Scolytinae), the most notable forest insect pest in western North America. Several

formulations are registered for tree protection, but efficacy is often inconsistent. We

evaluated the efficacy of a newly registered formulation of (—)-verbenone (SPLAT®

Verb, ISCA Technologies Inc., Riverside, CA, USA) for protecting individual

lodgepole pines, Pinus contorta Doug]. ex Loud, ponderosa pines, P. ponderosa

Dougl. ex Laws., and sugar pines, P. lambertiana Dougl., from mortality attributed to

D. ponderosae. Rather than a single release device, SPLAT® Verb is a flowable

emulsion that allows the user to adjust the size of each release point (dollop) according

to desired rates and distributions. SPLAT® Verb applied at 7.0 g of (—)-verbenone/tree

as four equally sized dollops to the tree bole was effective for protecting P. contorta,

but not P. ponderosa. In P. lambertiana, 4.0, 7.0, and 10.0 g of (—)-verbenone/tree

were effective. We discuss the implications of these and other results to the

management of D. ponderosae.

Key Words: mountain pine beetle, Scolytinae, semiochemicals, tree protection,

verbenone

INTRODUCTION

Mountain pine beetle, Dendroctonus ponderosae Hopkins (Coleoptera:

Curculionidae, Scolytinae), is a major disturbance agent in conifer forests of western

North America, where it colonizes at least 15 native pines, most notably lodgepole, Pinus

contorta Dougl. ex Loud., ponderosa, P. ponderosa Dougl. ex Laws., sugar, P.

lambertiana Dougl., limber, P. flexilis E. James, western white, P. monticola Dougl. ex

D. Don, and whitebark, P. albicaulis Engelm., pines (Negron and Fettig 2014). The

geographic distribution of D. ponderosae ranges from British Columbia, Canada, east to

South Dakota, United States, and south to Baja California, Mexico. Populations have

recently been reported in Nebraska, United States (Costello and Schaupp 2011), and the

insect is expanding its range northward in British Columbia and eastward in Alberta,

Canada (de la Giroday et al. 2012). In the last decade, outbreaks of D. ponderosae have

impacted > 27 million hectares of forest (USDA Forest Service 2012; British Columbia

| Corresponding Author: Pacific Southwest Research Station, USDA Forest Service, 1731 Research Park

Drive, Davis, CA 95618, USA; (530) 604-9152, cfettig@fs.fed.us

* Forest Health Protection, USDA Forest Service, P.O. Box 7669, Missoula, MT 59807, USA

3 Forest Health Protection, USDA Forest Service, 19777 Greenley Road, Sonora, CA 95370, USA

4 Pacific Northwest Research Station, USDA Forest Service, 1401 Gekeler Lane, La Grande, OR 97850,

USA

> Montana BioAgriculture Inc., 510 East Kent Avenue, Missoula, MT 59801, USA

6 Forest Health Protection, USDA Forest Service, 4746 South 1900 East, Ogden, UT 84403, USA

7 ISCA Technologies Inc., 1230 Spring Street, Riverside, CA 92507, USA

12 J. ENTOMOL. Soc. BRIT. COLUMBIA 113, DECEMBER 2016

Ministry of Forests, Lands and Natural Resource Operations 2013), and will continue to

be important forest disturbances—particularly given the magnitude of warming projected

for western North America, and its direct and indirect effects on D. ponderosae (Carroll

et al. 2003; Bentz et al. 2010). In conjunction with projected warming trends, susceptible

forest landscapes still exist throughout western North America. Although D. ponderosae

is native to North America and an important part of the ecology of North American

forests, tree mortality resulting from outbreaks may have undesirable social—ecological

impacts, for example, negatively affecting aesthetics, recreation, fire risk and severity,

human safety, timber production, wildlife habitat, and real estate values, among many

other resources.

Progar et al. (2014) provided a thorough review of the chemical ecology of D.

ponderosae relevant to host finding, selection, colonization, and mating behaviors. In

short, females initiate colonization of the lower tree bole in a behavioral sequence

mediated by aggregation pheromones (Vité and Gara 1962; Pitman ef al. 1968, 1969;

Ryker and Libbey 1982) and host kairomones (Renwick and Vité 1970; Borden et al.

1987; Miller and Lindgren 2000). Females are subsequently joined by males, and mass

attack ensues (Pitman et al. 1968), enabling D. ponderosae to overwhelm host tree

defenses consisting of anatomical and chemical components that are both constitutive

and inducible (Franceschi et al. 2005). During latter stages of colonization, increasing

amounts of verbenone (4,6,6-trimethylbicyclo[3.1.1]hept-3-en-2-one) are produced

(Pitman et al. 1969; Rudinsky et al. 1974), which inhibit additional D. ponderosae from

infesting the target tree. The first evidence of this effect was documented by Ryker and

Yandell (1983) in laboratory and field assays. In nature, verbenone is produced in small

amounts by autoxidation of the monoterpene a-pinene (Hunt ef a/. 1989), but the

principal route of production is through metabolic conversion by bark beetles of inhaled

and ingested a-pinene to the terpene alcohols cis- and trans-verbenol, which are then

metabolized to verbenone by yeasts in the alimentary system and within beetle galleries

(Hunt and Borden 1990). It is assumed that verbenone reduces intraspecific competition

and, perhaps, interspecific competition, by altering adult behavior to minimize

overcrowding of developing brood within the host (Byers and Wood 1980). Lindgren ef

al. (1996) proposed that verbenone is an indicator of host tissue quality and that its

quantity is a function of microbial degradation.

Fettig et al. (2014) discussed approaches for reducing the negative impacts of D.

ponderosae on forests. Direct control involves short-term tactics designed to address

current infestations by manipulating beetle populations, and commonly includes the use

of insecticides, semiochemicals (i.e., chemicals produced by one organism that elicit a

behavioral response in another organism), sanitation harvests, or a combination of these

treatments. The use of semiochemicals has largely focused on verbenone for protecting

individual, high-value trees or small groups of trees (e.g., in campgrounds). Results have

been favorable, but inconsistent (for a detailed explanation of associated factors, see

Progar et al. 2014). While several formulations of verbenone are registered for use in

Canada and the United States, pouches (several registrants) stapled at maximum reach

(~2 m in height) to individual trees or applied in a grid pattern for stand protection are

most commonly used (Gillette and Munson 2009).

Fettig et al. (2015) recently developed a novel formulation of (—)-verbenone

(SPLAT® Verb, ISCA Technologies Inc., Riverside, CA, USA) for protecting individual

P. contorta and stands of P. contorta from mortality attributed to D. ponderosae. Rather

than a single release device such as the pouch, SPLAT® Verb is a flowable emulsion that

allows the user to adjust the size of each release point (dollop) according to desired

distributions in the field. SPLAT® Verb is a “matrix-type” diffusion controlled-release

device specifically designed to release (—)-verbenone over a sustained period (~8—24

wks, depending on dollop size) at rates suitable to provide significant reductions in levels

of P. contorta mortality at relatively low doses (Mafra-Neto et al. 2013). Dollops

biodegrade within ~1 yr of application and, as such, do not need to be retrieved from the

J. ENTOMOL. SOC. BRIT. COLUMBIA 113, DECEMBER 2016 13

field as do most other release devices used to dispense verbenone. SPLAT® Verb was

registered by the United States Environmental Protection Agency (USEPA) for use on

pines, Pinus spp., in August 2013, and was first used commercially in the United States

in 2014. The objective of our research was to determine the efficacy of SPLAT® Verb for

protecting individual P. contorta, P. ponderosa, and P. lambertiana from mortality

attributed to D. ponderosae.

MATERIALS AND METHODS

Studies were conducted in three locations (see below) selected based on aerial and

ground surveys that indicated D. ponderosae was causing noticeable levels of tree

mortality in each area. Experimental trees were treated according to the criteria described

below for each study. Regardless of treatment, one commercially available two-

component tree bait [trans-verbenol (~1.2 mg/d) and exo-brevicomin (~0.3 mg/d);

Contech Inc., Delta, BC, Canada] was stapled to the bole of each experimental tree

immediately after treatment at ~2 m in height on the northern aspect, and left in place

until beetle flight had ceased (dates reported in Tables 1-3). The manufacturer estimates

the life expectancy of these baits is 100—150 days, depending on weather conditions,

covering most of the flight activity period of D. ponderosae at each location.

Initially, success of D. ponderosae attacks was based on visual assessments of pitch

tubes and boring dust (condition, distribution, and density) during August-September of

the year treatments were implemented. At that time, experimental trees were recorded as

not attacked, unsuccessfully attacked, strip attacked, or mass attacked (Gibson ef al.

2009). This allows for a surrogate estimate of treatment efficacy should the experimental

infrastructure be compromised or lost (e.g., due to wildfire, which is common in the

western USA). However, tree mortality was ultimately based on presence (dead) or

absence (live) of crown fade ~1 yr after treatments were implemented, except for

Experiment 3, when evaluations were conducted in late-September of the same year. The

only criterion used in determining the effectiveness of each treatment was whether

individual trees died due to colonization by bark beetles. Treatments were considered to

have experienced sufficient beetle “pressure” (i.e., a relative measure of population

density based on levels of tree mortality) to permit determination of efficacy if > 60% of

the untreated control trees were killed by bark beetles. SPLAT® Verb treatments were

considered efficacious if < 7 trees were killed by bark beetles (Hall et al. 1982; Shea et

al. 1984). These criteria were established based on a sample size of 22—35 trees and test

of the null hypothesis, Ho: S (survival > 90%). These parameters provide a conservative

binomial test (a = 0.05) to reject Ho when > 6 trees die. The power of this test, that is the

probability of having made the correct decision in rejecting Ho, is 0.84 (Hall et al. 1982;

Shea et al. 1984). This experimental design provides a very conservative test of efficacy,

and was originally developed for evaluating the efficacy of bole-applied insecticides to

protect individual trees from bark beetle attack. When properly applied, insecticides

typically provide higher levels of tree protection than verbenone (Fettig et al. 2013;

Progar et al. 2014), and as such, our experimental design represents a rigorous

examination of the efficacy of SPLAT® Verb.

Experiment 1 — Pinus contorta: This study was conducted on the Wisdom Ranger

District, Beaverhead—Deerlodge National Forest, Montana, United States (45° 24' 28.98"

N, 113° 39' 28.92" W; 2150 m elevation) during 2013/2014. Surrounding stands had a

mean live tree (= 12.7 cm dbh; diameter at 1.37 m in height) density of 18.0 m?/ha of

basal area (cross-sectional area of trees at 1.37 m in height), of which 87.2% was P.

contorta with a mean quadratic mean diameter (QMD, the diameter corresponding to

mean basal area) of 20.7 cm. The remainder was represented by Engelmann spruce,

Picea engelmannii Parry ex Engelm., and subalpine fir, Abies lasiocarpa (Hooker)

Nuttall. About 28.8% of P. contorta and 43.1% of P. contorta basal area had been killed

by D. ponderosae during the two years preceding the study (cause of death and time

14 J. ENTOMOL. SOC. BRIT. COLUMBIA 113, DECEMBER 2016

since death determined by gallery patterns in the phloem, and by color and needle

retention of the crown, respectively; Klutsch et al. 2009).

Thirty trees (min. dbh = 18.5 cm) were confirmed uninfested and randomly assigned

to each of two treatments (N = 60): (1) SPLAT® Verb [10.0% (—)-verbenone by weight;

EPA Reg. No. 80286-—20] applied at 7.0 g of (—)-verbenone/tree (70 g of SPLAT® Verb/

tree) as four 17.5-g dollops (~5.5 cm diam. X 1.2 cm ht.) to the tree bole at cardinal

directions at ~2.5 m in height using a caulking gun (Model X-Lite, Newborn Brothers

Co., Inc., Jessup, MD, USA); and (2) an untreated control. Treatments were applied on

29-30 June 2013. Adjacent experimental trees were separated by => 100 m. There was no

significant difference in tree dbh between treatments (F1, 53 = 0.3, P = 0.60; Table 1), a

factor that influences tree susceptibility to D. ponderosae (Shepherd 1966). The integrity

of dollops was visually inspected 18-19 September 2013 for evidence of contact or

consumption by animals.

Table 1

Efficacy of SPLAT® Verb (ISCA Technologies Inc., Riverside, CA, USA) for protecting

individual Pinus contorta from mortality attributed to Dendroctonus ponderosae, Wisdom

Ranger District, Beaverhead—Deerlodge National Forest, Montana (45° 24' 28.98" N, 113° 39'

28.92" W; 2150 m elevation), 2013/2014.

Treatment Dose? Mean dbh + SEM Mortality/n

Untreated 0 26.2 + 0.7 26/30

control

SPLAT® Verb 7 26.8 + 0.9 2/30

‘Values are grams of (—)-verbenone applied as four 17.5-g dollops (~5.5 cm diam. X 1.2 cm

ht.) to the tree bole at cardinal directions at ~2.5 m in height using a caulking gun. One tree

bait (Contech Inc., Delta, BC, Canada) was attached to the bole of each tree at ~2 m in height

on the northern aspect 29-30 June to 18-19 September 2013.

Experiment 2 — Pinus ponderosa: This study was conducted on the Darby Ranger

District, Bitterroot National Forest, Montana, United States (46° 04' 22.0" N, 114° 14'

17.7" W; 1344 m elevation) during 2013/2014. Surrounding stands had a mean live tree

(= 12.7 cm dbh) density of 27.2 m?/ha of basal area, of which 92.7% was P. ponderosa

with a mean QMD of 38.9 cm. The remainder was represented by Douglas-fir,

Pseudotsuga menziesii (Mirb.) Franc. About 20.8% of P. ponderosa and 12.4% of P.

ponderosa basal area had been killed by bark beetles, primarily D. ponderosae, during

the two years preceding the study.

Thirty trees (min. dbh = 21 cm) were confirmed uninfested and randomly assigned to

each of the two treatments described in Experiment 1 (V = 60). Treatments were applied

on 30 June—1 July 2013. Adjacent experimental trees were separated by > 100 m. There

was no significant difference in tree dbh between treatments (F', 53 = 0.7, P = 0.40; Table

2).

Experiment 3 — Pinus lambertiana: This study was conducted on the Groveland

Ranger District, Stanislaus National Forest, California, United States (37° 49' 48.63" N,

119° 51' 19.44" W; 1417 m elevation) during 2014/2015 in areas impacted by the Rim

Fire. The Rim Fire is the largest wildfire on record in the Sierra Nevada, California, and

burned 104,131 ha in August 2013 (Kirn and Dickman 2013). The study area was

impacted by mixed-severity fire, with ~26.1% of trees (all species) killed by fire, mostly

in the smaller-diameter classes (<< 31.8 cm dbh). Surrounding stands had a mean live tree

(> 16.5 cm dbh) density of 54.0 m?/ha of basal area, of which 30.7% was P. lambertiana

with a mean QMD of 75.4 cm. The remainder was represented by white fir, A. concolor

J. ENTOMOL. Soc. BRIT. COLUMBIA 113, DECEMBER 2016 15

(Gordon) Lindley ex Hildebrand, P. ponderosa, incense cedar, Calocedrus decurrens

(Torr.) Florin, and, to a much lesser extent, black oak, Quercus kelloggii Newb. About

5.1% of P. lambertiana died during the two years preceding the study, most of which was

attributed to fire, although several trees were colonized and killed by D. ponderosae.

Despite the low level of beetle “pressure” observed, substantial increases in levels of tree

mortality attributed to D. ponderosae were expected in 2014 due to the confounding

effects of fire on tree susceptibility to colonization by D. ponderosae (Jenkins et al.

2014).

Table 2

Efficacy of SPLAT® Verb (ISCA Technologies Inc., Riverside, CA, USA) for protecting

individual Pinus ponderosa from mortality attributed to Dendroctonus ponderosae, Darby

Ranger District, Bitterroot National Forest, Montana (46° 04' 22.0" N, 114° 14' 17.7" W; 1344

m elevation), 2013/2014. Some trees were also colonized by Dendroctonus brevicomis, Ips

pini, and I. emarginatus.

Treatment Dose? Mean dbh + SEM Mortality/n

Untreated 0 a2 244 28/29»

control

SPLAT® Verb fi 30:8 = 52 9/30

‘Values are grams of (—)-verbenone applied as four 17.5-g dollops (~5.5 cm diam. X 1.2 cm

ht.) to the tree bole at cardinal directions at ~2.5 m in height using a caulking gun. One tree

bait (Contech Inc., Delta, BC, Canada) was attached to the bole of each tree at ~2 m in height

on the northern aspect 30 June—1 July to 17-18 September 2013.

> One tree could not be located, and presumably was removed by woodcutters.

Twenty-five trees (min. dbh = 29 cm) were confirmed uninfested and randomly

assigned to each of four treatments (V = 100): (1) SPLAT® Verb applied at 4.0 g of (—)-

verbenone/tree (40 g of SPLAT® Verb/tree) as four 10.0-g dollops (~4.5 cm diam. X 1.5

cm ht.) to the tree bole at cardinal directions at ~2.5 m in height, (2) SPLAT® Verb

applied as in Experiments | and 2, (3) SPLAT® Verb applied at 10.0 g of (—)-verbenone/

tree (100 g of SPLAT® Verb/tree) as four 25.0-g dollops (~5.7 cm diam. X 2.8 cm ht.) to

the tree bole at cardinal directions at ~2.5 m in height, and (4) an untreated control (Table

3). Treatments were applied 23-24 May 2014. Adjacent experimental trees were

separated by > 50 m. There was no significant difference in tree dbh (F3, 96 = 0.5, P =

0.71) or percent crown volume scorched (F3, 96 = 0.5, P = 0.71) among treatments (Table

3). The latter is a significant predicator of the probability of P. Jambertiana mortality

following fire (Hood ef al. 2010). Although not well studied in P. lambertiana, pines

injured by fire are more susceptible to colonization by D. ponderosae (Jenkins et al.

2014).

RESULTS

Dendroctonus ponderosae “pressure” was sufficient to adequately challenge

treatments as 87%, 93%, and 72% of untreated, baited P. contorta, P. ponderosa, and P.

lambertiana, respectively, died from colonization by bark beetles. SPLAT® Verb was

effective for protecting P. contorta and P. lambertiana (tables | and 3) from mortality

attributed to D. ponderosae, but not P. ponderosa (Table 2). All doses evaluated for

protection of P. lambertiana were efficacious (Table 3).

16 J. ENTOMOL. SOC. BRIT. COLUMBIA 113, DECEMBER 2016

Table 3

Efficacy of SPLAT® Verb (ISCA Technologies Inc., Riverside, CA) for protecting individual

Pinus lambertiana from mortality attributed to Dendroctonus ponderosae, Groveland Ranger

District, Stanislaus National Forest, California (37° 49' 48.63" N, 119° 51' 19.44" W; 1417 m

elevation), 2014/2015.

Percent crown

Treatment Dose? Mean 200 i volume scorched + Mortality/n

SEM

Untreated control 0 62.0 + 4.1 378+.6.0 18/25

SPLAT® Verb 4 55.05 3.9 26.6 + 4.9 6/25

SPLAT® Verb 7 60.4 + 4.7 27.0 + 4.6 4/25

SPLAT® Verb 10 66.2 + 6.2 21.8+4.4 1/25

4 Values are grams of (—)-verbenone applied as four equally sized dollops to the tree bole at

cardinal directions at ~2.5 m in height using a caulking gun. One tree bait (Contech Inc.,

Delta, BC, Canada) was attached to the bole of each tree at ~2 m in height on the northern

aspect 23-24 May to 27 August 2014.

DISCUSSION

Experiment 1 confirms results of an earlier study conducted in Wyoming, United

States, demonstrating the efficacy of four 17.5-g dollops of SPLAT® Verb [7.0 g of (—)-

verbenone/tree] for protecting individual P. contorta from D. ponderosae (Fettig et al.

2015). In comparison, two pouches [13.5—15.0 g of (—)-verbenone/tree, depending on

manufacturer] are recommended per tree (Kegley and Gibson, 2009; Kegley ef a/. 2010)

and generally provide => 80% protection of P. contorta and P. albicaulis. For larger trees

(> 61 cm dbh), three to four pouches may be used. One registrant suggests using more

than six pouches: two pouches at ~1.5 and ~2.5 m in height on the northern aspect of

individual trees, with additional pouches placed at 4—5 m intervals on vertical substrates

around the treated tree. Alternatively, Fettig et al. (2015) reported that SPLAT® Verb

provided complete (100%) protection of individual P. contorta at much lower doses and

recommended applying four 17.5-g dollops/tree (1.e., one dollop placed at maximum

reach at each cardinal direction). They attributed the high level of tree protection

observed in their study to multiple release points per tree (Gillette et al. 2006) and the

larger zone of inhibition (i.e., demonstrated to be at least 8 m in radius in trapping

assays) provided by SPLAT® Verb when compared to other formulations of verbenone

that have been studied (Miller 2002; Fettig et al. 2009a, 2015). Fettig et al. (2015)

reported significantly fewer P. contorta (percentage of trees) killed on 0.041-ha circular

plots surrounding P. contorta treated with SPLAT® Verb compared to untreated trees,

suggesting attraction was disrupted at levels sufficient to impart tree protection within 11

m of the point of release. Although we did not measure this variable in Experiment 1, we

observed larger numbers of P. contorta killed by D. ponderosae within the vicinity (~10

m) of untreated, baited controls compared to SPLAT® Verb-treated trees.

As observed in Experiment 2, others have reported verbenone is ineffective for

reducing levels of P. ponderosa mortality attributed to D. ponderosae (e.g., Bentz et al.

1989; Lister et al. 1990; Gibson et al. 1991; Gibson and Kegley 2004; Negron ef al.

2006). However, to our surprise, several of the trees in Experiment 2 were also colonized

by western pine beetle, D. brevicomis LeConte, pine engraver, Ips pini Say, and

emarginated ips, J. emarginatus (LeConte), including eight of nine SPLAT® Verb-treated

trees that died. These bark beetle species are capable of causing tree mortality and are not

inhibited by (—)-verbenone at levels sufficient to impart tree protection (e.g., Devlin and

Borden 1994; Fettig et al. 2009a,b). As such, another evaluation of the efficacy of

J. ENTOMOL. SOC. BRIT. COLUMBIA 113, DECEMBER 2016 17

SPLAT® Verb for protecting P. ponderosa from mortality attributed to D. ponderosae

should be considered in areas where other bark beetles, specifically D. brevicomis, are

absent. This also serves as a reminder of the importance of carefully confirming the cause

of tree mortality when developing semiochemical-based technologies, many of which

impart species- or genera-specific responses.

Experiment 3 represents the first evaluation of a semiochemical-based tool for

protecting P. lambertiana from D. ponderosae. Doses as low as 40.0 g of SPLAT® Verb/

tree were efficacious (Table 3), suggesting lower doses may yield efficacy for protection

of P. contorta and should be evaluated. For decades, populations of P. lambertiana have

been heavily impacted by Cronartium ribicola J.C. Fisch, the exotic pathogen that causes

white pine blister rust (Maloney ef al. 2011). Although white pine blister rust can be fatal

to all species of white pine, a gene is present at low frequency in P. lambertiana that

confers immunity from C. ribicola (Kinloch et al. 1970). This gene controls a

hypersensitive response in needles that prevents further fungal growth (Kinloch and

Littlefield 1976). Restoring populations of P. Jambertiana involves, among other factors,

identifying white pine blister rust-resistant trees in the field and, where feasible,

protecting these individuals from colonization by D. ponderosae with insecticides,

particularly when epidemics occur. This is followed by selective breeding of these

individuals, and eventual outplanting of white pine blister rust-resistant seedlings.

SPLAT® Verb represents a more portable and less toxic alternative to insecticides for

protecting white pine blister rust-resistant P. lambertiana, other high-value P.

lambertiana (e.g., those growing in residential, recreational, and administrative sites), or

P. lambertiana that might otherwise be experiencing short-term stressors that increase

susceptibility to colonization by D. ponderosae. Relatedly, in 2014 we were asked to

treat several large-diameter, fire-injured P. lambertiana in areas impacted by the Rim

Fire within Yosemite National Park (37° 47' 37.32" N, 119° 51' 09.42" W; 1420 m

elevation). Given the size of these trees (mean dbh + SEM = 121.9 + 4.3 cm, max. =

223.1 cm), the height to the base of the crown was often > 15 m, and as such crown

scorch was only observed on one tree. However, the lower boles of all trees were heavily

charred by fire, and bark consumption was evident on some trees. We applied 100.0 g of

SPLAT® Verb to unbaited trees as four 25.0-g dollops (see Experiment 3 for complete

method). Of the 86 trees that were treated in 2014, none were colonized by D.

ponderosae that year. These trees were not retreated in 2015, and therefore left

unprotected. Many of these trees and nearby trees that had never been treated with

SPLAT® Verb were observed being colonized by D. ponderosae in May of 2015.

As with other formulations of verbenone, it is possible that animals could contact

and/or consume dollops of SPLAT® Verb. For example, while never observed directly, in

Experiment 1, we found evidence of small claw marks on several dollops on P. contorta

that we attributed to contact by red squirrels, Zamiasciurus hudsonicus (Erxleben).

Syracuse Environmental Research Associates (2000) conducted a risk assessment of

verbenone, and concluded it was unlikely that consumption by wildlife would have a

detectable impact on any species. While associated toxicology data are scarce, acute oral

LDso values for rats, Rattus spp., the only mammal studied, are estimated at 1,800 mg/kg

for females and 3,400 mg/kg for males (Syracuse Environmental Research Associates

2000). Verbenone administered to bobwhite quail, Colinus virginianus (L.), at doses of

39-300 mg/kg in corn oil had no effect on behavior or health (Syracuse Environmental

Research Associates (2000). Verbenone has mixed effects on several species of insects

(e.g., Lindgren and Miller 2002), however a common predator of bark beetles in western

North America, TZemnochila chlorodia (Mannerheim) (Coleoptera: Trogossitidae), is

attracted to (—)-verbenone, and its impact on bark beetles may therefore be enhanced by

treatments containing verbenone (Fettig et al. 2007). The inert ingredients of SPLAT®

Verb have been certified as food safe by the USEPA (Mafra-Neto et a/. 2013), and

SPLAT® Verb has been granted organic production status by the United States

Department of Agriculture.

18 J. ENTOMOL. SOC. BRIT. COLUMBIA 113, DECEMBER 2016

ACKNOWLEDGEMENTS

We thank K. Bradach, C. Hayes, M. MacKenzie, and J. Phewban (Forest Health

Protection, USDA Forest Service) A. Hermosillo and K. Sharma (ISCA Technologies

Inc.), Janina Bradley and Egan Jankowski (Montana BioAgriculture Inc.), Brian Mattos

(National Park Service), and Ross Gerrard (Pacific Southwest Research Station, USDA

Forest Service) for technical assistance. In addition, we thank the staff of the

Beaverhead—Deerlodge National Forest, Bitterroot National Forest, and Stanislaus

National Forest for providing access to study sites, and two anonymous reviewers for

their thoughtful critiques of an earlier version of this manuscript. This research was

supported, in part, by a collection agreement (14-CO-11272139-018) from ISCA

Technologies Inc. to the Pacific Southwest Research Station.

This publication reports research involving pesticides. It does not contain

recommendations for their use, nor does it imply that the uses discussed here have been

registered. All uses of pesticides in the United States must be registered by appropriate

state and/or federal agencies before they can be recommended. This article was written

and prepared by US Government employees on official time, and is, therefore, in the

public domain and not subject to copyright.

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J. ENTOMOL. SOc. BRIT. COLUMBIA 113, DECEMBER 2016 21

History of the balsam woolly adelgid, Adelges piceae

(Ratzeburg), in British Columbia, with notes on a recent

range expansion

G.M.G ZILAHI-BALOGH}!, L.M. HUMBLE’, R. FOOTTIT?, J.

BURLEIGH*, and A. STOCK?

ABSTRACT

The balsam woolly adelgid, Adelges piceae (Hemiptera: Adelgidae), was introduced

from Europe into eastern North America around 1900 and independently into western

North America sometime before 1928. It was first detected causing damage in North

Vancouver, British Columbia, in 1958. Since then, it has slowly spread to adjacent

areas of southwestern B.C. Surveys from 2011 to 2013 confirmed the presence of A.

piceae in the Cascades Forest District and in the town of Rossland, B.C., which are

outside the pre-2014 quarantine area. Until these recent detections, provincial

quarantine regulations have been the principle tool employed to prevent anthropogenic

spread of the adelgid through the restriction of movement of potentially infested

seedlings and nursery stock from infested coastal regions of British Columbia into the

highly susceptible high-elevation Abies lasiocarpa stands in the Interior forests. We

provide a historical overview of the quarantine regulations enacted since 1966, review

the distribution of Adelges piceae since the first confirmed records of establishment as

documented by historical survey records, and document the extent of recent survey

efforts and new detections in interior subalpine fir forests.

INTRODUCTION

The balsam woolly adelgid, Adelges piceae (Ratzeburg) (Hemiptera: Adelgidae),

occurs on both coasts of North America (NA) and can cause extensive tree damage and

mortality to native Abies species. It was introduced into eastern NA from Europe before

1900 (Foottit and Mackauer 1980) and independently into western North America (Hain

1988), where it was first reported near San Francisco in 1928 (Annand 1928). Adelges

piceae was first reported from British Columbia (B.C.) by E. P. Venables and R. Hopping

(Anon. 1938). Those reports noted its detection, along with Adelges niisslini (B6rner), on

Abies procera Rehder (=A. nobilis (Douglas ex D. Don) Lindley) in Vancouver. In 1958,

A. piceae was discovered damaging a Pacific silver fir, Abies amabilis (Douglas ex

Loud.) Dougl. ex J. Forbes, planted as an ornamental in North Vancouver, B.C. (Silver

1959). Surveys to delimit the range of this introduced pest in the province documented in

the Canadian Forest Invasive Alien Species (CanFIAS)® database (1958-1998; Nealis et

al. 2015) quickly demonstrated its presence on native and ornamental firs in drainages

near Vancouver, as well as on southern Vancouver Island. In 1959, it was found attacking

grand fir (Abies grandis (Douglas ex D. Don) Lindl.) at Thetis Lake, near Victoria, and

' Corresponding author: Canadian Food Inspection Agency, Plant Health and Biosecurity Directorate,

1853 Bredin Rd., Kelowna, B.C. V1Y 789; gabriella. Zilahi-Balogh@inspection.gc.ca

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

Victoria, B.C. V8Z 1M5

> Agriculture and Agri-Food Canada, Invertebrate Biodiversity - National Environmental Health Program

and Canadian National Collection of Insects, Arachnids and Nematodes, Ottawa, ON K1A 0C6

4 Ministry of Forests, Lands and Natural Resources, PO Box 9513 Stn Prov Govt, Victoria, B.C. V8W

9C2

5 Ministry of Forests, Lands and Natural Resources, Kootenay/Boundary Region, Nelson, B.C. V1L 6K1

6 CanFIAS database managed by Ian Demerchant, Natural Resources Canada, Atlantic Forestry

Centre, Fredericton, email: ian.demerchant@canada.ca

22 J. ENTOMOL. SOC. BRIT. COLUMBIA 113, DECEMBER 2016

the following year, it was detected at a commercial nursery near Victoria on white fir

(Abies concolor (Gord. & Glend.)) imported from Holland five years prior to the

detection.

The degree of susceptibility to damage and mortality varies amongst the three native

Abies species in B.C.: damage is generally moderate in grand fir, while that in both

Pacific silver fir and subalpine fir (A. lasiocarpa (Hook.) Nutt.) is more severe.

Subalpine fir is the most susceptible of the true firs in the Pacific Northwest (Mitchell

1966; Hain 1988). | |

In North America, A. piceae is anholocyclic on the ancestral secondary host, Abies

(Havill and Foottit 2007). There are two or more generations per year (Mitchell et al.

1961). The life stages consist of eggs, three nymphal instars, and the adults. Eggs hatch

into crawlers, which are the only stage capable of independent movement and dispersal

(Balch 1952). Crawlers select feeding sites, settling either on the bark of the main stem

and larger branches or at the base of vegetative or reproductive buds, then insert their

stylets into the cortical parenchyma. Once the stylets are inserted, the insect remains

sessile for its life. As the adelgids develop, they molt and secrete waxy threads that

appear as dense white, woolly, or “cottony’ masses. Adult females lay their eggs within

the woolly masses—each of which may contain more than 200 amber-coloured eggs

(Balch 1952).

Feeding by A. piceae on twigs produces gouting at nodes and inhibits new growth,

reducing tree vigor. Stem infestations are a more severe form of attack by A. piceae. The

tree responds to adelgid feeding by producing a type of compression wood in the

sapwood called “rotholz”. This abnormal growth of the sapwood tissue inhibits water

flow within the tree and eventually leads to tree death (Balch 1952; Livingston et al.

2000). The presence of rotholz in the annual rings of attacked trees in North Vancouver

suggested that A. piceae had been present in southwestern B.C. for 8—11 years before its

discovery (Silver 1959).

In British Columbia, A. piceae has historically been restricted primarily to

southwestern B.C., with the Coast and Cascade Mountains acting as natural barriers to

eastward range expansion into Interior stands of Abies lasiocarpa growing at high

elevations. To protect the susceptible subalpine fir stands in the interior of the province,

the Province of British Columbia has maintained regulatory restrictions on the

production and movement of all living Abies spp., as well as logs and cut Christmas trees

since 1966. Coulson and Witter (1984) observed that the initial quarantine restrictions

implemented against A. piceae in the mid-1960s were effective in stabilizing the

infestation boundary after 1967. In contrast, in the absence of any regulatory restrictions

in the western United States, A. piceae has expanded its range extensively through

Washington, Idaho and Montana. It is now present in all U.S. counties bordering B.C.

(Hayes 2015; Liebhold et a/. 2015) and has caused extensive mortality of A. lasiocarpa

in the Sawtooth National Forest in southern Idaho (Livingston ef al. 2000; Livingston

and Pederson 2010).

In 2008, a single branch sample was submitted from a 60-year-old Abies lasiocarpa

planted as an ornamental at low elevation near Rossland, B.C. The condition of the

sample precluded a definitive identification of the pest. In 2009, symptoms of branch

attack by A. piceae were reported from the Cascades Forest District immediately east of

the pre-2014 quarantine zone. These reports prompted additional surveillance for

A. piceae to determine the current extent of its range in B.C. Initial surveillance efforts

focused on detection of host trees with visible symptoms of attack, such as gouting or the

presence of white woolly masses associated with heavy stem attack. This study reviews

the historical records of A. piceae detections in B.C., the history of provincial regulations

to prevent anthropogenic dispersal of the adelgid to uninfested regions of the province,

and reports new locations in the interior of B.C. where it is now established.

J. ENTOMOL. SOC. BRIT. COLUMBIA 113, DECEMBER 2016 23

METHODS

Historical surveys and collections of Adelges piceae in B.C. Historical collections

of A. piceae documented by the Canadian Forest Service Forest Insect and Disease

Survey (FIDS), Pacific Forestry Centre, Victoria, B.C., and records of detections

extracted from both published and unpublished file reports compiled in the CanFIAS

database by Nealis et al. (2015) were retrieved and combined with locality records for

positive and negative collections of the pest documented in this study to provide an

overview of both the species’ occurrence in B.C., and the areas surveyed at which A.

piceae was not detected. Scatter plots of the positive and negative collections are also

presented to visualize the pattern of spread of the pest in coastal B.C. between 1957 and

1995.

In addition to latitude and longitude, the CanFIAS database provides an estimate of

the spatial accuracy of the locality information for all positive and negative collection

records for both point source collections made by FIDS between 1957 and 1995 and for

records generated from annual survey reports of FIDS, provincial aerial survey reports,

and other miscellaneous reports. The spatial accuracy of CanFIAS records for A. piceae

extracted from the aforementioned reports were compared to those generated during

individual collection events documented by the original collection records to estimate the

spatial accuracy of records derived from both types of records.

History of Adelges piceae Regulation in B.C. Copies of the text of all regulations

enacted under the Plant Health Act of B.C. pertaining to A. piceae were obtained from

the Legislative Library at the Provincial Legislature in Victoria, B.C. Titles of each

regulation and date of enactment, along with comments on the purpose of the Order in

Council (O.1.C.) or changes in the areas regulated, are summarized in Table 1. Maps of

the areas regulated for A. piceae were developed from the descriptions of the areas

regulated by each O.I.C. to illustrate the extent of the area regulated in each change.

Significant changes to the regulations documented in the applicable O.I.C.’s were

summarized.

2011-2014 Surveillance for Adelges piceae. In July and September 2011, Abies spp.

branches exhibiting symptoms of attack by A. piceae were sampled with pole or hand

pruners. Samples were collected from the lower one-third of the crown of trees that

showed evidence of gouting or tree decline in the Coquihalla Summit Recreation Area.

Branch samples were returned to the laboratory and held with the cut ends in water in

buckets at room temperature for approximately 10 days to induce adelgid development

and production of white woolly flocculence. Branch samples were inspected under a 10X

magnifying stereomicroscope for evidence of A. piceae life stages. In October 2013, two

lower branches of both mature trees and advanced regeneration of subalpine fir growing

in the vicinity of Rossland, B.C., were sampled by hand. Samples from individual trees

were bagged separately. Branches were examined under a stereomicroscope within 7

days of collection, and all adelgid life stages recovered were preserved in 95% ethanol. A

subsequent survey was done in the Rossland area and at high-elevation sites across the

southern interior of B.C. in 2014 to assess the extent of A. piceae establishment.

Adelgid samples were forwarded to RGF at the Canadian National Collection,

Agriculture and Agri-Food Canada, Ottawa, for identification. Species identifications

were based on an examination of species morphology, using slide-mounted specimens

and sequencing of the barcode region of the mitochondrial gene Cytochrome C oxidase

subunit I (COI). DNA was extracted from each of the submitted samples, and COI was

amplified, sequenced and compared to a reference library of adelgid sequences (Foottit et

al. 2009). COI sequences for all samples positive for A. piceae were deposited in

GenBank, National Center for Biotechnology Information, U.S. National Library of

Medicine, Bethesda MD, U.S.A. http://www.ncbi.nlm.nih.gov/genbank/.

J. ENTOMOL. Soc. BRIT. COLUMBIA 113, DECEMBER 2016

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26 J. ENTOMOL. SOC. BRIT. COLUMBIA 113, DECEMBER 2016

RESULTS

Historical surveys and collections of Adelges piceae in B.C. The original dataset

extracted from the CanFIAS database on 2 Sept. 2015 contained 480 positive records and

1318 negative records (1.e., records of collections made specifically to detect the adelgid

in which it was not found) for A. piceae. Two hundred and twelve of the positive records

documented actual collection events with the remaining 268 records being derived from

published and unpublished reports. The mean (+ standard deviation) and maximum

spatial accuracy of records derived from collection events were 6.18 + 2.1 km and 8.8

km, respectively, while the mean and maximum spatial accuracy of records derived from

literature were 67.9 + 61.8 km and 219.5 km, respectively. Sixty-five of the positive

records extracted from reports or literature exceeded the maximum spatial accuracy for

collection events (8.8 km) and all but one were excluded from mapping. The single

literature record that exceeded the spatial accuracy limit by 1.08 km was found to

represent a valid collection made on urban ornamentals in Penticton in 1967 (Wood

1968) and was retained. The final dataset of positive records from CanFIAS consisted of

the 212 collection events and 205 records generated from the literature. These occurrence

records, as well as the locations of positive collections obtained in this study, are mapped

in Fig. 1. Similarly, 47 literature records that exceeded the spatial accuracy limit and one

literature record based on damage only included in the negative collection records for A.

piceae extracted from the CanFIAS database were excluded from mapping. In total, 1270

CanFIAS negative records were mapped along with the 10 negative collection records

derived from this study to document the areas surveyed for A. piceae at which the pest

was not detected (Fig. 2).

Washington

kilometers

Figure 1. Location of positive detections of Adelges piceae in British Columbia between

1958 and 2014; locations denoted by red circles denote positive collection records extracted

from the CanFIAS database (Nealis et al. 2015); locations denoted by red squares are derived

from this study (see Table 2). The two collections denoted by red triangles represent

detections on urban ornaments that were eradicated.

J. ENTOMOL. SOC. BRIT. COLUMBIA 113, DECEMBER 2016 a

We also mapped the positive and negative detections of A. piceae generated from

point source collection data pooled with records with equivalent spatial accuracy derived

from survey reports and literature from 1958-1998 to illustrate the spread of A. piceae in

coastal B.C. (see Fig. 3). The importance of the latter data sources in documenting spread

is illustrated in Fig. 3b. Records to the northwest of the dashed line on the mainland

represent areas infested by the adelgid, as determined by surveys records documented in

unpublished reports (Ruppel and Allen 1964, 1965) that were not reported in collection

records. A chronology of significant range expansions of the adelgid in coastal B.C. are

summarized in Table 1.

426° 426° Aza Az 420° 118°

Figure 2. Locations surveyed for Adelges piceae in British Columbia between 1958 and

2014, at which the pest was not detected; locations denoted by unfilled circles denote

locations negative collection records extracted from the CanFIAS database (Nealis ef al.

2015); locations denoted by unfilled squares are derived from this study (see Table 3).

In the two years following the discovery of A. piceae in North Vancouver, populations

were detected in the mountains north of Vancouver and across Howe Sound, as well as at

a few localities near Victoria on Vancouver Island (Table 1; Fig. 3a). From 1959-1964,

infestations were discovered on the mainland as far northwest as Jervis Inlet, on the

Sunshine Coast, and as far east as Alouette Lake, in the Fraser Valley. On Vancouver

Island, expansion was limited to a few additional populations discovered on the west

shore of Saanich Inlet (Table 1; Fig. 3b). By 1969, additional infested stands had been

discovered on the east coast and interior of Vancouver Island as far north as Nanaimo,

and a significant northward expansion of the infested area in the Harrison Lake drainage

had been found on the mainland (Table 1; Fig. 3c). Between 1970 and 1979, the range of

A. piceae expanded slightly eastward on the mainland, and a further expansion to the

north within the quarantine zone was detected on Vancouver Island (Table 1; Fig. 3d). By

1989, the adelgid had been found in stands near the quarantine zone boundary on

Vancouver Island and near Powell River on the mainland. However, the most significant

event during the decade was the detection of mature stands of A. amabilis infested by the

adelgid on West Thurlow Island (WTI), between northern Vancouver Island and the

adjacent mainland coast, 100 km and 140 northwest of previously detected infestations

on the mainland and Vancouver Island, respectively (Table 1; Fig. 3e). The last

28 J. ENTOMOL. Soc. BRIT. COLUMBIA 113, DECEMBER 2016

collections of A. piceae documented in the CanFIAS database were made between 1990

and 1998. Expanded surveys to delimit the range of the adelgid on northern Vancouver

Island after the detection of infestations on WTI (Fig. 3e) led to the detection of A. piceae

as far north of Campbell River (Table 1; Fig. 3f). On the mainland, expansion of A.

piceae to both the north and east was detected in the Fraser River drainage, with the

adelgid being found for the first time east of the Fraser River. The adelgid had also

expanded northward along drainages emptying into the Squamish River and was present

beyond the 1992 regulated area at multiple locations in the Birkenhead Lake area (Table

1; Fig. 3f). After the completion of the 1995 survey year, organized surveys for forest

pests were no longer conducted by the Canadian Forest Service. In subsequent years,

surveys for A. piceae were conducted by provincial forestry staff. The single record from

1998 in the CanFIAS database was submitted by provincial forest service staff and

documents the presence of A. piceae on Texada Island in the Strait of Georgia, south of

Powell River (Fig. 3f).

History of Adelges piceae Regulation in B.C. By 1965, A. piceae was recognized as

a serious potential threat to Abies in B.C., and steps were taken to manage its impact

through establishment of quarantine regulations, supported by surveys of infestation

boundaries and damage, a ban on movement of logs from infested areas during periods

when the adelgid was actively reproducing, and expansion of research programs (Vyse

1971). These activities were cost-shared between the federal and provincial governments

and initially were enabled provincially through enactment of Order in Council (O.LC.)

1137 (14 Apr. 1966). They were completed the following year under a second agreement

(O.L.C. 2363, 25 July 1967; Table 2). The collaborative surveys enabled by these O.1.C.’s

led to the discovery of limited northwards range expansion on southern Vancouver Island

and no range expansion on the Lower Mainland. A benefit of these surveys was the first

extensive documentation of stands in the Lower Mainland and on Vancouver Island that

were free of A. piceae infestations (Table 1; Fig. 3d). Additionally, these extensive

surveys led to the discovery of A. piceae in Penticton, B.C.

Adelges piceae was first regulated under the Plant Protection Act (chapter 287 of the

Revised Statutes of British Columbia, 1960) with the approval of B.C. Reg. 58/66 (Table

2). The regulation prohibited the shipment or transport of any living Abies spp., as well

as the production of all Abies species for commercial purposes, eliminating the

production of Abies seedlings for reforestation, as well as the commercial production of

Abies species grown as ornamentals or Christmas trees. The initial regulation applied

province-wide to protect the highly susceptible A. Jasiocarpa in the high-elevation

Interior forests of B.C. and slow the spread of the insect (Wood 1968). Abies seedling

stocks at forest nurseries were destroyed and operational planting of all Abies ceased

with the imposition of the 1966 regulation (Vyse 1971), even though species of Abies

such as Pacific silver fir (A. amabilis) were desirable for silviculture at mid-elevations in

coastal forests where other conifers frequently failed to establish (Carrow 1973).

In 1977, the provisions of B.C. Reg. 58/66 were rescinded by O.1.C. 44 and replaced

by B.C. Reg. 7/77, the British Columbia Balsam Woolly Aphid Regulations, 1976 (Table

2). Annual permits were required to grow and sell Abies provincially and the first

regulated area, based on known infestations in Ranger Districts (Fig. 4a), was

established. Movement of all Abies species grown within the regulated area to areas

beyond its boundaries was prohibited, as was movement of cut trees or foliage of Abies

spp. between January 31 and November 1. The latter regulation allowed for the

movement of cut Christmas trees and foliage after research demonstrated that the adelgid

did not survive on cut trees (Woods 1967).

New finds of the adelgid beyond the boundaries of the area regulated by B.C. Reg.

7/77 (Table 1) led to an expansion of the regulated area (Table 2; Fig. 4b) with the

adoption of B.C. Reg. 414/92 established by O.1.C. 1604, approved on October 22, 1992

(Turnguist and Harris 1993). Movement of trees (living trees with roots, including both

seedlings and those produced by tissue culture) from inside to outside of quarantine area

J. ENTOMOL. SOc. BRIT. COLUMBIA 113, DECEMBER 2016 29

-129° “127° “125° -123° “121° = -429° -127° ~125° -123° “121°

50°

49° 49°

50°

49°

1958-79

50°

49°

48° q RUC Seba Wi eas. ee :

-129° “127° ~125° “123° ~121° = «429° -127° “425° -123° “121°

Figure 3. Expansion of Ade/ges piceae in coastal British Columbia between 1957 and 1995,

as documented by positive and negative collection records (denoted in red and black,

respectively), extracted from the CanFIAS database (Nealis et a/. 2015). Islands between

Vancouver Island and the mainland have been omitted for clarity. Red dashed lines delimit

areas in which A. piceae has been detected: a. collections recorded in 1958 and 1959; b.

cumulative collections between 1958 and 1964. Positive records to the northwest of the

dashed line on the mainland are derived from aerial survey records documented in Ruppel and

Allen (1964, 1965). Note significant expansion of the area infested to the southeast and

northwest; ¢. cumulative collections between 1958 and 1969. Expansion to the northwest is

evident on southern Vancouver Island and to the northeast on the mainland; d. cumulative

records between 1958 and 1979; e. cumulative records between 1958 and 1989. Populations

on the mainland and on Vancouver Island expanded to the northwest, and a satellite

population was discovered on West Thurlow Island (WTI); and f. cumulative records from

1958 to 1998. Between 1990 and 1995, survey efforts documented significant northward

expansion of A. piceae in the Fraser, Lillooet and Squamish river drainages on the mainland,

as well as expansion to the northwest along the east coast of Vancouver Island to Campbell

River.

30 J. ENTOMOL. Soc. BRIT. COLUMBIA 113, DECEMBER 2016

boundaries continued to be prohibited; however, the modified regulations allowed the

movement of logs of Abies spp. out of a quarantine zone if the logs were transported and

stored in water and promptly processed as research had found that they posed minimal

risk for dispersal of the adelgid (Atkins and Woods 1968). The sale of cut trees or foliage

continued to be prohibited in B.C. between January 31 and November 1.

In 2000, a minor change was made to the Balsam Woolly Adelgid Regulation (Table

2). B.C. Reg. 169/2000, enabled by O.1.C. 726/2000, eliminated the need for the tagging

of trees offered for commercial sale to identify that they had been grown under permit.

~428° ~126° -124° “122° -120° -118° 116°

@ Williams Lake

50 100 450

kilometers

~128° -126° “124° -122° -120° -118°

Figure 4. Historical change in areas of southwestern British Columbia regulated for Ade/ges

piceae: a. area regulated in 1977. As no official record of the boundaries of the “Ranger

Districts” named in Order in Council (O.1.C.) 1977-44 was retained, the area regulated on the

mainland was approximated from other sources; b. area regulated by O.I.C. 1992-1604; and ec.

area regulated by O.I.C. 2006-493 (light grey) and the area added by O.I.C. 2014-361 (dark

grey). The locations of the Adelges piceae detections at Falls Creek and Rossland are

designated by unfilled and filled black squares, respectively.

With the detection of new finds east of the known distribution (Table 1), the Balsam

Woolly Aphid Regulation was amended by B.C. Reg. 213/2006, established by O.LC.

493/2006. The definition of “log” was changed to clarify that branches were not

permitted on logs, and the definition of “quarantine zone” was repealed and replaced by

“quarantine area” (described in the regulation’s appendix). The only substantive change

to the regulation was that Appendix A of B.C. Reg. 414/92 was repealed and an expanded

quarantine area defined in the Appendix to B.C. Reg. 213/2006 was created (Table 2; Fig.

J. ENTOMOL. Soc. BRIT. COLUMBIA 113, DECEMBER 2016 31

4c). The most recent amendment to the Balsam Woolly Adelgid Regulation, B.C. Reg.

137/2014, of June 30, 2014, added the Cascade Forest District to the quarantine area after

recent surveillance indicated establishment of the adelgid beyond the boundaries of the

2006 regulated area (Table 2; Figure 4d).

Table 2

Summary of the provincial Orders in Council and regulations arising from each Order in

Council (O.I.C.) enacted against Adelges piceae. Changes to the area regulated for Adelges

piceae by specific O.I.C. are noted in the comments.

Order in Council!

Yea Numbe _§ Enactment

r r Date

Regulati

Comments

(196 | 460 16Feb. | B.C. Reg. _ All of British Columbia

| 6 1966 58/66

196 1137 Cost sharing agreement

6 for surveys and

| acquisition of

| contaminated nursery

stock

er a ah ST Bc ne De 9 ae Sa SANG. avis Binds seek ea ae ta

196 2363 25 Jul. 1967 Cost sharing agreement

my for surveys and

| acquisition of

| contaminated nursery

| stock

197 7 Jan. 1977 British Columbia All portions of 10

7 Balsam Woolly Aphid | mainland and Vancouver

| Regulations, 1976 Island Ranger Districts

| and the infested portions

of two Ranger district in

| Vancouver Region (see

Fig 3a)

| 199 1604 B.C. Reg. | Balsam Woolly Adelgid

| 2 414/92 | Regulation

/ 200 726 18 May B.C. Reg. | Balsam Woolly Adelgid

| 0 2000 |: 169/2000 | Regulation

200 493 13 Jul. 2006 | B.C. Reg. | Balsam Woolly Adelgid | South Coast Forest

6 213/2006 | Regulation Region and West Coast

2 Forest Region

201 361 5 Jun. 2014 | B.C. Reg. | Balsam Woolly Adelgid | Cascades Forest District;

4 137/2014 | Regulation South Coast Forest

Region and West Coast

Forest Region

137/204, June 30 2014, pertaining to Adelges piceae were accessed at the following URL: http://

www.bclaws.ca/Recon/document/ID/freeside/11_ 414 92 (accessed 10 October 2015).

* Amendment to B.C. Reg. 414/92

32 J. ENTOMOL. SOC. BRIT. COLUMBIA 113, DECEMBER 2016

2011-2014 Surveillance for Adelges piceae. Locations sampled between 2011 and

2014 at which the presence of Adelges piceae was confirmed are documented in Table 3

and mapped in Figure 1; locations negative for the presence of A. piceae are presented in

Table 4 and mapped in Figure 2. The presence of the adelgid in the Coquihalla Summit

Recreation Area in the Cascades Forest District is not surprising, as the adelgid was

known to be present in high-elevation coastal forests to the west of the detection. The

Cascade Forest District was added to the quarantine area in June 2014 as a result of the

surveys conducted. However, the presence of established populations of A. piceae in the

Southern Interior of the province around Rossland (Table 2; Fig. 1), more than 200 km

east of known infestations in B.C., was unexpected.

DISCUSSION

The discovery of A. piceae at multiple locations near Rossland is the first detection of

this pest in the subalpine-fir forests in the interior of the province. Infested stands were

detected in this study up to 9.5 km north and 3.8 km south of Rossland. Although

A. piceae was detected in the southern Okanagan in 1967, the incursions were restricted

to a few non-native urban ornamentals (A. alba and A. concolor) in Oliver and Penticton,

and the populations were subsequently eradicated. Surveys of subalpine fir from

Penticton south to the U.S. border provided no evidence that A. piceae had dispersed

from the two low-elevation urban infestations into natural stands (Wood 1968; Wood et

al. 1968). Symptoms of attack were not evident at the majority of the locations surveyed

for A. piceae in the Rossland area. Severe gouting, stem attack and limited mortality

were only evident at the golf course in Rossland, the lowest-elevation site sampled.

Gouting was also documented on subalpine fir at the Highway 3B location (Table 2). At

all other locations sampled, gouting was not evident, although a slight thickening of

some nodes was noted at two other locations (Table 3; LH, personal observations).

It is unlikely that the A. /asiocarpa growing at the Rossland golf course was infested

with A. piceae when planted some 60 years ago, as subalpine fir is highly susceptible to

infestations and generally succumbs rapidly after infestation. Those subalpine firs that

succumb first after infestation by A. piceae are often growing on the best sites (stream

bottoms, benches and around meadows), with the severest damage occurring at the

lowest elevations starting at around 915 meters (3000 ft.; Mitchell 1966), a description

that closely resembles the golf course site. The source of this infestation could not be

determined with any certainty; however, it is suspected that it has arisen from aerial

dispersal of crawlers from nearby counties in Washington or Idaho, where populations of

A. piceae have been documented (Liebhold et al. 2015). Alternatively, because

asymptomatic A. lasiocarpa infested with A. piceae were easily found within Rossland

and adjacent forested areas, the possibility remains that the adelgid may have originally

been introduced on ornamental firs transported from other infested regions of the

province. Unlike earlier detections of A. piceae on non-native urban ornamentals in the

southern Okanagan (European silver fir, Abies alba Mill. and white fir, A. concolor;

Wood 1968; Wood ef al. 1968), plantings of non-native firs were not observed during

surveys in Rossland. In addition, the latter scenario is improbable, as quarantine

regulations have restricted the movement of ornamental Abies spp. from infested areas of

the province to uninfested areas since 1966. While quarantine restrictions were also in

place in 1967, the infested A. alba detected at Oliver were imported from Europe 29

years prior to the discovery of the adelgid. Neither the origin nor date of planting for the

two white firs in Penticton was determined, although the trees were of similar height and

diameter at breast height to the silver firs in Oliver, suggesting that they also were

planted prior to the introduction of any quarantine regulations. Neither of these

ornamental firs were killed by A. piceae, and gouting is not expressed on either species in

the Pacific Northwest (Mitchell 1966).

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34 J. ENTOMOL. SOC. BRIT. COLUMBIA 113, DECEMBER 2016

The detection of A. piceae at Falls Creek in the Cascades Forest District (Table 2)

was the first detection of the adelgid east of the Coast Mountains in B.C. The collection

site is to the northeast of the previously most-eastern collections of A. piceae (Fig. 1, 3f),

and appears to represent an eastward range expansion in B.C. Gouting was evident on

A. amabilis infested by A. piceae at the sites sampled.

The implementation of provincial regulations defining regulated areas for A. piceae in

B.C., which are supported by federal restrictions, has been successful in slowing the

expansion of the infestation boundary of A. piceae in southwestern B.C. (Coulson and

Witter 1984). As the range of the adelgid infesting Pacific silver fir and grand fir in the

coastal forests of B.C. has slowly expanded (Fig. 3), the quarantine boundaries have been

expanded and strategies have been developed to produce and distribute clean Abies

seedlings for reforestation to prevent anthropogenic dispersal of A. piceae. Additionally,

the growth, sale and distribution of ornamental Abies spp. within the province has been

restricted, as have imports of all Abies spp. into B.C. to prevent redistribution of the pest

to uninfested regions.

The records presented in this study document the known distribution of A. piceae in

B.C. from historical survey records and recent detections. The records noted in Anon.

(1938) indicate that A. piceae was introduced into southwestern B.C. on ornamental

Abies before 1937, at least 20 years prior to the first reports of damage in North

Vancouver (Silver 1959). Vyse (1971) states that the adelgid was first introduced into

southwestern B.C. in about 1938, although no evidence is provided to document that

assertion. The rapid expansion of the infested area detected in the first seven years after

its initial discovery on the mainland (Fig. 3a, b) may not represent active dispersal and

rapid range extensions of the pest from the infestations detected in 1957 (Silver 1959),

but rather may have arisen from the detection of previously established populations

introduced in the late 1930s. It is apparent from the historical records that the distribution

of this destructive forest pest was actively expanding when federal survey efforts ceased

after 1995 (Fig. 3f). The recent detections at Falls Creek demonstrate that this adelgid has

continued to expand its range inland beyond the coastal forests during the last two

decades.

The historical detections of the adelgid in mature stands have generally occurred well

after its initial establishment in the stand—most often when visible damage such as

gouting or mortality is discovered at any one location. Subsequent delimitation surveys

beyond the initial find have often demonstrated that the pest is more widely distributed.

The detection of A. piceae beyond the regulated area on Vancouver Island in 1993 led to

the discovery of multiple infested stands well north of the boundary in subsequent years

(Table 1). Negative survey records from northern Vancouver Island made after the

discovery of populations on WTI (Fig. 3e, f) suggest that the adelgid was not yet active

in those areas. However, difficulty of access has limited survey efforts on the adjacent

mainland between WTI and known infested areas near Powell River first discovered in

1986. It is highly probable that A. piceae was established at least as far north as the

mainland adjacent to WTI by 1987. Vyse (1971) estimated that over the two decades

ending in 1987, the maximum range expansion of A. piceae to the northwest along

mainland would reach the Campbell River area. The WTI infestations exceed the

estimated maximum range expansion along the coast under the most pessimistic scenario

discussed by Vyse (1971) by more than 50 km.

The first detections of A. piceae in Interior A. lasiocarpa stands near Rossland pose

serious challenges to the quarantine management of this pest in B.C. The cryptic nature

of A. piceae in the initial stages of an infestation makes detection of attacked stands and

thus delimitation of boundaries for regulated areas extremely difficult. Although Mitchell

(1966) noted that “significant gouting always accompanies the decline of subalpine fir,

but is often not conspicuous because the trees die so quickly”, gouting was not apparent

at many of the locations sampled near Rossland. The absence of such readily apparent

symptoms of attack makes detection of infested trees considerably more difficult,

J. ENTOMOL. Soc. BRIT. COLUMBIA 113, DECEMBER 2016 35

requiring the collection and microscopic examination of branch samples for presence of

the adelgid under bud scales, at the base of staminate flowers, or at branch nodes. The

massive numbers of white woolly masses associated with stem attack by A. piceae are

more visually apparent than life stages associated with branch attack; however, stem

attack usually begins high in the crown on subalpine fir and progresses down the stem

(Mitchell 1966), also making the initial stages of such attacks difficult to detect. The

presence of a native adelgid, Pineus abietinus (Underwood & Balch), attacking true firs

in B.C. (Underwood and Balch 1964; Maw et al. 2000) further complicates recognition

of stem attack by A. piceae, as the former species also develops dense populations on

Abies stems.

Current quarantine regulations were developed to address potential sources of

anthropogenic dispersal of A. piceae during harvest and reforestation activities in coastal

forests, as well as to address that risk also posed by commercial distribution of

potentially infested ornamental Abies spp. in urban areas. Regulations related to the

movement of logs from within a regulated area will likely need to be modified, should

quarantine restrictions be considered for management of this pest in Interior stands.

Current regulations requiring the transport and storage of Abies logs in water are not

feasible in the Interior of the province. Furthermore, current restrictions related to the

production of seedlings for reforestation rely on the production of clean nursery stock in

nurseries situated well beyond areas of known infestation to prevent infestation of the

seedlings by aerially dispersed crawlers. To ensure that any sites used for the production

of reforestation seedlings are free of populations of A. piceae, true firs present in the

surrounding forests or cultivated as ornamentals near nurseries should be surveyed for

the presence of A. piceae. Ideally, nursery stock should be produced in nurseries situated

well beyond the range of native fir stands or ornamental plantings. Should this not be

feasible, we recommend the use of sentinel plantings of Abies species that easily express

persistent and apparent symptoms of infestation (e.g., gouting) by low numbers of A.

piceae. Trees to be planted as sentinels should be grown from seed at nurseries remote

from any populations of A. piceae in B.C.

Subalpine fir in southeastern B.C. is restricted to the upper elevations in the

mountains, resulting in an extremely patchy distribution with very limited road access in

most areas. Given the constraints noted above, development of a survey strategy will be

extremely difficult. Should surveys for A. piceae be undertaken in subalpine fir forests,

we recommend that presence of the pest be documented with properly preserved samples

of life stages suitable for both morphological (i.e., preserved in 70% ethanol) and

molecular identification (preserved in 95% ethanol). These samples are also essential to

separate attack by Ade/ges piceae from that of the non-damaging native species, Pineus

abietinus (Cook et al. 2010).

ACKNOWLEDGEMENTS

We are grateful to Don Hepner for providing information on the infestation of balsam

woolly adelgid near Falls Lake. Troy Kimoto, David Holden, Tracy Hueppelsheuser, and

Don Hepner assisted with surveys near Falls Lake, and D. Hepner assisted in the

Kootenays. Amanda Biernacka-Larocque, J-P Nadeau, and E. Maw (Agriculture and

Agri-Food Canada, Ottawa carried out the sequencing and compilation of the barcodes.

Gurp Thandi (Natural Resources Canada) is thanked for his assistance in generating the

maps of A. piceae distribution. Funding for the genetic analyses for the 2012-2014

collections was provided by the Interdepartmental Genomics Research and Development

Project on Quarantine and Invasive Species to RF and LH.

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J. ENTOMOL. SOc. BRIT. COLUMBIA 113, DECEMBER 2016 39

Pollen preferences of two species of Andrena in British

Columbia’s oak-savannah ecosystem

JULIE C. WRAY! and ELIZABETH ELLE!

ABSTRACT

Although understanding the requirements of species is an essential component of their

conservation, the extent of dietary specialisation is unknown for most pollinators in

Canada. In this paper, we investigate pollen preference of two bees, Andrena

angustitarsata Vierick and A. auricoma Smith [Hymenoptera: Andrenidae]. Both

species range widely throughout Western North America and associated floral records

are diverse. However, these species were primarily associated with spring-blooming

Apiaceae in the oak-savannah ecosystem of Vancouver Island, BC, specifically

Lomatium utriculatum [Nutt. ex Torr. & A. Gray] J.M. Coult. & Rose, L. nudicaule

[Pursh] J.M. Coult. & Rose, and Sanicula crassicaulis Poepp ex. DC. Floral visit

records and scopal pollen composition for these species from two regions on

Vancouver Island indicate dietary specialisation in oak-savannah habitats where

Apiaceae are present. Both species were also caught in low abundances in residential

gardens where Apiaceae were scarce, sometimes on unrelated plants with

inflorescence morphology similar (to our eyes) to Apiaceae. Further study of these

species is needed to understand whether preferences observed locally in BC exist

elsewhere in their range. Our findings contribute to understanding pollen preference in

natural and urban areas, and highlight an important factor to consider for species-

specific conservation action in a highly sensitive fragmented ecosystem.

Key Words: Andrenidae, Apoidea, oligolecty, pollen preference

INTRODUCTION

Relationships between flowering plants and bees (Hymenoptera: Apoidea, Apiformes)

range from extreme pollen specialisation, or oligolecty, to extreme generalisation, or

polylecty. Perhaps as many as half of all non-parasitic bee species exhibit dietary

specialisation at some level, but for most bees the extent of specialisation is unknown

(Cane and Sipes 2006; Michener 2007). Bee species are considered to exhibit narrow

oligolecty if females consistently provision offspring with pollen from a small related

clade of plants (typically genus level; Linsley and MacSwain 1958; Miller 1996; Cane

and Sipes 2006). Broad oligolecty could potentially comprise a larger number of related

plant species or genera, but differs from polylecty in that many to most of the available

pollen sources are not utilized (Cane and Sipes 2006). Facultative oligolecty occurs in

some bee species, which normally specialize on a host plant, but may use non-host pollen

when the usual host is unavailable (e.g., Williams 2003; Sipes and Tepedino 2005). Use

of non-host pollen by oligolectic species can sometimes have substantial fitness

consequences (Praz et al. 2008a), which may have conservation implications and explain

why specialists tend to be more sensitive to disturbance (Elle et al. 2012).

Evaluating specialisation can be challenging. The identity of plant species from

which bees are collected has frequently been used to describe foraging preferences, but

we have known since the landmark work of Charles Robertson that bees visit a broader

array of flowers for nectar than for pollen (Robertson 1925). Similarly, foraging

preferences of male bees, which do not provision nests with pollen (they forage only for

1 Evolutionary and Behavioural Ecology Research Group, Department of Biological Sciences, Simon

Fraser University, 8888 University Drive, Burnaby, British Columbia, Canada, V5A 1S6; (778)

782-4592, eelle@sfu.ca

40 J. ENTOMOL. Soc. BRIT. COLUMBIA 113, DECEMBER 2016

nectar), should not be considered when evaluating dietary specialisation. Floral records

that do not include information on gender and behaviour simply have limited utility

(Cane and Sipes 2006). Instead, it is important to consider not only constancy of floral

visits by female non-parasitic bees, but also the composition of pollen they collect for

offspring provisioning.

Habitat loss and fragmentation is predicted to have negative effects on biodiversity

unless organisms can use the surrounding landscape (or “matrix”) for food or nesting

resources (Dunning ef al. 1992; Fahrig 2001; Wray and Elle 2015). Dietary specialists

are expected to be negatively impacted by habitat conversion because the plant species

upon which they rely are generally not available in the non-habitat matrix. In studies

examining the effects of habitat loss on pollinating insects, specialists consistently

decline with reduction in natural habitat (Steffan-Dewenter and Tscharntke 2000; Burkle

et al. 2013). Britain and the Netherlands experienced parallel declines in bee-dependent

plants and specialist bees (Biesmeijer et al. 2006), in some cases, because the plant

species required by the bees have been locally extirpated. Documenting dietary

specialisation, therefore, is important for recognition of threats to bees of particular

conservation concern, the plants upon which they rely for offspring provisioning, and

potentially how reproductive output of those plants would be impacted by pollinator loss.

Such research can be especially interesting in human-dominated landscapes where host

plants may vary in availability (Maclvor et al. 2014).

We investigated potential pollen specialisation of two mining bees, Andrena

angustitarsata Vierick and A. auricoma Smith (Hymenoptera: Andrenidae) in a highly

fragmented oak-savannah ecosystem. Less than 5% of this habitat remains due to

residential expansion, agricultural development, and the introduction of invasive species

(Fuchs 2001). We noticed that collections of these bees at our study sites were primarily

from two species of Lomatium (Apiaceae), spring gold (Z. utriculatum [Nutt. ex Torr. &

A. Gray] J.M. Coult. & Rose) and Indian consumption plant (L. nudicaule [Pursh] J.M.

Coult. & Rose), suggesting the bees may be oligolectic. However, the range of both bees

comprises the majority of Western North America, and the floral records included in

species descriptions are quite diverse (LaBerge and Ribble 1975; LaBerge 1989). We

have also more recently been working in urban and suburban neighborhoods near oak-

savannah fragments, where the host plants are not available (Wray and Elle 2015). We

were therefore interested in documenting the proportion of visits by A. angustitarsata

and A. auricoma females to Apiaceae (at our field sites, the two Lomatium species and

Sanicula crassicaulis Poepp ex. DC) vs. other plant species, and the composition of

pollen collected by females for nest provisioning, to better understand whether these

Andrena spp. are facultative oligoleges in our region.

METHODS

As part of two larger research projects, bees were collected directly from flowers in

oak-savannah fragments of the Cowichan Valley (“North OS”, 2008-2010), and oak—

savannah fragments and urban residential gardens on the Saanich Peninsula (“South OS”,

“South Gardens”, 2012) of Vancouver Island, British Columbia (Fig. 1, Gielens 2012;

Wray and Elle 2015). Bees collected from flowers of a single plant species (or genus in

gardens, where botanical cultivars were common) were euthanized in the same cyanide

tube during the duration of a sample period. For North OS sites, two 15-minute plant-

species-specific samples were collected on each day sampling occurred (Gielens 2012).

In South OS sites and gardens, two 30-minute samples were taken on each day of

sampling, with collections made from any plant species in flower, although collected

insects were still kept separate by plant species (Wray and Elle 2015). For both studies,

collection dates comprised the majority of the flowering season of our sites, from April to

July. Bees of the species of interest were collected from 11 April to 27 June, depending

on year and site, with the majority collected in May. Because of the variability in

J. ENTOMOL. SOc. BRIT. COLUMBIA 113, DECEMBER 2016 41

numbers of bees collected at different sites, we calculated the proportion of 4A.

angustitarsata and A. auricoma collected on different plant species within a geographic

region (i.e., North OS, South OS, South Gardens), rather than by sites within a region.

C)Nortthos |

B) Locator map

Andrena angustitarsata present

~ Andrena auricoma present

Neither species present

een

toe

Figure 1. Maps illustrating locations of study sites and which focal species were collected at

each. Land is white and ocean is grey. Dashed boxes indicate the area magnified in

subsequent panels. A) Vancouver Island in the broader regional context of British Columbia

(from iMapBC, http://maps.gov.bc.ca/ess/sv/imapbc/). B) Locator map of southern Vancouver

Island, with two regions of study outlined. C) North Oak-Savannah (OS) sites (squares). C)

South OS (circles) and South Gardens (triangles). Symbol colour indicates species collected

(black: Andrena angustitarsta; grey: A. auricoma; split symbols: both species; white: neither).

Female bees were visually examined for the presence of scopal pollen, and a random

sample of those individuals was included in the analysis. Only pollen from the tibial

scopa and propodeal corbicula (i.e., pollen-carrying regions of Andrena) was analysed

because our intention was to investigate the pollen collected for offspring provisioning.

Using pollen from these areas also reduced the likelihood of any contamination by non-

scopal pollen from our cyanide tubes. The pollen load from the scopa of one leg was

scraped into a l-mL centrifuge tube with a sterilized needle. In addition, the leg and

propodeum were washed with 1mL of 70% alcohol. The pollen/alcohol mixture was

centrifuged at 3500 rpm for 5 min, discarding the supernatant after centrifuging. Pollen

pellets were left in centrifuge tubes to dry for 30 min. We added 10uL Caberla solution

(6mL glycerin, 12mL 95% ethyl alcohol, 18mL distilled water, 50mg basic fuchsin stain

[pararosaniline]), and 10uL glycerin to the pollen pellet, and tubes were agitated to

suspend pollen in solution. We transferred the entire 20-uL sample to a glass slide,

covered the pollen solution with a cover slip, and used clear nail polish to seal the edges.

42 J. ENTOMOL. SOc. BRIT. COLUMBIA 113, DECEMBER 2016

We identified pollen under a light microscope at 400X magnification with the

assistance of reference pollen slides made from flowers collected from our sites and from

the SFU Palynology Lab. Pollen grains were identified to “type” at the lowest possible

taxonomic level—species, genus, or family (Table 1). Pollen morphology is often

conserved within a genus or family and so identification to species is frequently not

possible. When pollen was identified to genus or family, but could not be unambiguously

assigned to species using our reference collection, we inferred likely species based on

vegetation-sampling data from our sites (Gielens 2012; Wray and Elle 2015) and include

a note in the table. Pollen grains were counted in four random “transects” of the cover

slip (as in Miiller 1996). Samples with insufficient pollen (<50 grains) were not included

in results. Any pollen types that contributed less than five percent to the total pollen

count on a slide were deleted to account for the possibility of trace contamination (Cane

and Sipes 2006). However when averaged across all samples within a geographic region

(North OS, South OS, South Gardens), some pollen types still contributed less than five

percent to the total pollen count across all slides from that region. In addition to

presenting the average proportion of pollen types for different regions, we include

information on the incidence of “pure” loads (100% host pollen once trace contaminants

comprising <5% removed, as in Cane and Sipes 2006)

Table 1

Description of pollen types, including the family, genera, and species combined into the

different types. When pollen could not be attributed to a particular plant species using our

reference collection, likely species identities listed in the table are based on our vegetation

surveys, as indicated.

Pollen type Family/genera/species included

Apiaceae-1 Lomatium utriculatum, L. nudicaule

Apiaceae-2 Sanicula crassicaulis

Asteraceae Many species possible. Based on our vegetation surveys, this pollen

type could include Achillea millefolium, Balsamorhiza deltoidea,

Bellis perennis, Eriophyllum lanatum, Hypochaeris radicata, and/or

Taraxacum officinale

Brassicaceae Likely Brassica spp., based on our vegetation survey

Rhamnaceae Ceanothus spp. (probably C. thyrsiflorus cv. “Victoria” as this was

commonly planted in gardens)

Sorbus spp. (usually S. aucuparia, according to our vegetation

survey)

Rosaceae-1

Rosaceae-2 Rosa nutkana

Trifolium pratense Trifolium pratense

Trifolium repens Trifolium repens

Other Caryophyllaceae, Ranunculaceae, Unknown-1

RESULTS

Two pollen types that could be assigned via morphology to Apiaceae were detected in

our samples. Based on our reference collection, Apiaceae-1 is consistent with Lomatium

spp., and so could be either L. utriculatum or L. nudicale. The larger grains of Apiaceae-2

J: ENTOMOL. Soc. BRIT. COLUMBIA 113, DECEMBER 2016 43

are consistent with Sanicula crassicaulis (Fig. 2). Inflorescence morphology for these

three species is also shown in Figure 2, and all are comprised of dense heads of small

flowers.

Andrena angustitarsata. We collected a total of 141 females of this species from

oak-savannah fragments in the Cowichan Valley (“North OS’) and on the Saanich

peninsula (“South OS”), and from residential gardens on the Saanich Peninsula (“South

Gardens”; Table 2). Most were collected from flowers of Apiaceae; 89% in North OS

sites, and 98% in the South OS sites (Table 2). In South Gardens, only 11 A.

angustitarsata were collected, none from flowers of Apiaceae. No plants from this family

were blooming in residential gardens during the spring flight period of this bee, although

some Apiaceae genera bloom in gardens in July and August (Astrantia, Eryngium,

Foeniculum, Pastinaca). In gardens, bees were collected from a number of plant species

with diverse floral morphologies, including small flowers densely arranged in compact

heads or umbels (e.g., Ceanothus Linnaeus [Rhamnaceae], Sorbus Linnaeus [Rosaceae],

Spiraea Linnaeus [Rosaceae]) and simple flowers with easily accessible pollen and

nectar rewards (Brassica spp. Linnaeus [Brassicaceae]).

Figure 2. Pollen grains and inflorescence architecture of focal plant species: A) Smaller

pollen grains are Apiaceae-1 (includes both Lomatium utriculatium and L. nudicaule), and

larger grains are Apiaceae-2 (Sanicula crassicaulis); B) L. utriculatum; C) L. nudicaule; D) S.

crassicaulis.

44 J. ENTOMOL. SOC. BRIT. COLUMBIA 113, DECEMBER 2016

We investigated composition of actively collected pollen (from scopae and

corbiculae) for 73 bees (Table 2). Pollen grains counted per slide ranged from 117 to

11,225 grains, with an average of 2,334 grains per slide. Across all North OS sites, A.

angustitarsata pollen loads were comprised primarily of Apiaceae-1 (83%, Lomatium

spp.), followed by Rosaceae-1 (13%, Sorbus spp.). Brassicaceae and Rosaceae-2 each

contributed less than 2% to the total pollen sample, while unknown pollen types

combined into the “Other” category were less than 1% (Fig. 3A). Of the 34 North OS

bees included in the pollen analysis, 16 (47%) had “pure” pollen loads of Apiaceae-1.

In South OS sites, pollen was almost exclusively Apiaceae-1 (98%). Brassicaceae,

Rosaceae-1 and Other each contributed less than 1% to the total pollen composition (Fig.

3C). Of the 31 South OS bees included in the pollen analysis, 28 (90%) had “pure”

pollen loads.

In South Garden sites, there was no Apiaceae pollen in samples that could not be

attributed to contamination (e.g., 1-2 grains out of several hundred counted per sample).

Instead, samples were comprised largely of Rosaceae-1 (42%), Brassicaceae (21%),

Rhamnaceae (17%), and Trifolium repens (13%, Fig. 3E). Trifolium pratense contributed

less than four percent to the total, and Other types less than one percent.

Andrena auricoma. There were a total of 49 female A. auricoma collected from oak-

savannah fragments (19 North OS sites, 30 South OS sites). Only two females were

collected from South Garden sites. Females were predominantly collected from flowers

of Apiaceae (95% in North OS, 83% in South OS), the majority from plants in the genus

Lomatium (Table 2).

Pollen composition was investigated for a total of 32 female bees. We counted

between 58 and 4,984 grains per slide, with an average of 1,465 grains. The two females

collected from South Garden sites were not included in pollen analysis, due to low

numbers of pollen grains (<50 grains counted). In both OS regions (North and South),

Apiaceae-l1 was the dominant pollen type (North: 61%, South: 75%) followed by

Apiaceae-2 (North: 37%, South: 19%; Fig. 3B, D). In North OS sites, Rosaceae-1 and

Trifolium repens each contributed less than 2% (Fig. 3B). In South OS sites, Rosaceae-1

contributed 5% to total pollen counts, and Asteraceae less than 1% (Fig. 3D).

Of the 11 female bees included from North OS, 9 of 11 (82%) had “pure” pollen

loads (combined Apiaceae | and 2). Of the 19 females included from South OS sites, 17

(89%) had “pure” loads.

DISCUSSION

We found evidence of preference for Apiaceae pollen for two bee species in oak-

savannah habitats on Vancouver Island. Female Andrena angustitarsata and A. auricoma

predominantly visit Lomatium spp. and Sanicula crassicaulis in this region, and pollen

collected to provision nests is largely from the Apiaceae. Dietary specialization 1s

normally considered a characteristic of species, rather than something that varies across a

species’ range. The published floral host records for these species are diverse (LaBerge

and Ribble 1975; LaBerge 1989), suggesting polylecty. Further evaluation of pollen

collection behaviour throughout the ranges of these species would be useful, as 1t would

allow visits for nectar or by male bees to be distinguished from visits by females actively

provisioning nests with pollen. Confirmation of observed preferences, an evaluation of

preferences in other parts of the species’ ranges, and clarification of the ecological

conditions that promote these preferences in otherwise polylectic bees is clearly required.

It is unlikely that these species are simply foraging on Apiaceae due to

disproportionate availability of Apiaceae flowers. For example, during the spring bloom

of Lomatium spp. and Sanicula crassicaulis, these plants combined contributed less than

5% to the total cover of blooming flowers recorded in surveys of South OS sites (Wray

and Elle 2015). In addition, other pollen sources (e.g., common camas, Camassia

quamash [Pursh] Greene [Asparagaceae]; great camas, Camassia leichtlinii [Baker] S.

J. ENTOMOL. SOC. BRIT. COLUMBIA 113, DECEMBER 2016 45

Watson [Asparagaceae]) were abundant and available and are consistently used by other

solitary mining bees, mason bees, social bumble bees, and honey bees (Gielens 2012;

Wray and Elle 2015). Instead, our results indicate a preference for Apiaceae for these two

mining bees, particularly plants in the genus Lomatium.

Table 2

Total numbers of Andrena angustitarsata and Andrena auricoma collected from different

study locations. Of the total collected, we state the number (and percentage, in brackets) of all

collected bees that were caught foraging on plants in the Apiaceae, and within Apiaceae, the

total that were collected from Lomatium utriculatum and L. nudicaule combined. The number

randomly chosen for pollen analysis is also indicated.

Andrena angustitarsata Andrena auricoma

North OS South OS South NorthOS South OS

Gardens

Total number 99 42 1] 19 30

collected

Collected from 88 41 (97.6%) 0 (0%) 18 (94.7%) 25 (83.3%)

Apiaceae (88.9%)

Collected from 85 40 (95.2%) 0 (0%) 11. (87.9%) .24:(70.0%)

Lomatium (85.9%)

Included in pollen 34 Sil 8 1] 21

analysis

Documenting dietary specialisation and/or preference is highly dependent on

considering pollinator sex, and evaluating pollen loads used for rearing offspring (Cane

and Sipes 2006). Andrena angustitarsata is considered a polyletic species, with over

2000 females and 1400 males collected on plants from 61 genera in 24 families in

Western North America and included in the revision by LaBerge (1989). These

collections include records from multiple species of Lomatium and Sanicula, as well as

Ceanothus, Ranunculus Linnaeus (Ranunculaceae), Salix Linnaeus (Salicaceae),

Brassica, and Spiraea. Similarly, Andrena auricoma has been collected from flowers of

44 genera and is also documented to be polylectic (LaBerge and Ribble 1975). The most

frequent collections in decreasing order were from Ranunculus, Descurainea Webb and

Berth (Brassicaceae), Salix, and Potentilla Linnaeus (Rosaceae). However, these

collection records do not indicate the sex of the specimen, foraging behaviour (nectar vs.

pollen foraging), nor has the pollen collected by female bees been examined. Without

this information, one cannot confidently assume these species are polylectic or

oligolectic. As such, it is difficult to determine whether Apiaceae preference by 4.

angustitarsata and A. auricoma is simply localised to our study region of oak-savannah

habitat on Vancouver Island, or if it may be more widespread and present in other habitat

types within the broad range of these species. In our region, at least, our data suggest the

species should be considered facultative oligoleges, sensu Cane and Sipes (2006).

Specialist bees are predicted to be more sensitive to the effects of habitat loss and

fragmentation (Davies et al. 2000; Henle et al. 2004), and our study species were caught

in low abundances outside of natural oak-savannah habitat (A. angustitarsata: 11 total at

three South Garden sites, average 1.4/site out of eight total South Garden sites sampled;

A. auricoma: 2 total at a single South Garden site, average 0.25/site). The fitness

consequences for these bees in gardens are not known, but for other species, the

46 J. ENTOMOL. SOC. BRIT. COLUMBIA 113, DECEMBER 2016

consequences vary. For example, Praz et al. (2008a) found in some oligolectic species,

larvae failed to develop on non-host pollen; Haider et a/. (2013) found within- and

among-population variation in offspring development on non-host pollen; and Williams

(2003) found no effect on development for specialist larvae reared on non-host pollen.

However, successful development on non-host pollen does not necessarily change

foraging preference of adults, as Praz et al. (2008b) found that larvae successfully reared

on non-host pollen preferred their normal host in choice-tests as adults. We do not know

if non-Apiaceae pollen would support successful offspring production by these two

species in our region, or if such offspring would subsequently maintain oligolecty; this

should be studied.

Andrena angustitarsata Andrena auricoma

North OS

South OS

WA Apiaceae-1 amm@m™ Rosaceae-1

[ss Apiaceae-2 WZ Rosaceae-2

South Gardens Mmmm Asteraceae @@@@— Trifolium pratense

[== Brassicaceae MES Trifolium repens

Wm Rhamnaceae ([__! Other

Figure 3. Average proportional pollen composition for Andrena angustitarsata and A.

auricoma in North OS (A, B), South OS (C, D), and South Garden sites (E). Numbers in pie

sections indicate the percentage a pollen type contributed to the total pollen count.

We found it curious that bees in gardens were collecting pollen from non-host plants

that appear (to our eyes) morphologically similar to Apiaceae. Lomatium utriculatum has

small flowers arranged in flat umbels, and L. nudicaule and S. crassicaulis have small

flowers arranged in spherical clusters (Fig. 2). In gardens, Sorbus and Ceanothus

comprised a large proportion of the diet of the few A. angustitarsata collected, and have

similar inflorescence architecture to the native host plants. Pollen chemistry has been

invoked as an important cue for oligolectic species (e.g., Miller and Kuhlman 2008), but

floral morphology may also be important if specialists are limited in their ability to

extract resources from flowers with morphology different than that of hosts (Thorp 1979;

Miiller 1996; Williams 2003). The apparent similarity of visual cues between host and

J. ENTOMOL. SOc. BRIT. COLUMBIA 113, DECEMBER 2016 47

non-host flowers suggests an investigation of foraging decisions when Apiaceae are

unavailable would be worth pursuing in future studies.

Our study provides only a baseline of information on pollen preference in A.

angustitarsata and A. auricoma within BC, and clearly further assessment of pollen

provisioning behaviour needs to be done throughout these species’ ranges. Our data do

suggest that, in the geographically restricted oak-savannah ecosystem on Vancouver

Island, L. utriculatum, L. nudicaule, and Sanicula crassicaulis provide vital resources for

A. angustitarsata and A. auricoma. Urban residential gardens in our area support a

diversity of bees, including two species in the Megachilidae that are Asteraceae

specialists (Megachile perihirta Cockerell (Megachilidae) and Osmia coloradensis

Cresson (Megachilidae); Wray and Elle 2015). Homeowners could consider planting

native oak-savannah wildflowers to sustain these, and other specialist bees.

ACKNOWLEDGEMENTS

Emily Helmer and Rolf Mathewes of the SFU Palynology Lab provided pollen

reference slides to complement those made by our research group. Funding was provided

by the Natural Sciences and Engineering Council (NSERC) of Canada (Discovery Grant

to E. Elle) and by a donation from the SFU “Be Bee Friendly” Campaign (D. McKinley,

C. Choi, P. Ficzyez, S. Ghuman, and P. Raimbault). We thank two anonymous reviewers

for their comments on the manuscript.

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J. ENTOMOL. SOc. BRIT. COLUMBIA 113, DECEMBER 2016 49

Relative efficacies of sticky yellow rectangles against three

Rhagoletis fly species (Diptera: Tephritidae) in Washington

State and possible role of adhesives

W. L. YEE! and R. B. GOUGHNOUR?

ABSTRACT

Sticky yellow rectangle traps are used to monitor various pest Rhagoletis flies

(Diptera: Tephritidae), but it is unclear if relative efficacies of these traps differ with

fly species. Here, the main objective was to identify the most efficacious of five

commercial sticky yellow rectangles baited with ammonium carbonate against western

cherry fruit fly, R. indifferens Curran, apple maggot fly, R. pomonella (Walsh), and

walnut husk fly, R. completa Cresson, in Washington State, U.S.A. Two plastic yellow

sticky strips (PL1 and PL2) supplemented with Tanglefoot adhesive and three sticky

yellow cardboards, the Pherocon AM (PA1), Multigard AM (PA2) and Alpha Scents

Yellow Card (PA3), were tested. Across all three species, the PL1 and PL2 +

Tanglefoot generally caught the most flies, the PA3 sometimes caught more than the

PAI, and all caught more than the PA2. Adding Tanglefoot to the PA1 did not make

the trap as efficacious as the PL1 + Tanglefoot against R. indifferens, but it did against

R. pomonella and R. completa. Results suggest the plastic rectangles tested here are

better than standard cardboard rectangles for capturing high numbers of all three

Rhagoletis species, implying they should be the rectangles of choice for monitoring

these flies. Results also suggest that similar trap efficacies against the three species

may have different underlying causes.

Key Words: Rhagoletis indifferens, Rhagoletis pomonella, Rhagoletis completa,

yellow plastic traps, yellow cardboard traps, Tanglefoot® adhesive

INTRODUCTION

Traps are used to monitor and detect various pest Rhagoletis flies (Diptera:

Tephritidae) as a first step in a multi-pronged approach for protecting fruit commodities.

Of all the different trap types developed, sticky yellow rectangles baited with ammonia

compounds, in particular ammonium carbonate, remain the most widely used against

these flies in North America (e.g., Riedl et al. 1989; Liburd et al. 2001; Yee et al. 2012).

These traps are commercially available, flat, light, easy to store and deploy, and the dark

flies are easy to see on and remove from them. Other commercial sticky traps used in

North America or Europe are red or green spheres, the Ladd trap (AliNiazee et al. 1987;

Riedl et al. 1989; Jones and Davis 1989), and the Rebell trap (Remund and Boller 1978).

Non-commercial, experimental sticky traps include yellow spheres (AliNiazee 1981), a

bell trap (Burditt 1988), and cylinder traps (Opp ef a/. 2003).

Efficacies (that is, how well a trap performs in controlled experiments relative to

other traps) between rectangle and some other trap types differ among Rhagoletis species

(e.g., Prokopy and Hauschild 1979; Liburd et al., 2001; Lampe et al. 2005). For example,

against European cherry fruit fly, R. cerasi (L.), three-dimensional yellow Rebell traps

are more efficacious than Pherocon AM traps, whereas the reverse is true for eastern

cherry fruit fly, R. cingulata (Loew) (Katsoyannos et al. 2000; Lampe ef a/. 2005).

1 Corresponding author: Yakima Agricultural Research Laboratory, United States Department of

Agriculture, Agricultural Research Service, 5230 Konnowac Pass Road, Wapato, WA, USA 98951,

(509) 454-6558, wee. yee@ars.usda.goc

2 Washington State University Extension, 1919 NE 78" Street, Vancouver, WA, USA 98665

50 J. ENTOMOL. SOC. BRIT. COLUMBIA 113, DECEMBER 2016

However, which yellow rectangles are most efficacious and whether efficacies of those

rectangles differ among Rhagoletis species have not been well studied.

In particular, sticky yellow plastic rectangles that catch more western cherry fruit fly,

R. indifferens Curran, than conventional sticky yellow cardboard rectangles in

Washington State, U.S.A. (Yee 2014), have not been tested against other Rhagoletis

species. It can be predicted that plastic yellow rectangles are efficacious against them as

well, based on similarities in spectral sensitivities in representative species and fly

responses to color (Prokopy 1968; Agee et al. 1982; Agee 1985). The availability of

different commercial yellow rectangles presents an opportunity to test this hypothesis.

Rejection of this hypothesis could lead to work identifying factors responsible for trap

efficacy and thus more species-specific traps.

In western North America where fruit commodities are of high economic value,

trapping is critical for quarantine and control measures against three Rhagoletis species.

In Washington State and other northwestern U.S. states, as well as in British Columbia,

Canada, R. indifferens and apple maggot fly, R. pomonella (Walsh), are major quarantine

pests of cherries (Prunus spp.) and apple (Malus domestica Borkhausen), respectively. In

California, walnut husk fly, R. completa Cresson, is a major pest of walnuts (Juglans

spp.). Annually, in Washington State, cherries are valued at ~$US300—-$400 million

(National Agricultural Statistics Service 2013a) and apples at ~$US1.5 billion (National

Agricultural Statistics Service 2012); in California, walnuts are valued at oe 3 billion

(National Agricultural Statistics Service 2013b).

In this study, the main objective was to identify the most efficacious Ee five

commercial sticky yellow rectangles baited with ammonium carbonate against R.

indifferens, R. pomonella, and R. completa in Washington State. The hypothesis that

relative efficacies of these traps against all three species are similar was tested. A

secondary objective was to determine which factors could affect their efficacies. In

particular, traps were supplemented with Tanglefoot® adhesive (Contech Enterprises,

Inc., Victoria, B.C., Canada) to determine if this can increase the efficacy of a trap, as

adhesive type effects on fly captures may vary (Yee 2011).

MATERIALS AND METHODS

Five-trap comparisons. Five commercial sticky yellow rectangles were tested in the

first set of experiments (Table 1, Fig. 1). The two plastic traps were the Agri-Sense

Yellow Sticky Strip (PL1) and the Olson Yellow Sticky Strip (PL2) (Agri-Sense-BCS,

South Wales, U.K., and Olson Products, Medina, OH, U.S.A., respectively), both covered

with pressure-sensitive adhesives. These adhesives are thin, solvent-/water-free tacky

materials unlike conventional thick, Vaseline-like adhesives. Both traps were 14 x 23 cm.

The pressure-sensitive adhesives on the traps were supplemented with Tanglefoot®

(Tangle-Trap™ Insect Trap Tropical Formula). Tanglefoot was added, because it is

known from previous tests (Yee 2014; W. L.Y., unpublished) that the pressure sensitive

adhesives on these traps can lose stickiness within 2-3 weeks. More importantly, the

PL1, as received from the manufacturer, had variable amounts of pressure-sensitive

adhesive, and occasional lots were not sticky. About 10 g of Tanglefoot (5 g each side)

was spread onto each plastic trap.

The three cardboard or paper traps were the Pherocon AM (PA1), used for the last 40

years against Rhagoletis flies (e.g., Prokopy and Hauschild 1979; Riedl et al. 1989;

Liburd et al. 2001), the Multigard AM (PA2; it or its variations have been available since

at least 1994 [Katsoyannos et al. 2000]), and the Alpha Scents Yellow Card (PA3;

available within the last six years [Yee 2011]; Table 1, Fig. 1). These three traps were

initially tested without adding Tanglefoot to them, because they were assumed to retain

their tackiness over test trap durations of <4 weeks. The sticky adhesives on the PA] and

PA2 were Vaseline-like, but differed from the Tanglefoot (compositions of commercial

adhesives are proprietary). About 5.0 g and 5.5 g of sticky adhesives were present on the

J. ENTOMOL. Soc. BRIT. COLUMBIA 113, DECEMBER 2016

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52 J. ENTOMOL. Soc. BRIT. COLUMBIA 113, DECEMBER 2016

PA1 and PA2, respectively. The sticky material on PA3 was a hot-melt pressure-sensitive

adhesive that was tackier than the pressure-sensitive adhesive on the PL1 (Yee 2011).

A vial containing 10 g of ammonium carbonate (Keystone Universal, Melvindale, MI,

U.S.A.) with a plastic lid and two 1-mm holes was hung ~1 cm above each trap.

Study sites and experimental types in five-trap comparisons. All sites were

located in central or western Washington State (WA). Sites were unmanaged orchards,

homeowners’ yards, or wild habitats. Twelve tests were conducted between May and

September 2014 in sweet cherry (Prunus avium (L.) L.), apple, black hawthorn

(Crataegus douglasii Lindley), and English walnut trees (Juglans regia (L.); Table 2).

For each fly species, three to five tests were conducted, each using a randomized

complete block design with three to five replicate blocks.

TTT Te ddd ddd ddd ddd dedad,

jot

8° Hemispherical Reflectance Factor

320 360 400 440 480 520 560 600 640 680 720 760 800

Wavelength (nm)

PL1i + Tanglefoot

PL2 + Tanglefoot

PA1

PA2

PAS

Figure 1. Yellow rectangle traps used in study: (A) PL1 + Tanglefoot, (B) PL2 + Tanglefoot,

Elmer Lambda QUV-Vis-NIR Spectrometer Ser. No. 1611; Avian Technologies LLC, Sunapee,

NH).

J. ENTOMOL. SOc. BRIT. COLUMBIA 113, DECEMBER 2016 53

A block was a defined location comprising a set of trees, a single tree, or a sector of a

tree containing all five trap types and within which all trap types were rotated to further

reduce spatial effects. The number of blocks equaled the number of replicate traps.

Availability of trees and layout of trees determined the number of replicates and possible

blocking schemes, as no sets of trees in the field are neatly arranged like trees in

orchards.

One trap per tree was set up in five-tree blocks when at least 25 trees (five trap types

x five replicate trees) were available to use at a site. Here, each trap was spaced 3—5 m

apart, depending on inter-tree distances. This scheme was used for R. indifferens and R.

pomonella (Table 2). When only about five large trees were present, all five trap types

were placed in one tree; each tree was a block. All traps were placed in the south half of a

tree, ~2 m apart. This scheme was used for R. indifferens, R. pomonella, and R. completa

(Table 2). Two or three blocks per tree were set up when there were only two large, 15—

18 m wide walnut trees at a site. Here each block was a 4—5 m sector of a tree with five

traps, each trap 1—2 m apart within the sector (Table 2).

In all tests, traps were hung from branches ~1.5—2 m above ground. Traps within

blocks were rotated 2 to 18 times (Table 2). Flies were removed from traps every time

positions were changed and were saved in cups and later sexed in the laboratory. Traps

were replaced after three weeks if needed, with one to three replacements occurring over

the 3-8 week tests. Particulars of each test site and its trees follow.

Four five-tree block tests were conducted (Table 2). Two tests were conducted for R.

indifferens: one in an unmanaged cherry orchard in Yakima with 145 trees ~4—5 m tall

and wide, and the other in an unmanaged cherry grove in Vancouver with 50 trees ~6—8

m tall and ~3—5 m wide. Two tests were conducted for R. pomonella in an old homestead

in Skamania County with approximately 100 apple and 25 black hawthorn trees ~5—8 m

tall and wide.

Six tests using blocks of one tree were conducted (Table 2). One test for R. indifferens

was conducted in five seedling cherry trees ~5—7 m tall and wide in Roslyn. Three tests

for R. pomonella were conducted in a contiguous stand of black hawthorn trees ~6—8 m

tall and wide in the Nile Valley and in individual black hawthorn and apple trees ~5—7 m

tall and wide in Vancouver. Two tests for R. completa were conducted in five walnut trees

~3—4 m tall and wide in Zillah, Site 1, and in 12 walnut trees ~8—17 m tall and wide in

Naches.

Two tests employing multiple blocks per tree were conducted for R. completa (Table

2) in English walnut in homeowners’ yards. The first test was at Zillah, Site 2, with two

trees, each ~15—18 m tall and wide; the second was at Donald, with two trees, each ~15

m tall and wide.

PL1 + Tanglefoot vs. PA1 and other traps + Tanglefoot. The main purpose of this

second set of experiments was to determine if adding or replacing the adhesive already

present on a trap with Tanglefoot improves the trap’s efficacy. Emphasis was placed on

comparing the PL1 + Tanglefoot with the PA] + Tanglefoot, in most cases with the same

sticky surface areas. However, to gain additional information, other yellow traps +

Tanglefoot (all baited with ammonium carbonate) were also compared in 2014 and 2015.

Blocking schemes and other procedures followed those described for the five-trap

comparison tests.

For R. indifferens, one test was conducted from 18-29 May 2015 in three sweet

cherry trees with one or two blocks per tree for five total blocks in Kennewick, WA. The

PL1 + Tanglefoot vs. PAl + Tanglefoot (each with 596 cm? sticky surface) was the major

comparison. The adhesive on the PA] was scraped off and replaced with Tanglefoot (~5

g each side). To obtain additional information, the traps were compared with three non-

rectangle traps. The first was a yellow PALz trap, a plastic rectangle with ends tied

together (30.5 x 22.9 cm, 614 cm* sticky surface; Plant Protection Institute, Budapest,

Hungary). This trap was covered with a thick Tanglefoot-like adhesive, so no Tanglefoot

was added to it.

J. ENTOMOL. SOC. BRIT. COLUMBIA 113, DECEMBER 2016

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J. ENTOMOL. Soc. BRIT. COLUMBIA 113, DECEMBER 2016 55

The second trap was a yellow ‘Fly Trap’ with a “modular” design (24.8 cm high x 8.9

cm wide, 482 cm? sticky surface; PIC Corporation, Linden, NJ), and the third trap was a

9.0-cm diameter yellow ball (283 cm? sticky surface; laboratory made). The Fly Trap had

a thin layer of pressure-sensitive adhesive, and the ball had no adhesive; Tanglefoot was

added to both. Traps within blocks were rotated four times; due to high fly numbers,

traps were replaced each time.

For R. pomonella (Table 3), three tests were conducted in 2014 and 2015. All tests

used one trap per tree. In Test 1, five blocks of five trees each were set up. In Test 2, five

blocks of two trees each were set up. In Test 3, three blocks of two trees each were set

up.

For R. completa (Table 3), four tests were conducted in 2014 using multiple blocks

per tree. In all four R. completa tests, five blocks were set up. In Tests 1, 3, and 4, there

were two blocks in each of two trees and one block in one. In Test 2, there were two

blocks in one tree and three in a second tree.

For both species, traps within blocks were rotated four to seven times, except at

Skamania in 2015 (twice; Table 3). In 2014, Tanglefoot was added on top of adhesives

already present, but in 2015, the adhesives were scraped off and replaced with

Tanglefoot.

Statistics. For each test, fly counts were summed over all collection dates and square-

root transformed (Zar 1999; data met normality and homogenous variance assumptions)

and then subjected to randomized complete block analysis of variance (ANOVA),

followed by Tukey’s HSD test for means separation (SAS Institute Inc. 2010). In a

second analysis, counts were adjusted for sticky surface area (cm?) before analysis with

ANOVA and Tukey’s HSD test. In addition, orthogonal contrasts were conducted after

ANOVA, using the contrast statement in SAS for fly counts per cm? to identify possible

common factors affecting captures. Specifically, for the five trap-comparison tests, three

contrasts of plastic vs. paper traps or Tanglefoot vs. other adhesives were made: the

means of PL1 + PL2 with Tanglefoot vs. means of PAI + PA2, PA] + PA3, and PA2 +

PA3. For the PL1 + Tanglefoot vs. PAl and other traps + Tanglefoot tests against R.

pomonella and R. completa, two or four contrasts of Tanglefoot vs. other adhesives were

made (none was made for the R. indifferens test). Female- and male-fly data were

combined to simplify the results, as catch patterns of the sexes were similar.

RESULTS

Five-trap comparisons. Within fly species, relative trap efficacy patterns using five-

tree block, one-tree block, and multiple blocks per tree designs were similar, especially

for the best performing traps (Figs. 2-4), so the way blocking was performed made no

difference in the conclusions. Across all three Rhagoletis species, the PL1 and PL2 +

Tanglefoot were generally the most efficacious of the five traps tested. Compared with

the PL2 + Tanglefoot, the PL1 + Tanglefoot caught statistically more R. indifferens in one

(Fig. 3A) of three tests, more R. pomonella in one (Fig. 3D) of five tests, and more R.

completa in one (Fig. 4A) of four tests.

The PA3 was the next most efficacious trap, but the PL1 + Tanglefoot caught

statistically more flies than the PA3 in nine of twelve tests across species. The PA3

caught statistically more R. pomonella than the PA1 in one (Fig. 3D) of five tests and

more R. completa than the PA1 in three (Figs. 3E, 3F, and 4B) of four tests. In no case

did the PAI catch statistically more flies than the PA3. The PA2 was the least effective of

all five traps.

Combining data from all tests, more females than male flies were caught on all trap

types. For R. indifferens, 58-60% caught on the five trap types were females. For R.

pomonella, 65-69% of flies caught were females; for R. completa, 55-66% were

females.

J. ENTOMOL. SOC. BRIT. COLUMBIA 113, DECEMBER 2016

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Five-Tree Blocks

28 B. R. indifferens, Vancouver, 12 June-

14 July (206 flies)

A. R. indifferens, Yakima, 19 May

-25 June (133 flies)

A 20

st 5

0 0

300 250

D. R. pomonella, Skamania, 11 Aug-

2 Sept (2,176 flies)

C.R. pomonella, Skamania, 4 July-

7 Aug (3,063 flies)

200

150

Mean No. Flies per Replicate + SE

150

100

PL1 PL2 PA1 PA2 PA3 PL1 PL2 PA1 P PA3

+ Tanglefoot + Tanglefoot

Figure 2. Five-tree block tests: mean numbers of flies (sexes combined) caught per replicate

+ SE in 2014: (A) R. indifferens in Yakima; (B) R. indifferens in Vancouver; (C) R. pomonella

in Skamania in apple and hawthorn; (D) R. pomonella in Skamania in apple. (A) F' = 5.96; df

= 4, 16; P = 0.0039; (B) F = 17.70; df = 4, 16; P < 0.0001; (C) F = 16.77; df = 4, 16; P<

0.0001; (D) F = 19.34; df = 4, 16; P < 0.0001. Means within tests with same letters are not

significantly different (Tukey’s HSD test, P > 0.05).

PL1 + Tanglefoot vs. PAl and other traps + Tanglefoot. The PL1 + Tanglefoot

caught statistically more R. indifferens than the PA1 + Tanglefoot when both had 596 cm?

sticky surfaces (Fig. 5). It also caught more flies than two non-rectangle traps with

Tanglefoot, although statistically not more than the PALz (Fig. 5), which had a different

adhesive. However, unlike for R. indifferens, mean catches of R. pomonella on the PL1 +

Tanglefoot and PA1 + Tanglefoot with 596 cm sticky surfaces did not differ statistically

in three tests (Figs. 6A-6C). Similarly to those of R. pomonella, mean catches of R.

completa on the PL1 + Tanglefoot with a 596 cm? sticky surface and PA] + Tanglefoot

with 407 and 596 cm? sticky surfaces did not differ (Figs. 7A and 7B). However, the PL1

+ Tanglefoot caught more R. pomonella than the PA2 + Tanglefoot (Fig. 6A) and more R.

completa than both the PA2 + Tanglefoot (Fig. 7C) and PA3 + Tanglefoot (Fig. 7D) when

sticky surface areas were equal.

Fly captures adjusted for sticky surface area. In the first set of the five-trap

comparisons, the relative efficacies of traps based on catch numbers not adjusted and

adjusted for sticky surface area differed, but the major patterns were the same (Table 4).

58 J. ENTOMOL. Soc. BRIT. COLUMBIA 113, DECEMBER 2016

Notably, for R. indifferens, the PL1 + Tanglefoot was still more efficacious than the PA2

and PA3 in two of three tests; for R. pomonella, the PL1 + Tanglefoot was more

efficacious than the PA] in three of five tests, and the PL] and PL2 + Tanglefoot were

more efficacious than the PA2 in all five tests; the PL1 + Tanglefoot was more so than the

PA3 in four of five tests. For R. completa, the PL1 + Tanglefoot was more efficacious

than the PAI and PA2 in all four tests—more so than the PA3 in two of four tests (Table

4),

Blocks of One Tree Each

eee A. R. indifferens, Roslyn, 9 July-18 Aug a B. R. pomonella, Nile Hawthorn, 9 July-

(16,035 flies) 3 Sept (724 flies)

joi A

1200

900

600

300

C. R. pomonella, Vancouver Hawthorn,

23 June-16 July (580 flies)

A 200

160

D. R. pomonella, Vancouver Apple,

3-23 July (1,529 flies)

120

Mean No. Flies per Replicate + SE

300

E.R. completa, Zillah 1, 2 July-2 Sept F.R. completa, Naches, 16 Jul-29 Aug

(6,572 flies) (3,116 flies)

600 250

A PL ok,

500 A

200

400

150

300

100

200

100 30 |

O ERSTE EES, PERERA Redeseaeeee ROR E een '

Pits PE2 PA. PA2* PAS Ald Pho PAL: PA2.“SPAS

+ Tanglefoot + Tanglefoot

Figure 3. Blocks of one-tree tests: mean numbers of flies (sexes combined) caught per

replicate + SE in 2014: (A) R. indifferens in Roslyn; (B) R. pomonella in Nile; (C) R.

pomonella in Vancouver in hawthorn; (D) R. pomonella in Vancouver in apple; (E) R.

completa in Zillah, Site 1; (F) R. completa in Naches. (A) F = 116.27; df = 4, 16; P<

0.0001; (B) F = 33.51; df= 4, 16; P < 0.0001; (C) F = 1,631.26; df = 4, 8; P < 0.0001; (D) F

= 47.77; df = 4, 16; P < 0.0001; (E) F = 90.56; df = 4, 16; P < 0.0001; (F) F = 90.56; df = 4,

16; P < 0.0001. Means within tests with same letters are not significantly different (Tukey’s

HSD test, P > 0.05).

J. ENTOMOL. SOC. BRIT. COLUMBIA 113, DECEMBER 2016 59

Orthogonal contrasts. For the first set of the five-trap comparisons, contrasts

between the means of PL1 + PL2 with Tanglefoot vs. means of PA1l + PA2, PAI +

PA3, and PA2 + PA3 differed regardless of fly species (Table 5). This suggests

plastic material or Tanglefoot contributed to higher fly catches. In the PL1 +

Tanglefoot and other traps + Tanglefoot comparisons for R. pomonella, results

suggested Tanglefoot increased captures on PL1, PA2, and PA3, but not on PAI

(Table 5); for R. completa, results suggest Tanglefoot increased captures on PL1 and

PA1 (Table 5).

Multiple Blocks Per Tree

400

A. R. completa, Zillah 2, 2 July-1 Aug (2,750 flies)

300

200

100

400

300

Mean No. R. completa per Replicate + SE

ee |

PL1 Pi2 . PAL PAZ

+ Tanglefoot

Figure 4. Multiple blocks per tree tests: mean numbers of flies (sexes combined) caught per

replicate + SE in 2014: (A) R. completa in Zillah, Site 2; (B) R. completa in Donald. (A) F =

88.45; df= 4, 12; P < 0.0001; (B) F = 80.80; df = 4, 12; P < 0.0001. Means within tests with

the same letters are not significantly different (Tukey’s HSD test, P > 0.05).

J. ENTOMOL. SOC. BRIT. COLUMBIA 113, DECEMBER 2016

Multiple Blocks Per Cherry Tree

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2100

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Shue?

PL1 + Tanglefoot vs. PAI + Tanglefoot and other traps in Kennewick in 2015. Sticky surface

Figure 5. Mean captures of Rhagoletis indifferens (sexes combined) + SE per replicate on

areas are shown below trap types. F

J. ENTOMOL. SOc. BRIT. COLUMBIA 113, DECEMBER 2016 61

Two- or Five-Tree Blocks of Apple

200

A. Test 1: R. pomonella, Skamania, 4 Sept-16 Oct

2014 (1,975 flies)

160

120

rit PA1 PA2 PA3

+ Tanglefoot

150 596 cm* 596 cm? 596 cm? 596 cm” 478 cm?

B. Test 2: R. pomonella, Skamania, 3-29 Sept

2015 (631 flies)

C. Test 3: R. pomonella, Roslyn, 17 Aug-24 Sept

2015 (135 flies)

Mean No. R. pomonella per Replicate + SE

PL1 + Tanglefoot PA1 + Tanglefoot

596 cm* 596 cm@

Figure 6. Mean captures of Rhagoletis pomonella (sexes combined) + SE per replicate in

2014 and 2015 on (A) PLI and PL1 + Tanglefoot vs. PAl + Tanglefoot, PA2 + Tanglefoot,

and PA3 + Tanglefoot; PL1 + Tanglefoot vs. PAl + Tanglefoot in (B) Skamania and in (C)

Roslyn. Sticky surface areas are shown below trap types. (A) F = 7.10; df= 4, 16; P = 0.0017;

(B) F = 1.66; df = 1, 4; P = 0.2669; (C) F = 8.13; df = 1, 2; P = 0.1041. Means within tests

with same letters are not significantly different (Tukey’s HSD test, P > 0.05).

62 J. ENTOMOL. SOC. BRIT. COLUMBIA 113, DECEMBER 2016

Multiple Blocks Per Walnut Tree |

100 100

A. Test 1: R. completa, Naches, 27 B. Test 2: R. completa, Donald, 27

Aug-26 Sept (772 flies) Aug-26 Sept (838 flies)

80 80

vy 60 60

40 40

20 20

0 0

PL1 PL1+ PAY‘ PA1 + PL1 PL1 PA1 PA1

Tanglefoot Tanglefoot + Tanglefoot

596cm* §96cm* 407cm* 407 cm” 596cm* 596cm2 407cm2 596 cm”

40 40

D. Test 4: R. completa, Naches, 29

Aug-26 Sept (257 flies)

C. Test 3: R. completa, Naches, 29

Aug-26 Sept (201 flies)

Mean No. R. completa per Replicate +

PL PL1 PA2 PL1 PUTO <7 PA

+ Tanglefoot + Tanglefoot

All 596 cm? All 596 cm?

Figure 7. Mean captures of Rhagoletis completa (sexes combined) + SE per replicate in 2014

on PL1 and PL1 + Tanglefoot vs. (A, B) PAl and PAI + Tanglefoot; (C) PA2 + Tanglefoot;

and (D) PA3 + Tanglefoot. Sticky surface areas are shown below trap types. (A) F' = 12.35; df

= 3, 12; P = 0.0006; (B) F = 8.64; df = 3, 12; P = 0.0025; (C) F = 19.34; df = 2, 8; P=

0.0009; (D) F = 7.63; df = 2, 8; P = 0.0140. Means within tests with same letters are not

significantly different (Tukey’s HSD test, P > 0.05).

J. ENTOMOL. SOC. BRIT. COLUMBIA 113, DECEMBER 2016

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J. ENTOMOL. SOC. BRIT. COLUMBIA 113, DECEMBER 2016 67

DISCUSSION

The plastic rectangles supplemented with Tanglefoot were the most efficacious of the

five sticky yellow rectangle traps tested against R. indifferens, R. pomonella, and R.

completa. Until recently (Yee 2011, 2014), the PAI (Pherocon AM) could be assumed to

be the most efficacious yellow rectangle against most Rhagoletis flies in North America.

However, the PA] had usually been compared with spheres and Ladd traps and not with

other yellow rectangles against R. pomonella and R. completa, as well as the blueberry

maggot, R. mendax Curran (e.g., Prokopy and Hauschild 1979; AliNiazee et al. 1987;

Riedl et al. 1989; Liburd et al. 2001; Teixeira and Polavarapu 2001). In the current study,

the PA1 was only more efficacious than the PA2, as it was against R. cerasi (Katsoyannos

et al. 2000). It is unclear whether traits of the PAl have changed over the years. For this

reason, the trap traits documented in Fig. 1 and Table 1 can be useful for future

comparisons of traps tested here with other, newer traps.

Based on their superior performance against high Rhagoletis populations, the plastic

traps + Tanglefoot may be able to detect lower fly populations than all the cardboard

traps tested here, perhaps making them better options for monitoring. Plastic and

cardboard traps cost about the same (~U.S.$1.10 per trap; cost of 10 g TF per trap is ~9

cents), but have the advantage of not fading over the season, as can the PA2 and PA3 (W.

L. Y., personal observations). There are, however, several disadvantages to the plastic

traps. One is that they are thinner and lighter, so are more prone to flap in heavy winds.

This sometimes causes them to tear loose from branches; however, this can be prevented

by securing the traps to branches using three ties instead of one. Another disadvantage is

that, as currently manufactured, plastic traps would need to be supplemented with

Tanglefoot. Thus, caution should be taken when deciding which traps to use, because

overall trap catch is not always necessarily the deciding factor in selecting an ‘optimal’

trap for monitoring purposes. Traps need only be effective enough to provide a

consistent, reliable ‘sample’ or estimate of a population, with minimal cost and time in

servicing.

All traps generally caught more females than males, consistent with findings for R.

mendax (e.g., Liburd et al. 2001; Teixeira and Polavarapu 2001). Other studies showed

that 46%, 57%, and 50% of R. indifferens, R. pomonella, and R. completa that emerged

from soil under cages, respectively, were females (Frick et al. 1954; Dean and Chapman

1973; Boyce 1934). Because females comprised 55-69% of trap catches here, the traps

may be slightly biased towards females, suggesting males may be less attracted to yellow

rectangles than females.

The major objective here was to identify the most efficacious of five commercial

sticky yellow rectangles against flies, but a secondary objective was to determine which

factors might affect their efficacies. Differences in sticky surface areas, color,

translucence, and adhesive type make identifying factors responsible for the greater

efficacies of the PL] and PL2 + Tanglefoot vs. the PA1, PA2, and PA3 (Figs. 2-4)

difficult, but there are at least three possible factors. One is that the sticky surface areas

of the plastic traps were larger; however, analyses of catches per sticky area suggest this

was of minimal importance for all three species. A second possible factor is that all three

fly species were most stimulated visually by color and other cues in the plastic traps. The

third possible factor is that supplementing the plastic traps with Tanglefoot increased

their efficacy either by making them tackier due to composition or amount or by altering

their visual properties.

Results comparing the PL1 and PAI both with Tanglefoot and with equal sticky

surface areas against R. indifferens in 2015 suggest that Tanglefoot did not cause the

greater efficacy of the PL1 + Tanglefoot vs. the PA1 against R. indifferens in 2014 tests.

More likely, the yellow color and translucence of the PL1 caused this (Yee 2014). The

PL1 + Tanglefoot was also better than two non-rectangle traps with Tanglefoot for

catching high numbers of R. indifferens, further suggesting traits of the PL] itself

68 J. ENTOMOL. Soc. BRIT. COLUMBIA 113, DECEMBER 2016

independent of Tanglefoot were responsible for its high efficacy. Perhaps not

coincidentally, the PaLz trap was plastic and also performed well.

In contrast to results for R. indifferens, Tanglefoot on the PL1 appeared responsible

for the higher efficacy of the PL1 + Tanglefoot vs. the PAI against R. pomonella and R.

completa, based on tests where sticky surface areas were equal. Tanglefoot may be

tackier and/or had a lower viscosity than the adhesive on the PAI, making flies stick

faster. Less likely, it increased the visual attractiveness of the trap. Polybutene is the

active ingredient in Tanglefoot (Contech 2014) and presumably also in the adhesive on

the PAI (exact chemical compositions of the adhesives are unpublished). Even though

both adhesives are clear or slightly cloudy, particular polymers, grade, or amount of

polybutene in Tanglefoot and other adhesives probably differ.

Supplementing the PA2 and PA3 with Tanglefoot resulted in variable outcomes for R.

pomonella and R. completa. For both flies, Tanglefoot was not a factor in why the PL1 +

Tanglefoot performed better than the PA2. The distinct reflectance/color of the PA2 (Fig.

1F) may simply have been less attractive. In contrast, for R. pomonella, the PL1 +

Tanglefoot appeared better than the PA3 solely because of the Tanglefoot. Traits of the

PA3, PL1, and PAI thus may be similarly attractive to R. pomonella. However, for R.

completa, the PL1 + Tanglefoot caught more flies than the PA3 + Tanglefoot, so traits of

the PL1 may have been more attractive for this species, although more tests are needed to

confirm this. 7

In summary, results suggest the plastic rectangles + Tanglefoot tested here are better

than standard cardboard rectangles for capturing high numbers of all three Rhagoletis

species. This implies these should be the rectangles of choice for monitoring these

species. The efficacy of some cardboard rectangles tested against R. pomonella and R.

completa but not against R. indifferens may be increased simply by using an alternative

or more adhesive. This suggests similar trap efficacies against the three species may have

different underlying causes, which if true has implications for the development of more

species-specific and efficacious traps.

ACKNOWLEDGEMENTS

We thank Pete Chapman, Janine Jewett, and Nicholas Ward (USDA-ARS) for

laboratory and field assistance, the USDA Forest Service for use of the Saint Cloud

Recreational Area in Skamania County, the City of Vancouver Parks and Recreation

Greenway’s Sensitive Wetlands, Clark County Heritage Farm, and various homeowners

for allowing us to use their sites for tests. We also thank Grant McQuate (Daniel K.

Inouye U.S. Pacific Basin Agricultural Research Center, USDA-ARS, Hilo, HI), Jana

Lee (Horticulture Crops Research Unit, USDA-ARS, Corvallis, OR), and two

anonymous reviewers for comments on the manuscript. This article reports results of

research only. Mention of a proprietary product does not constitute an endorsement or a

recommendation for its use by USDA.

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- -12- | Print: versions: 2015, «¥2(¢1): doi:

10.1673/031.012.12401.

Zar, J. H. 1999. Biostatistical analysis. Fourth edition. Prentice Hall, Upper Saddle River, NJ.

J. ENTOMOL. SOC. BRIT. COLUMBIA 113, DECEMBER 2016 71

SCIENTIFIC NOTE

Pacific Flatheaded Borer, Chrysobothris mali Horn

(Coleoptera: Buprestidae), found attacking apple saplings

in the Southern Interior of British Columbia

SUSANNA ACHEAMPONG |, GABRIELLA M.G. ZILAHI-

BALOGH’, ROBERT G. FOOTTIT®?, GARY J.R. JUDD‘, AND T.

DIMARIA®

Key words: Chrysobothris mali, Pacific flatheaded borer, Malus domestica

The Pacific flatheaded borer, Chrysobothris mali Horn (Coleoptera: Buprestidae), is

widely distributed throughout western North America, occurring west of the Rocky

Mountains from California to British Columbia and western provinces (Furness and

Carolin 1977; Soloman 1995). It feeds on 41 genera of plants in 20 families, including

Malus (Burke 1929). We investigated borer damage to young apple, Malus domestica,

saplings reported by a grower in July 2015 in Kelowna, British Columbia. The saplings

were planted in April 2014 and 2015. Larvae were actively feeding on B9 and M9

rootstocks imported from the Netherlands and grafted with gala and honeycrisp apple

varieties. The report was a concern, because flatheaded borer is not a pest of apples in

British Columbia. Apple clearwing moth, a serious pest of apples in British Columbia

was imported on rootstock from the Netherlands. The study was conducted to identify the

Chrysobothris species, its distribution and infestation levels to help guide management

recommendations.

Infested trees had dark-coloured, cracked bark with frass showing through. Frass was

evident around the base of some infested trees. Larvae were found beneath the bark and

inside the wood, and larval galleries were exposed when bark was removed; one larva

was found in a sapling, rarely two in a sapling. The galleries occasionally encircled the

stem, which killed the sapling. Leaf symptoms on infested saplings varied from yellow to

purple.

Surveys were conducted in 19 young apple plantings and nurseries in the Okanagan

and Similkameen valleys of British Columbia, from July to October 2015, to determine

distribution and infestation levels. All saplings at each survey site were visually inspected

for damage and presence of larvae (Table 1). Survey sites were located in Kelowna,

Oyama, Winfield, Lavington, Cawston, and Keremeos. Site selections were based on

information about young apple plantings provided by B.C. Tree Fruits Ltd. We inspected

infested saplings at the original site, but it was not included in our surveys, because the

grower had removed some of the infested trees.

To confirm whether the borer was a native or imported Chrysobothris species, 20

infested saplings, including six from the original site, were sent to the Canadian Food

Inspection Agency (CFIA) Entomology laboratory in Ottawa for larval and adult

identifications. Infested saplings were also kept in the B.C. Ministry of Agriculture

' Corresponding Author: BC Ministry of Agriculture, 1690 Powick Rd., Kelowna, BC V1X 7G5; (250)

861-7681, Susanna.Acheampong@gov.bc.ca

2 Canadian Food Inspection Agency, 1853 Bredin Rd., Kelowna, BC V1Y 7S9

3 Agriculture and Agri-Food Canada, Ottawa Research and Development Centre, K.W. Neatby Building,

960 Carling Ave, Ottawa, ON K1A 0C6

4 Agriculture and Agri-Food Canada, Summerland Research and Development Centre, Summerland, BC

VOH 1Z0

5 BC tree Fruits, 880 Vaughan Ave., Kelowna, BC V1Y 7E4

72 J. ENTOMOL. SOC. BRIT. COLUMBIA 113, DECEMBER 2016

Laboratory in Kelowna for rearing out adults. Four larvae were sent to Robert Foottit

(AAFC) for DNA barcoding.

Two of four larval specimens were barcode sequenced, producing identical

sequences, and matched to C. mali. Species identifications were based on sequencing of

the DNA barcode region of the mitochondrial gene Cytochrome C oxidase subunit I

(COI). DNA was extracted, amplified, and sequenced according to standard DNA

barcode protocols (http://www.barcodeoflife.org/sites/default/files/

Protocols for High Volume DNA Barcode _Analysis.pdf), then compared to reference

sequences in BOLD (Barcode of Life Data Systems (http://www.boldsystems.org), based

on specimens in the Canadian National Collection, collected in Alberta, Saskatchewan

and California. The match to C. mali ranged from 99.5 to 98.8 percent, whereas the

similarity to 22 other species ranged from 82.0 to 95.0 percent. Sequences were

deposited in GenBank, National Center for Biotechnology Information, U.S. National

Library of Medicine, Bethesda, MD, USA (http:/Avww.ncbi.nlm.nih.gov/genbank/;

GenBank Accession No. KX283168).

The CFIA and the B.C. Ministry of Agriculture laboratories reared out adult C. maii.

Chrysobothris mali larvae were detected at low infestation levels (0.01 to 0.22%) in 15 of

the 19 survey sites (Table 1).

Table 1

Flatheaded borer larval infestation levels in young apple orchards/nurseries in the

Okanagan and Similkameen valleys of British Columbia in 2015.

Number of Number of Infestation

Orchard/Nursery trees examined infested trees Level (%)

Pe A 42,000 10 0.02 9

B 14,000 1 0.01

C 14,000 1 0.01

D 90,000 45 0.05

E 15,000 33 0.22

F 10,000 0 0

G 17,500 21 0.12

H 35,000 11 0.03

| 55,000 10 0.02

J 4,000 0 0

K 150,000 147 0.10

L 40,000 0 0

M 15,000 4. 0.02

N 245,000 1 0

J. ENTOMOL. SOC. BRIT. COLUMBIA 113, DECEMBER 2016 ois:

There were previous unconfirmed reports of C. mali damage to young apple trees in

Kelowna in 2003 (Philip 2003). Our study provides the first confirmed record of C. mali

causing damage on young fruit trees in the Okanagan. Chrysobothris mali has previously

been reported as a pest of newly planted fruit trees and of young nursery trees in

California (Burke 1919; Burke 1929; McNelly et al. 1969). McNelly et al. (1969)

reported that young trees stressed by sunburn, drought, or bark injury, or planted too late

in the spring were particularly subject to attack.

Chrysobothris mali produces a single generation per year. It overwinters as mature

larvae in the heartwood. Pupation occurs between April and May. Adult emergence and

oviposition occurs in June and July (Burke 1929). Eggs are laid in cracks and crevices in

the bark. Eggs hatch, and larvae mine into the cambium and pack frass in the mine

behind it (Davis et al. 1968).

ACKNOWLEDGEMENTS

We thank Caitlin Miller, Diane Thomas, Hugh Philip, Carl Withler, and B.C. Tree

Fruits Ltd. staff for help with surveys, and participating apple nurseries, rootstock

importers and growers for access to survey sites. We thank Amanda Biernacka-Larocque

(AAFC, Ottawa) for carrying out the barcode sequencing, and Eduard Jendek (CFIA,

Ottawa) for larvae and adult identifications. Funding for the survey was provided by the

B.C. Fruit Growers’ Association.

REFERENCES

Burke, H. E. 1919. Biological Notes on the Flatheaded Apple Tree Borer (Chrysobothris femorata Fab.)

and the Pacific Flatheaded Apple Tree Borer (Chrysobothris mali Horn). Journal of Economic

Entomology 12:26—330.

Burke, H. E. 1929. The Pacific Flathead Borer. USDA Technical Bulletin 83. 36 pp.

Davis, C. S., J. H. Black, K. W. Hench, and C. V. Carlson. 1968. Controlling Pacific Flatheaded Borer.

California Agriculture 22:6—7.

Furness, R. L., and V. M. Carolin. 1977. Western Forest Insects. USDA Forest Service. Miscellaneous

Publication No. 1339.

McNelly, L. B., D. H. Chaney, G. R. Post, and C. S. Davis. 1969. Protecting Young trees from Attack by

the Pacific Flatheaded Borer. California Agriculture 23:12—13.

Philip, H (compiler). 2003. British Columbia Ministry of Agriculture, Food & Fisheries 2003 Insect Pest

Report to the Western Committee on Crop Pests. Pages 6-8 in Appendix I, Minutes of the Western

Committee on Crop Pests 43rd Annual Meeting (prepared by Ian Wise). 98 pp. lhittp://

vww.westernforum.org/Documents/WCCP/WCCP%20Minu VCCP2003Minutes.pdf [accessed 4

May 2016].

Soloman, J. D. 1995. Guide to Insect Borers in North American Broadleaf Trees and Shrubs. USDA

Forest Service Handbook. AH-706. 735 pp. http://www.srs.fs.usda.gov/pubs/misc/ah_.706/

ah-706.htm [accessed 29 April 2016].

74 J. ENTOMOL. SOc. BRIT. COLUMBIA 113, DECEMBER 2016

NATURAL HISTORY AND OBSERVATIONS

First Canadian records for two invasive seed-feeding bugs,

Arocatus melanocephalus (Fabricius, 1798) and Raglius

alboacuminatus (Goeze, 1778), and a range extension for a

third species, Rhyparochromus vulgaris (Schilling, 1829)

(Hemiptera: Heteroptera)

SUSANNA ACHEAMPONG]!, WARD B. STRONG?, MICHAEL D.

SCHWARTZ?3, ROBERT J. HIGGINS‘, MOLLY A.THURSTONS®,

EMMA J. WALKER®, AND J. ROBERTS’

New invasive insect species affect agriculture, the environment, landscapes, and

homeowners. Invasive species are difficult and challenging to manage due to limited

availability of direct control products or other management strategies. Nuisance insect

outbreaks can have negative impacts on homeowners and businesses; if they reach high

densities, they can cause anxiety and discomfort and additional management costs. We

report here the first records from Canada for two new invasive pests, Arocatus

melanocephalus and Raglius alboacuminatus, and provide a range extension for

Rhyparochromus vulgaris (Schilling), reported as new to British Columbia by Scudder

(in press, this volume).

The elm seed bug, Arocatus melanocephalus Fabricius (Hemiptera: Heteroptera:

Lygaeidae), is native to Europe and widely distributed in central and southern Europe

(Ferracini and Alma 2008); it was reported in China in 2013 (Gao et al. 2013). Arocatus

melanocephalus was first detected in the United States, in Idaho, in 2009 and is present

in Oregon, Washington, and Utah (Collman and Bush 2016). An infestation of

A. melanocephalus was reported by a homeowner in the Rutland area of Kelowna,

British Columbia, in June 2016. There were large numbers of adults in and around the

home and on Chinese elm wood piles in the yard. Three homeowners in the area also

reported A. melanocephalus outbreaks.

Adults are 6.5~—7 mm long, strongly punctate, with conspicuously contrasting dark-red

and black coloration—black on head, posterior lobe of pronotum, scutellum, and

posterior half of corium, and rusty red on anterior lobe of pronotum, outer portion of

clavus, and anterior portion of corium (Gao et al. 2013; figures la and 1b).

The life cycle of A. melanocephalus in Canada has not been determined. In Europe

and the United States, there is one generation per year (Maistrello et al. 2006; Idaho State

Department of Agriculture 2013). They overwinter as adults and emerge in the spring to

lay eggs on elm. Nymphs feed on elm seeds from May to June and develop into adults in

the summer (Idaho State Department of Agriculture 2013). Arocatus melanocephalus

1 Corresponding Author: B.C. Ministry of Agriculture, 1690 Powick Rd., Kelowna, B.C. V1X 7G5;

(250) 861-7681, susanna.acheampong@gov.bc.ca

2 B.C. Ministry of Forests, Lands, and Natural Resource Operations, Kalamalka Forestry Centre, 3401

Reservoir Rd, Vernon, B.C. V1B 2C7

3 Agriculture and Agri-Food Canada, Ottawa Research and Development Centre, K.W. Neatby Building,

960 Carling Ave, Ottawa, ON K1A 0C6

4 Biological Sciences, Thompson Rivers University, 900 McGill Road, Kamloops, B.C. V2C 0C8

° BC tTee Fruits, 880 Vaughan Ave., Kelowna, B.C. V1Y 7E4

6 Department of Ecology and Evolutionary Biology, University of Toronto, 25 Wilcox Street, Toronto,

ON, MS5S 3B2

7 Canadian Food Inspection Agency, 506 West Burnside Road, Victoria, B.C., V8Z 1M5

J. ENTOMOL. SOC. BRIT. COLUMBIA 113, DECEMBER 2016 BB

feed primarily on elm seeds (U/mus spp.), but have been collected from oak (Quercus)

and linden (Tilia) (Bechinski et al. 2012).

Figure 1a. Dorsal view of adult Arocatus melanocephalus. Photograph by W. B. Strong

Figure 1b. Lateral view of adult Arocatus melanocephalus. Photograph by W. B. Strong

An outbreak of Raglius alboacuminatus (Hemiptera: Heteroptera:

Rhyparochromidae) was reported by an orchardist in Kelowna, British Columbia, in

August 2016. The source of the infestation appeared to be a cut hayfield next to the

76 J. ENTOMOL. SOC. BRIT. COLUMBIA 113, DECEMBER 2016

property. There were large numbers of R. alboacuminatus at the grower’s fruit shop,

apple bins, and in and around the home and other buildings and structures on the

property.

Raglius alboacuminatus is native to Europe and the Mediterranean. It was first

detected in North America, in Utah, in 1999, and it is present in California, Oregon, and

Washington (Henry 2004; Bechinski and Merickel 2007). Adults are about 5-6 mm long,

with dark-brown to black coloration, white markings on the posterior pronotal lobe,

anterior half of the corium, and three conspicuous white spots—one on the apex of each

corium, and one on the apex of hemelytral membrane (figures 2a and 2b).

Figure 2b. Lateral view of Raglius alboacuminatus. Photograph by W. B. Strong

J. ENTOMOL. SOc. BRIT. COLUMBIA 113, DECEMBER 2016 fe:

The life cycle of R. alboacuminatus in Canada has not been determined. There is one

generation per year in the United States (Bechinski and Merickel 2007), but there are two

in England and three in Russia (Henry 2004; Southwood and Leston 1959). Raglius

alboacuminatus overwinters as adult. Overwintered adults lay eggs in the soil or ground

litter in early spring. Nymphs feed on developing seeds of plants in the mint family

(Lamiaceae). The preferred host in the U.K. is black horehound, Ballota nigra (Bantock

and Botting 2013).

In addition to the two species new to British Columbia, we also report the occurrence

of a third species of the exotic seed bug, Rhyparochromus vulgaris (Schilling)

(Hemiptera: Heteroptera: Rhyparochromidae), from two locations in Kelowna. It was

collected in conjunction with A. melanocephalus in September 2016 and as by-catch in

surveillance traps for invasive alien species from the Kelowna landfill (49.948597° N,

119.424989° W) in June 2016. The initial discovery of R. vulgaris in British Columbia is

reported by Scudder (in press). All three species are the first representatives of these

genera in Canada.

Although A. melanocephalus and R. alboacuminatus are not known as agricultural

pests, the presence of large numbers could be problematic for homeowners and farmers.

Elm is a common landscape tree in Interior British Columbia, which may lead to more

reports of infestations by A. melanocephalus. Years with high summer temperatures may

see particularly large numbers, as was found in Italy (Maistrello et al. 2006). The large

numbers of R. alboacuminatus were a nuisance for the affected orchardist, who was

worried about the negative impact on his business. The spread of R. alboacuminatus via

farm equipment, vehicles, and transportation of farm produce is likely; we had to remove

R. alboacuminatus hitchhikers from our vehicle before leaving the infested property.

Vouchers of the three species have been deposited at the Canadian National Insect

Collection, and Spencer and Royal BC museums. A R. vulgaris voucher from the

invasive alien species landfill traps has been deposited in the Canadian Forest Service,

Pacific Forestry Centre reference collection, Victoria, British Columbia. Photographs are

of specimens collected at the sites.

ACKNOWLEDGEMENT

We thank two anonymous reviewers for their constructive comments, edits, and

timely response.

REFERENCES

Bantock, T., and J. Botting. 2013. British bugs: An online identification guide to UK Hemiptera. http://

www.britishbugs.org.uk/index.html [Accessed 3 November 2016].

Bechinski, E. J., J. Barbour, and F. Merickel. 2012. Elm seed bug, Arocatus melanocephalus, a new bug

in Idaho. http://vickisgardentips.com/wp-content/uploads/2013/01/Elm-seed-bug-handout.pdf

[Accessed 2 November 2016].

Bechinski, E. J., and F. Merickel. 2007. Tuxedo bug. A new home-invading insect in Idaho. University of

Idaho, College of Agricultural and Life Sciences, Extension publication CIS 1147. http://

www.cals.uidaho.edu/edcomm/pdf/cis/cis1147.pdf [Accessed 3 November 2016].

Collman, S. J., and M. R. Bush. 2016. Emerging Pests in Pacific Northwest Ornamentals https://

pnwhandbooks.org/insect/emerging-pacific-northwest-ornamentals [Accessed 2 November 2016].

Ferracini, C., and A. Alma. 2008. Arocatus melanocephalus a hemipteran pest on elm in the urban

environment. Bulletin of Insectology, 61(1):193—194.

Gao, C., E. Kondorosy, and W. Bu. 2013. A review of the genus Arocatus from Palaearctic and Oriental

regions (Hemiptera: Heteroptera: Lygaeidae). Raffles Bulletin of Zoology 61(2):687—704.

Henry T. J. 2004. Raglius alboacuminatus (Goeze) and Rhyparochromus vulgaris (Schilling)

(Lygaeoidea: Rhyparochromidae): Two palearctic bugs newly discovered in North America.

Proceedings of the Entomological Society of Washinton 106:513—522

78 J. ENTOMOL. Soc. BRIT. COLUMBIA 113, DECEMBER 2016

Idaho State Department of Agriculture. 2013. Elm seed bug, Arocatus melanocephalus: an exotic

invasive pest new to the USS.

spring 2013 esb fact _sheet.pdf [Accessed 2 November 2016].

Maistrello, L., L. Lombroso, E. Pedroni, A. Reggiani, and S. Vanin. 2006. Summer raids of Arocatus

melanocephalus (Heteroptera, Lygaeidae) in urban buildings in Northern Italy: Is climate change to

blame? Journal of Thermal Biology, 31(8):594—-598.

Scudder, G. G. E. in press. Rhyparochromus vulgaris (Schilling) (Hemiptera: Heteroptera:

Rhyparochromidae): newly discovered in the interior of British Columbia. Journal of the

Entomological Society of British Columbia

Southwood, T. R. E., and D. Leston. 1959. Land and Water Bugs of the British Isles. Frederick Warner &

Co. London. 436 pp.

J. ENTOMOL. SOc. BRIT. COLUMBIA 113, DECEMBER 2016 79

NATURAL HISTORY AND OBSERVATIONS

Notes on insects recently introduced to Metro Vancouver

and other newly recorded species from British Columbia

C. G. RATZLAFF'!, K.M. NEEDHAM, and G. G. E. SCUDDER

ABSTRACT

Sixteen insects species are recorded for the first time from British Columbia, including

seven new to Canada. These records are comprised of nine introduced species,

including the first record of Rhopalum gracile Wesmael (Hymenoptera: Crabronidae)

from North America, and range extensions for seven species native to the Nearctic

region. Colour images of selected specimens are included.

Key words: Introduced, Coleoptera, Diptera, Hemiptera, Hymenoptera, Trichoptera,

British Columbia, Canada

INTRODUCTION

During the course of our fieldwork over the past few years, and by determining older

unidentified material in the Spencer Entomological Collection, we have discovered 16

species never before recorded from British Columbia, including seven species new to

Canada. Nine of the species have been introduced from their native ranges, and seven are

range extensions for Nearctic species. We report these new records here.

Our field work during this time consisted mainly of monthly forays to local areas,

participation in bioblitzes and species surveys outside the Lower Mainland (which

concentrate efforts on a select locale for a short period of time), and an ongoing survey of

the "green roof" atop the Vancouver Convention Centre.

All specimens recorded here are deposited in either the Spencer Entomological

Collection [SEM] or the first author’s personal collection [CGR]. All photos were taken

by the first author using a Leica DFC490 digital camera mounted on a Leica M205C

stereomicroscope. Post-processing of the images was done using Adobe Photoshop CS4.

INTRODUCED INSECTS

The following nine species are native to the Palearctic region and have been recently

introduced to British Columbia. Three of these are new to Canada.

Coleoptera: Chrysomelidae

Cassida rubiginosa Miller

The leaf beetle, Cassida rubiginosa Miiller, has been recorded from British Columbia

for the first time. Native to Europe and Asia, this species was first introduced to North

America in Quebec around 1902 and is now found throughout most of Canada east of the

Rockies and in the northeastern United States. Members of the Family Asteraceae,

specifically thistles, are the preferred host plant for C. rubiginosa, although they are

polyphagous (Majka and Lesage 2008). The specimens collected in Richmond at both

Terra Nova Rural Park and Iona Beach Regional Park were found feeding on thistle.

New Records: 3 adults, Richmond, Terra Nova Rural Pk., 49.1660, —123.1956,

15.viii.2013 (C. G. Ratzlaff) [CGR, SEM]; 1 adult, Richmond, Iona Beach Reg. Pk.,

49.21940, —123.20839, 12.ix.2014 (C. G. Ratzlaff) [CGR]; 1 adult, Richmond, Iona

! Corresponding Author: Spencer Entomological Collection, Beaty Biodiversity Museum, 2212 Main

Mall, Vancouver, BC V6T 1Z4; chris.ratzlaff@gmail.com

80 J. ENTOMOL. SOC. BRIT. COLUMBIA 113, DECEMBER 2016

Beach Reg. Pk., 49.2196, —123.2083, 13.v.2015 (C. G. Ratzlaff) [CGR] (Fig. 1a); 1 adult,

Richmond, Iona Beach Reg. Pk., 49.220, —123.211, 1.vii.2015 (C. G. Ratzlaff) [SEM]

Figure 1. (a) Cassida rubiginosa Miller from Iona Beach Regional Park, in Richmond, B.C.;

(b) Brumus quadripustulatus Linnaeus from the University of British Columbia (UBC)

Botanical Garden, in Vancouver, B.C.; and, (c) Chilocoris bipustulatus (Linnaeus) from the

Beaty Biodiversity Museum at UBC, in Vancouver.

Coleoptera: Coccinellidae

Brumus quadripustulatus Linnaeus

The pine ladybird, Brumus quadripustulatus Linnaeus, used widely in the Palearctic

region as a biocontrol agent, was first introduced and established in California between

1915 and 1928 as a control for woolly hemlock adelgid (Gordon 1985). There are no

known further releases of B. quadripustulatus, and the species has apparently dispersed

north along the Pacific Coast. Year-round specimen records show that this species can

overwinter as an adult. These represent the first records of this species in Canada.

New Records: | adult, Richmond, 49.1888, —123.0959, 16.vi.2013 (C. G. Ratzlaff)

[CGR]; 1 adult, Vancouver, 49.2281, —123.0870, 13.iv.2014 (C. G. Ratzlaff) [CGR]; 2

adults, Vancouver, UBC Botanical Garden, 49.254, —123.247, 16.v.2014 (C. G. Ratzlaff)

[CGR, SEM] (Fig. 1b); 1 adult, Vancouver, UBC Botanical Garden, 49.254, —123.247,

4.x1.2014 (C. G. Ratzlaff) [CGR]; 1 adult, Vancouver, 49.2281, —123.0870, xi.2014 (C.

G. Ratzlaff) [CGR]; 1 adult, Vancouver, Pacific Spirit Reg. Pk., 49.2445, —123.2002, 8.11.

2016 (C. G. Ratzlaff) [SEM]; 1 adult, Vancouver, UBC, Beaty Biodiversity Museum,

49.2632, —123.2502, 22.11.2016 (C. G. Ratzlaff) [SEM]; 2 adults, Vancouver, UBC,

Beaty Biodiversity Museum, 49.2632, —123.2502, 1.iv.2016 (C. G. Ratzlaff) [SEM]; 2

adults, New Westminster, Sapperton Landing Pk., 49.2166, —12.8929, 11.iv.2016 (C. G.

Ratzlaff) [SEM]; 1 adult, Burnaby, Deer Lake Pk., 18.viii.2016 (K. Needham) [SEM]

Coleoptera: Coccinellidae

Chilocoris bipustulatus (Linnaeus)

Chilocoris bipustulatus (Linnaeus), a species widespread in the Palearctic region, first

became established in North America when it was introduced in California in 1951 as a

biocontrol agent for scale insects (Gordon 1985). Although it likely has been released in

a number of locations throughout North America, C. bipustulatus is poorly adapted to

low temperatures and will not survive cold winters (Kehat et a/. 1970). Our records from

Vancouver, the first for Canada, may be the direct result of biocontrol releases where

recent mild winters may have enabled this species to persist. Our first specimens were

J. ENTOMOL. Soc. BRIT. COLUMBIA 113, DECEMBER 2016 81

collected at Burns Bog during a B.C. Ministry of Environment, Lands, and Parks

Ecosystem Review (Kenner and Needham 1999), but were misidentified as Chilocoris

tricyclus Smith, a similar-looking native species.

New Records: 3 adults, Delta, Delta Nature Reserve, 11.x.1999 (R. Kenner and K.

Needham) [SEM]; 1 adult, Vancouver, Queen Elizabeth Pk., 49.2442, —123.1121, 15.v.

2013 (C. G. Ratzlaff) [SEM]; 1 adult, Vancouver, 49.2281, —123.0870, 17.x.2013 (C. G.

Ratzlaff) [CGR]; 1 adult, Vancouver, UBC Botanical Garden, 49.254, —123.247, 17.1v.

2015 (C. G. Ratzlaff) [SEM]; 2 adults, Vancouver, UBC, Beaty Biodiversity Museum,

49.2632, —123.2502, 19.iv.2016 (C. G. Ratzlaff) [SEM] (Fig. 1c)

Diptera: Opomyzidae

Geomyza tripunctata (Fallén)

The Palearctic opomyzid, Geomyza tripunctata (Fallén), commonly known as the

cereal fly, is recorded for the first time in British Columbia. In Canada, it has previously

been recorded from Ontario, Quebec, Prince Edward Island, and Nova Scotia (Wheeler ef

al. 1999). The larvae feed on the shoots of various grasses and have the potential to be a

crop pest.

New Records: | adult, Vancouver, 49.2281, —123.0870, ix.2010 (C. G. Ratzlaff)

[CGR]; 1 adult, Vancouver, 49.2281, —123.0870, 27.vi.2013 (C. G. Ratzlaff) [SEM]; 1

adult, Abbotsford, 49.0782, —122.3130, 18.viii.2013 (C. G. Ratzlaff) [SEM]; 1 adult,

Vancouver, Memorial South Pk., pond, 49.2321, —123.0869, 5.ix.2013 (C. G. Ratzlaff)

[CGR]; 1 adult, Richmond, Iona Beach Reg. Pk., 49.2276, —123.2299, 11.iv.2014 (C. G.

Ratzlaff) [CGR]; 1 adult, Mayne I., Miner's Bay, 48.8504, —123.3013, 15.x1.2014 (C. G.

Ratzlaff) [SEM]; 2 adults, Vancouver, Vancouver Convention Centre, Green Roof,

49.2887, —123.1162, 8.iv.2016 (C. G. Ratzlaff and K. Needham) [SEM] (Fig. 2a); 1 adult,

Sidney I., Dragonfly Pond, 48.6033, —123.3046, 14.vi1i.2016 (SEM Team) [SEM]

Figure 2. (a) Geomyza tripunctata (Fallen) from the green roof at the Vancouver Convention

Centre, Vancouver, B.C.; (b) Pseudomalus auratus (Linnaeus) from Woods Island Park, in

Richmond, B.C.

Hemiptera: Miridae

Atractotomus magnicornis (Fallén)

The plant bug, Atractotomus magnicornis (Fallén), is recorded for the first time in

British Columbia. Native and widespread in Europe, A. magnicornis is currently found in

Ontario, Quebec, Nova Scotia, and Newfoundland, as well as in several states in the

82 J. ENTOMOL. SOc. BRIT. COLUMBIA 113, DECEMBER 2016

eastern United States (Stonedahl 1990; Maw et al. 2000). This species is associated with

conifers, primarily members of Abies and Picea, often in gardens or other ornamental

environments (Stonedahl 1990).

New Records: 19, Coquitlam, Westwood Plateau Ridge Pk., 6.vii.2013 (T. Loh)

[SEM]; 32, Vancouver, UBC, Beaty Biodiversity Museum, 49.2632, —123.2502, 15.vii.

2016 (C. G. Ratzlaff) [SEM]

Hymenoptera: Chrysididae

Chrysis angolensis Radoszkowski

The large cuckoo wasp, Chrysis angolensis Radoszkowski, a cleptoparasite of mud-

daubers in the genus Sceliphron, is recorded for the first time in British Columbia. A

common host, Sceliphron caementarium (Drury), often builds mud nests on human-made

structures that are then transported and has been spread throughout many parts of the

world, facilitating the cosmopolitan distribution of C. angolensis (Kimsey 2006). In

North America, C. angolensis is thought to have been originally introduced to the eastern

US during World War II and has since spread to the western states, eastern Canada, and

parts of Mexico (Bohart and Kimsey 1982).

New Records: 1°, Vancouver, UBC Campus, 49.2593, —123.2477, 12.1x.2013 (C. G.

Ratzlaff) [SEM]; 19, Richmond, Terra Nova Rural Pk., 49.173, —123.199, 4.x.2015 (J.

Chan) [SEM] (Fig. 3)

Figure 3. Female Chrysis angolensis Radoszkowski collected from Terra Nova Rural Park, in

Richmond, B.C.

Hymenoptera: Chrysididae

Pseudomalus auratus (Linnaeus)

The small cuckoo wasp, Pseudomalus auratus (Linnaeus), has recently been

introduced to the Vancouver area. Native to Europe, Asia, and North Africa, this species

was introduced to the eastern United States sometime before 1828. It was recorded from

Utah in the late 1960s and then in California in 1980 (Bohart and Kimsey 1982). It has

since spread to a number of other states, as well as to eastern Canada. The hosts for P.

auratus, a cleptoparasite, are stem-nesting pemphredonine (Crabronidae) wasps, and it

has been suggested that its spread is a result of the transportation of garden plants (Danks

1971; Bohart and Kimsey 1982).

J. ENTOMOL. SOC. BRIT. COLUMBIA 113, DECEMBER 2016 83

New Records: 1¢ 29, Vancouver, 49.2281, —123.0870, 17.vi.2013 (C. G. Ratzlaff)

[CGR, SEM]; 14, Vancouver, 49.2281, —123.0870, 10.vi.2014 (C. G. Ratzlaff) [SEM]; 1

adult, Richmond, Iona Beach Reg. Pk., 49.22177, —123.21223, 25.vi.2014 (C. G.

Ratzlaff) [CGR]; 1 adult, Vancouver, 49.2281, —123.0870, 29.vi.2014 (C. G. Ratzlaff)

[CGR]; 1 adult, North Saanich, Schwartz Bay Terminal, 49.6883, —123.4099, 17.vi1.2015

(C. G. Ratzlaff) [CGR]; 1 adult, Vancouver, 49.2281, —123.0870, 6.v.2016 (C. G.

Ratzlaff) [SEM]; 1 adult, Richmond, Woods I. Pk., 49.2119, —123.1646, 14.vii1.2016 (C.

G. Ratzlaff) [SEM] (Fig. 2b)

Hymenoptera: Crabronidae

Rhopalum gracile Wesmael

The crabronid wasp, Rhopalum gracile Wesmael, is recorded here for the first time in

North America from Iona Beach Regional Park in Richmond, British Columbia. This

small wasp lives throughout the Palearctic region from England to Japan, although it is

typically uncommon (Bitsch and Leclercq 2009). It nests in the stems of plants such as

Euthamia occidentalis Nuttall and species of Phragmites (Bitsch & Leclercq 2009); the

latter genus grows near the specimen collection location. Rhopalum gracile was probably

introduced by means of aircraft, because Iona Beach is close to Vancouver International

Airport. It is unknown whether a population will become established; further monitoring

is needed.

New Record: 14 19, Richmond, Iona Beach Reg. Pk., 49.2196, —123.2083, 22.vii.

2015 (C. G. Ratzlaff) [SEM] (Fig. 4)

Figure 4. Rhopalum gracile Wesmael (a) male and (b) female from Iona Beach Regional

Park, in Richmond, B.C.

Hymenoptera: Torymidae

Megastigmus aculeatus (Swederus)

The parasitic wasp, Megastigmus aculeatus (Swederus), is recorded for the first time

in western North America. This species develops in the seed buds of Rosa species and

reproduces primarily through thelytokous parthenogenesis. Although native to western

Europe, M. aculeatus has spread over much of the globe and now occurs throughout

eastern Europe, parts of Africa, Iran, Iraq, China, Japan, Australia, and eastern North

America, including Ontario and Quebec. More than 23 host species of Rosa have been

recorded in the West Palearctic region, including several that have been introduced to

North America (Roques and Skrzypcezynska 2003). The wide commercial transport of

Rosa species, along with their parthenogenic reproduction, allow for easy colonization of

areas by M. aculeatus.

84 J. ENTOMOL. SOC. BRIT. COLUMBIA 113, DECEMBER 2016

New Record: 19, Whistler, Creekside Field, 50.0942, —122.9874, 9.vii.2016, 650m,

light trap (SEM Team) [SEM] (Fig. 5)

Figure 5. Female Megastigmus aculeatus (Swederus) from Creekside, in Whistler, B.C.

SIGNIFICANT RANGE EXTENSIONS OF NEARCTIC INSECTS

The following seven species are native to North America; these records represent

range extensions into British Columbia. Four of these species are also new to Canada.

Diptera: Blephariceridae

Philorus californicus (Hogue)

The net-winged midge, Philorus californicus (Hogue), is recorded for the first time in

Canada. This also represents the first record of the genus in Canada. Members of this

family of flies are associated with fast-flowing streams, where the larvae live on the

surfaces of rocks, often in cracks and crevices (Hogue 1973).

New Record: 1 larva, Lindell Beach, nr. Stillwood Camp, Watt Cr., 49.024,

—121.999, 17.vi.2014, under rock in fast-flowing stream (C. G. and N. A. Ratzlaff)

[SEM] (Fig. 6)

Diptera: Ulidiidae

Chaetopsis massyla (Walker)

The picture-wing fly, Chaetopsis massyla (Walker), is recorded for the first time from

British Columbia, collected during the 2015 Gulf Islands National Park Reserve BioBlitz

on Saturna Island. This species is widespread throughout most of North America and

feeds on damaged stems of wetland monocots, such as cattails and sedges (Steyskal

1965; Allen and Foote 1992). Chaetopsis massyla is also a pest of corn in several south-

eastern states (Goyal et al. 2010). |

New Record: | adult, Saturna I., Gulf Islands Nat. Pk. Res., Winter Cove, 48.8123,

—123.1879, 17.vii.2015 (C. G. Ratzlaff) [SEM] (Fig. 7)

Hemiptera: Miridae

Orectoderus montanus Knight

The plant bug, Orectoderus montanus Knight, is recorded for the first time in British

Columbia. The genus Orectoderus Uhler was recently revised and now contains five

J. ENTOMOL. SOC. BRIT. COLUMBIA 113, DECEMBER 2016 85

species, with three of those species reported from Canada (Nyniger 2010). To date, O.

montanus has been recorded in Canada only from Alberta and Saskatchewan (Kelton

1980; Maw et al. 2000; Nyniger 2010). In the United States, it has been recorded from

Colorado, Idaho, Montana, Nevada, North Dakota, Utah, and Wyoming. Hosts for O.

montanus include Ericameria nauseosa (Pall. ex Pursh) (Asteraceae), Symphoricarpos

spp. (Caprifoliaceae), and Potentilla fruticosa Linnaeus (Roseacae) (Nyniger 2010).

New Record: 13, Pink Mt., 57.0487, —122.8687, 2.vii.2016, 1715m (C. G. and N. A.

Ratzlaff) [SEM]

Figure 6. (a) Dorsal and (b) ventral view of a Philorus californicus (Hogue) larva from Watt

Creek, south of Cultus Lake, B.C., showing the suction cups on each segment used to hold

onto the surface of rocks in fast-flowing streams.

Hymenoptera: Crabronidae

Spilomena barberi Krombein

The small cryptic wasp, Spilomena barberi Krombein, is identified for the first time

from British Columbia. It has been recorded in Canada, from Ontario and Quebec, and in

the United States, from coast to coast (Buck 2004). Little is known about the biology of

S. barberi, and it is rarely collected.

New Record: 19, Osoyoos, Haynes Ecological Reserve, 13.vii. - 17.vii.1988, pitfall

trap, Purshia/Aristida shrub-steppe (S.G. Cannings) [SEM]; 19, Hornby Island, Norman

Pt., 9.vil.1989 (S. G. Cannings) [SEM]

Hymenoptera: Platygastridae

Synopeas anomaliventre (Ashmead)

The parasitic wasp, Synopeas anomaliventre (Ashmead), is identified for the first

time from Canada. It has previously been recorded from Maryland, Pennsylvania,

Florida, Louisiana, and New Hampshire, in the eastern United States (Muesebeck 1979).

86 J. ENTOMOL. SOC. BRIT. COLUMBIA 113, DECEMBER 2016

Many small parasitic wasps have widespread distributions, and whether this disjunct

record is natural or the result of an accidental introduction is unknown. Nothing is known

of the biology of this species, but others in the genus are parasitic on flies in the Family

Cecidomyliidae (Fouts 1924).

New Record: 1, Galiano I., north end, 19.iv.1981 (S. G. Cannings) [SEM]

Figure 7. Chaetopsis massyla (Walker) from Winter Cove in Gulf Islands National Park

Reserve, on Saturna Island, B.C.

Hymenoptera: Pompilidae

Ceropales pacifica Townes

The spider wasp, Ceropales pacifica Townes, is recorded from Canada for the first

time, previously being recorded only from Oregon and California. No specific

information about the biology of C. pacifica is known, but members of the genus are

kleptoparasites of other wasps in the Family Pompilidae. Females lay an egg in the book

lungs of a host wasp’s unattended spider prey before it is deposited in the host’s nest.

When hatched, the Ceropales larva will consume both the spider and the egg laid by the

host wasp (Townes 1957). :

New Records: 1419, Oliver, UBC Geology Camp, 20.vii.1989, malaise trap, pine/

thicket edge (S. G. Cannings) [SEM]; 23, Oliver, UBC Geology Camp, 22.vii.1989,

malaise trap, pine/thicket edge (S. G. Cannings) [SEM]; 1, Oliver, UBC Geology

Camp, 22.vii.1990, malaise trap, ponderosa pine forest (S. G. Cannings) [SEM]; 19,

Oliver, UBC Geology Camp, 27.vii.1990, malaise trap, hawthorn thicket edge (S. G.

Cannings) [SEM] (Fig. 8); 1¢', Oliver, UBC Geology Camp, 28.vii.1990, malaise trap,

hawthorn thicket edge (S. G. Cannings) [SEM]

Trichoptera: Limnephilidae

Desmona mono (Denning)

Desmona is an unusual genus of caddisfly in the Family Limnephilidae. Its members

inhabit the shallow edges of high-alpine lakes and springs or seeps of alpine meadows

(Wiggins 1996). Although most limnephilids are detritivores, feeding on dead and

decaying plant material at the bottom of these lakes, Desmona crawl out of the water at

night to feed on living plants near the water's edge (Wiggins and Wisseman 1990).

In North America, three species are recognized in the genus (Nimmo 2012). Desmona

bethula Denning and Desmona denningi Nimmo are found in California, while Desmona

J. ENTOMOL. Soc. BRIT. COLUMBIA 113, DECEMBER 2016 87

mono (Denning) (= Monophylax mono) inhabits the Pacific Northwest of the United

States.

During surveys of some high-alpine lakes, streams, and seeps atop Whistler Mountain

as part of the 2014 Whistler Bioblitz, we collected several larvae and cases of Desmona,

which were later identified as D. mono. These represent the first record of this genus and

species for British Columbia and Canada.

New Records: 9 larvae, Whistler, Blackcomb Mt., Overlord Trail wetland, 50.080,

—122.888, 23.viii.2014 (C. M. Stinson) [SEM] (Fig. 9); 3 larvae, Whistler, Whistler Mt.,

Harmony L., 50.0640, —122.9379, 3.ix.2014, 1760m (C. G. Ratzlaff) [SEM]; 8 larvae,

Whistler, Whistler Mt., Harmony L., 50.0640, —122.9379, 28.ix.2014, 1780m (C. G.

Ratzlaff) [SEM]; 1 larva, Whistler, Whistler Mt., alpine stream, 50.0522, —122.9350,

28.ix.2014, 1810m (C. G. Ratzlaff) [SEM]; 1 larva, Whistler, Blackcomb Mt.,

Blackcomb L., 50.0813, —122.8812, 27.vi.2015, 1800m (C. G. Ratzlaff) [SEM]

Figure 8. Ceropales pacifica (Townes) from the U eology Camp, in Oliver, B.

CONCLUSION

Examination of these new records highlights the importance of concentrated surveys

targeting specific locales, as well as the usefulness of repeated forays to the same area.

Seasonal and yearly variation in species presence makes it imperative to revisit a site

periodically in order to produce a comprehensive species list for an area.

The value of the taxonomic expertise of previous researchers and detailed reference

material, such as exists in provincial and national research collections, cannot be

underestimated in projects such as these. Cataloguing the biodiversity of an area would

not be possible without these resources. Adding to the historical record of species

distributions and seasonal occurrences is valuable both in and of itself and as a

foundation for further research.

88 J. ENTOMOL. SOC. BRIT. COLUMBIA 113, DECEMBER 2016

Figure 9. (a) Larva and case of Desmona mono (Denning) from Overlord Trail on Blackcomb

Mountain, in Whistler, B.C.; (b) The lateral hump showing the two ring-like sclerites found in

the larvae of Desmona species.

ACKNOWLEDGEMENTS

We’d like to thank organizers of the Whistler Bioblitz, Bob Brett and Kristina

Swerhun, and of the Parks Canada Bioblitz, Athena George and Madelin Emery. We’d

also like to thank Paul Hunter of the Vancouver Convention Centre and Patrick Lewis of

the UBC Botanical Gardens for allowing us access and facilitating our sampling efforts.

All photographs in this paper were taken by the first author with digital imaging

equipment funded by a Canadian Foundation for Innovation infrastructure grant.

REFERENCES

Allen, E. J., and B. A. Foote. 1992. Biology and immature stages of Chaetopsis massyla (Diptera:

Otitidae), a secondary invader of herbaceous stems of wetland monocots. Proceedings of the

Entomological Society of Washington 94:320-328.

Bitsch, J., and J. Leclercq. 2009. Complément au volume 1 des Hyménoptéres Sphecidae d’Europe

occidentale (Faune de France 79). Bulletin de la Société Entomologique de France 114:211—244.

Bohart, R. M., and L. S. Kimsey. 1982. A synopsis of the Chrysididae in America North of Mexico.

Memoirs of the American Entomological Institute 33:1—266.

Buck, M. 2004. An annotated checklist of the Spheciform wasps of Ontario (Hymenoptera: Ampulicidae,

Sphecidae and Crabronidae). Journal of the Entomological Society of Ontario 134:19-84.

Danks, H. 1971. Biology of some stem-nesting aculeate Hymenoptera. Transactions of the Royal

Entomological Society of London 122:323-399.

Fouts, R. M. 1924. Revision of the North American wasps of the subfamily Platygasterinae. Proceedings

of the United States National Museum 63:1—145.

Gordon, R. D. 1985. The Coccinellidae (Coleoptera) of America North of Mexico. Journal of the New

York Entomological Society 93(1):1—912.

Goyal, G., G. S. Nuessly, G. J. Steck, D. R. Seal, J. L. Capinera, and K. J. Boote. 2010. New report of

Chaetopsis massyla (Diptera: Ulidiidae) as a primary pest of corn in Florida. Florida Entomologist

93:198-202.

Hogue, C. L. 1973. The netwinged midges or Blephaceridae of California. Bulletin of the California

Insect Survey, Vol. 15. University of California Press, Berkeley. 83 pp.

Kehat, M., S. Greenberg, and D. Gordon. 1970. Factors causing seasonal decline in Chilocoris

bipustulatus L. (Coccinellidae) in citrus groves in Israel. Entomophaga 15:337—345.

Kelton, L. A. 1980. The plant bugs of the prairie provinces of Canada. Heteroptera: Miridae. The Insects

and Arachnids of Canada. Part 8. Research Branch, Agriculture Canada Publications 1703. 408 pp.

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Kenner, R. D., and K. M. Needham. 1999. Burns Bog Ecosystem Review: Invertebrate Component.

Environmental Assessment Office. B.C. Ministry of Environment, Lands, and Parks, Victoria, BC.

271 pp.

Kimsey, L. S. 2006. California Cuckoo Wasps in the Family Chrysididae. University of California

Publications in Entomology 28:1—319.

Majka, C. G., and L. Lesage. 2008. Introduced leaf beetles of the Maritime Provinces, 7: Cassida

rubiginosa Miiller and Cassida flaveola Thunberg (Coleoptera: Chrysomelidae). Zootaxa 1811:37—

56.

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

of Canada and Alaska. NRC Research Press. Ottawa. 220 pp.

Muesebeck, C. F. W. 1979. Superfamily Proctotrupoidea, pp. 1121-1186. Jn K. V. Krombein, P. D. Hurd,

Jr, D. R. Smith, and B. D. Burks. Catalog of Hymenoptera in America north of Mexico: Volume 1,

Symphyta & Apocrita (Parasitica). Smithsonian Institution Press, Washington, DC. 1198 pp.

Nimmo, A. P. 2012. Review of the three genera Desmona Denning, Monophylax Nimmo, and

Psychoglypha Ross, with descriptions of 12 new species (Insecta, Trichoptera: Limnemphilidae;

Limnephilinae; Chilostigmini). Occasional Papers on Trichoptera Taxonomy Number 2. Edmonton.

73 pp.

Nyniger, D. 2010. Resurrection of the Protocrepini Knight, with revisions of the Nearctic genera

Orectoderus Uhler, Pronotocrepis Knight, and Teleorhinus Uhler and comments on the Palearctic

Ethelastia Reuter (Heteroptera: Miridae: Phylinae). American Museum Novitates 3703. 67 pp.

Roques, A., and M. Skrzypczynska. 2003. Seed-infesting chalcids of the genus Megastigmus Dalman,

1820 (Hymenoptera: Torymidae) native and introduced to the West Palearctic region: taxonomy, host

specificity and distribution. Journal of Natural History 37:127-238.

Steyskal, G. C. 1965. Family Otitidae, pp. 642-654. Jn Stone, A., C.W. Sabrosky, W.W. Wirth, R.H.

Foote, and J.R. Couslon, eds., A Catalog of the Diptera of America North of Mexico. USDA

Agriculture Handbook 276, 1696 pp.

Stonedahl, G. M. 1990. Revision and cladistic analysis of the Holarctic genus Atractotomus Fieber

(Heteroptera: Miridae: Phylinae). Bulletin of the American Museum of Natural History 198. 88 pp.

Townes, H. 1957. Nearctic wasps in the subfamilies Pepsinae and Ceropalinae. United States National

Museum Bulletin 209. 286 pp.

Wiggins, G. B. 1996. Larvae of the North American caddisfly genera (Trichoptera). 2nd Edition.

University of Toronto Press, Toronto. 457 pp.

Wiggins, G. B., and R. W. Wisseman. 1990. Revision of the North American caddisfly genus Desmona

(Trichoptera: Limnephilidae). Annals of the Entomological Society of America 83(2):155—161.

Wheeler, T. A., J. R. Vockeroth, and S. Boucher. 1999. Geomyza tripunctata Fallén, a Palaearctic

opomyzid fly new to North America, with notes on range expansions in Holarctic Opomyzidae

(Diptera). Proceedings of the Entomological Society of Ontario 130:5-20.

90 J. ENTOMOL Soc. BRIT. COLUMBIA 113, DECEMBER 2016

NATURAL HISTORY AND OBSERVATIONS

Rhyparochromus vulgaris (Schilling) (Hemiptera:

Heteroptera: Rhyparochromidae): newly discovered in the

interior of British Columbia

G. G. E. SCUDDER!

The Palaearctic seed bug Rhyparochromus vulgaris (Schilling) was first reported

from Oregon and Washington in North America by Henry (2004). The species is here

reported for the first time in the interior of British Columbia (B.C.).

In early 2015, Mike Art observed many specimens in a woodshed in Creston, B.C.

Through the cooperation of Ward Strong, Arthur Stock and Karen Needham, Kristine

Sacenieks collected a series of specimens for me on 1 May 2015, at Creston, Goat River,

851 Aldich Road 49.075986°N 116.517452°W (UTM 11 —0535241E x 5436015W) at an

elevation of 546 metres. Using the keys and description in Peéricart (1998) and

comparison with specimens from several localities in Europe, I confirmed that the

species was Rhyparochromus vulgaris (Schilling) (Fig. 1). The sample received

contained nineteen males and four females of Rhyparochromus vulgaris, plus two males

of Arhyssus scutatus (Stal) (Rhopalidae). This is the first record of Rhyparochromus

vulgaris from the interior of British Columbia. However, previous to this, Dr. M. D.

Schwartz (Ottawa) had identified three males and six females of Rhyparochromus

vulgaris for the Canadian Food Inspection Agency (CFIA) that had been collected at a

commercial nursery near Langley, B.C., on 28 February 2013. These were discovered in

an outdoor covered storage area near boxes of supplies. These specimens are deposited in

the Canadian National Collection of Insects (CNC). The Creston specimens have been

deposited in the CNC, Royal British Columbia Museum (RBCM), University of British

Columbia (UBC), and my own collection.

According to Péricart (1998), Rhyparochromus vulgaris occurs naturally throughout

Europe and the Mediterranean region, from Belgium, Germany and Russia in the north

around the 55" parallel, to Asia Minor in the south. Péricart (1998) had no records for the

British Isles.

Figure 1. Rhyparochromus vulgaris (Schilling). B.C. specimen, male, dorsal view. Length

6.75 mm.

! Department of Zoology and Biodiversity Research Centre, University of British Columbia, 6270

University Boulevard, Vancouver, B.C. V6T 1Z4.

J. ENTOMOL SOc. BRIT. COLUMBIA 113, DECEMBER 2016 91

ACKNOWLEDGEMENTS

I thank Kristine Sacenieks, Arthur Stock and Ward Strong (Government of British

Columbia), as well as Karen Needham (UBC) for providing me with the specimens from

Creston. Dr. M. D. Schwartz (Ottawa) kindly provided information on the CFIA

collection. Don Griffiths (UBC) took the photograph for Fig. 1, and Launi Lucas (UBC)

kindly processed the manuscript.

REFERENCES

Henry, T. J. 2004. Raglius alboacuminatus (Goeze) and Rhyparochromus vulgaris (Schilling): two

Palearctic bugs newly discovered in North America. Proceedings of the Entomological Society of

Washington 106(3): 513-522.

Péricart, J. 1998. Hémiptéres Lygaeidae Euro-Méditerranéens. Volume 3. Systématique: Troisiéme

Partie. Faune de France 84C: 487 pp.

92 J. ENTOMOL. SOC. BRIT. COLUMBIA 113, DECEMBER 2016

Proceedings of the Pollination Science and Stewardship

Symposium

Okanagan College Campus, Penticton, British Columbia, March 17,

2016

INTRODUCTION

Jennifer Heron, Cory S. Sheffield, and Cara Dawson

Pollination is a vital ecological process in terrestrial ecosystems. Pollinators are the

organisms that provide this service, thus facilitating reproductive success in plant

communities. Awareness of the importance of pollinators and pollination has increased in

the last few decades, largely due to global declines in honey bee colonies and

documented widespread declines of some bumble bee and other pollinator species. In

Canada, the main pollinators are insects, with thousands of species from a wide range of

taxa that regularly visit flowers. Among the most familiar and important insect

pollinators are the butterflies and moths (Lepidoptera), many groups of flies (Diptera)

and beetles (Coleoptera), and the Hymenoptera, which include the bees and many other

types of wasps.

The symposium, Pollination Science and Stewardship, featured 12 presenters who

spoke about current research and on topics relating to the diversity, conservation,

ecosystem services, pesticide management, agriculture, citizen science, and stewardship

for pollinators, with focus on Canada. The workshop also aimed to facilitate connections

between pollination specialists and land managers, owners, stewards and biologists, thus

enabling information and idea exchange such that these practitioners could then apply it

to their own conservation work. More than 90 people attended the symposium.

Participants included members of academia, industry professionals, agriculturalists,

citizen scientists, artists, students, gardeners, and landowners interested in enhancing

their properties for pollinators.

The symposium was supported by funding from Environment and Climate Change

Canada Habitat Stewardship Program for Species at Risk, the British Columbia Ministry

of Environment, the Royal Saskatchewan Museum, and the Entomological Society of

British Columbia.

Butterflies, conservation, and citizen science in Canada

John Acorn, University of Alberta, Faculty of Agricultural, Life and Environmental

Sciences, Department of Renewable Resources, 777 General Services, Edmonton, AB

T6G 2H1; email: jacorn@Qualberta.ca

Butterflies are colourful, relatively easy to identify, and popular with naturalists.

Butterflies are also potential pollinators, even though they appear to be relatively delicate

and clean when compared to furry, flower-wrestling, pollen-covered bees and syrphid

flies. Recent work, however, shows the importance of non-syrphid flies to pollination

(Orford et al. 2015, DOI: 10.1098/rspb.2014.2934), and by analogy we know little about

the importance of butterflies. What we care about as biodiversity conservationists,

however, is not just pollinators, or the “ecosystem service” of pollination, but the

diversity and abundance of wild flowers and flower-visiting insects. In this regard,

butterflies are ideal “flagship” organisms for the conservation cause.

Butterfly collecting laid the groundwork for the understanding of butterfly faunistics,

and amateur collecting can be considered the original butterfly citizen science project.

The non-consumptive approach to butterfly citizen science began in Canada as “Fourth

of July Butterfly Counts” some 20 years ago, coordinated originally by the Xerces

Society, and more recently by the North American Butterfly Association. There are still a

J. ENTOMOL. SOC. BRIT. COLUMBIA 113, DECEMBER 2016 93

few such counts, including the Dry Island Buffalo Jump Provincial Park Butterfly Count,

in Alberta. This count involves park rangers with nets, and counters the common

perception that nets are evil. However, the data from such counts is poor, because of the

effect of weather on shifting phenologies and the difficulty of comparing a one-day

sample from one year to similar samples from other years. A better approach is to

_ conduct weekly Pollard Walks, along a standard transect or route. This method results in

much more valuable data, but it requires considerable effort, and has not caught on in

Canada, with a few exceptions, including my own Pollard route in Edmonton, which |

have visited regularly during the butterfly seasons since 1999.

Another approach involves the creation of butterfly atlas projects, such as the British

Columbia Butterfly Atlas, the Ontario Butterfly Atlas, and the Maritimes Butterfly Atlas.

These projects update both distributional and phenological databases, and they can be

popular. In Alberta, the Alberta Lepidopterists’ Guild (ALG) considered an atlas project,

but declined because of 1) a small number of potential participants for such a large

geographic area, 2) the fact that atlas projects are not open-ended, and are intended to

produce a book-like end product, and 3) the perceived redundancy of atlassing and the

citizen science project eButterfly.org.

Instead of an atlas, the ALG initiated the Alberta Butterfly Roundup in the spring of

2015, an open-ended attempt to reconfirm the 173 species of butterflies known from the

province. In the 2015 season, 53 naturalists participated, and of those, 33 contributed at

least one species to the count. A total of 120 species were reported, and the top

contributor found 24 of these. The Roundup helped document a new species for the

province (regal fritillary, Speyeria idalia) and the westward spread of a “native” species,

the dun skipper (Euphyes vestris). Although participants were encouraged to submit

records through eButterfly, most records were submitted through the ALG Facebook

Page and the ALG listserver (Albertaleps), both two to three times as popular as

eButterfly. eButterfly received about the same number of submissions as the Albertabugs

listserver, the Edmonton Natural History Club listserver, and emails directly to me. In the

future, and to find the remaining 52 species, ALG is encouraging directed searches for

species that are rare and localized in areas that are not often visited. The Roundup

approach may not possess the same scientific rigour as an atlas project, but it does focus

attention on butterflies, with potential benefits for conservation, and the monitoring of

pollinators and floral resources in general.

Biology and diversity of moths in Canada: a conservation perspective

Greg Pohl, Natural Resources Canada, Canadian Forest Service, Northern Forestry

Centre, 5320 122 Street NW, Edmonton, AB T6H 385; email: greg.pohl@canada.ca

Butterflies and moths comprise the order Lepidoptera, one of the four most diverse

orders of insects. Moths make up about 90% of Lepidoptera; the butterflies are just one

small branch of the group. Approximately 5,100 species of moths live in Canada, with

about 2,650 species known in B.C. Most moth larvae feed on living plant tissue, as

exposed or concealed feeders on leaves, or as borers in stems, roots, flowers, and fruit.

Not all moths feed as adults, but those that do are looking for food energy such as

nectar from flowers, which makes them potential pollinators. Many diurnal species are

generalist flower visitors, but some species are very specialized. Males of many diurnal

micromoth species, such as choreutids (Choreutidae) and fairy moths (Adelidae), patrol

around nectar sources to find females, and exploit whatever flower species are present.

Other moths, such as Greya and Lampronia (Prodoxidae), are more specialized. Females

lay eggs in the flowers of their host plants, and they have been observed pollinating while

ovipositing. In yucca moths (the Genus TJegeticula in the Prodoxidae), this specialization

has developed into the famous mutualism between moths and plants. Yucca plants are

completely dependent on yucca moths for pollination. Female yucca moths deposit

pollen on the flower stigma after injecting eggs into the yucca flower ovary. The larvae

then hatch and feed in the developing seed pods of the yucca plant. Thus, they require

94 . J. ENTOMOL. SOC. BRIT. COLUMBIA 113, DECEMBER 2016

pollination of the flower for their food to develop. The larvae eat only some of the

developing seeds, leaving some to produce the next generation of plants. This text-book

case of mutualism gets more complicated; two species of "cheating" yucca moths have

evolved. These cheaters lay eggs on developing seeds that have been fertilized by the

pollinator moth species, so they are completely dependent on the yuccas and the

pollinating yucca moth species.

Many orchids rely on specialized moth pollinators that have co-evolved with the

plants. Some orchids have pollinia—sticky anther tips that break off and attach to the

visiting insect—to aid pollen transfer. Although moth—orchid pollination information is

often scanty, several moth species have been observed with pollinia attached, so they

clearly play a role in orchid pollination. Some orchids have evolved a system of placing

the nectar at the base of a long spur, so the pollinating insect must reach deep into the

plant and increase its chances of bumping the pollinia. This has triggered an evolutionary

arms race where the insects evolve longer mouthparts to access nectar, and the plants

evolve longer spurs to promote pollination. An extreme case is Darwin's Orchid

(Angraecum sesquipedale), with a 40+ cm spur. At the time, no insect was known with

long enough mouthparts to reach the nectar, and Charles Darwin predicted in 1862 that a

moth must exist with the necessary mouthparts. It was discovered in 1902: Xanthopan

morganii praedicta has a 40-cm-long proboscis.

Biology, conservation & stewardship for flies in Canada

Andrew Young, Carleton University, Ottawa, ON; email: a.d.young@gmail.com

Many species of true flies (Diptera), especially flower flies (Syrphidae), are

significant wild pollinators. Diptera contribute approximately half of the pollination

services in most environments and become increasingly important with increasing

latitude. For example, flies are the primary (and often only) pollinators north of the

Arctic Circle, but despite this their ecological impact has been historically

underappreciated, and relatively little is known about the specifics of many plant—

pollinator interactions. Potential conservation efforts are currently hampered by

inadequate taxonomic knowledge, coupled with poor knowledge of distribution for most

species. Even within the taxonomically well-known family Syrphidae, there is a need for

increased collecting effort and development of widely accessible, user-friendly

identification tools. Several case studies of potentially endangered species and

conservation efforts are discussed.

Biology, conservation and stewardship for bees in Canada

Cory S. Sheffield, Royal Saskatchewan Museum, 2340 Albert Street, Regina, SK S4P

2V7; email: Cory.Sheffield@gov.sk.ca; website: royalsaskmuseum.ca/research-and-

collections/biology/cory-sheffield

Bees are the most important pollinators of both crop and native plant species.

Although a few managed species are thought to provide most of the pollination services

to crops, a growing body of evidence suggests that wild bee species play an important

role. This is especially true for native plant species, some of which have intimate

relationships with their bee pollinators. Over 800 species of bee are recorded from

Canada, most of these occurring in ecozones in the southern half of the country. Our

recent efforts have focused on documenting the patterns of diversity and distribution of

Canada's bees, this being facilitated with DNA barcoding. These initiatives have also

allowed for the first comprehensive assessment of the conservation status of the bee

fauna of Canada. However, there is still much to learn. This session covers our current

knowledge of bees in Canada, including their diversity, distribution, gaps in taxonomic

knowledge, and conservation status.

J. ENTOMOL. SOC. BRIT. COLUMBIA 113, DECEMBER 2016 95

Global bee diversity and conservation

Laurence Packer, York University, Lumbers Building 345, 4700 Keele Street, Toronto, ON

M3J 1P3, Canada; email: xeromelissa@gmail.com

More than 20,000 species of wild bee exist in the world, and almost none of them fit

the standard archetype people have as to what bees are. Those archetypes relate to the

domesticated honey bee, which is not native to North America; further, honey bees have

been shown to often not be as important for pollination of crops as wild bees are. This

presentation focuses on the taxonomic, ecological and behavioural diversity of the bees

of the world, and compare bee diversity globally with that from Canada.

The Federal General Status Program of Canada, Wild Species 2015

Leah Ramsay, British Columbia Ministry of Environment, British Columbia Conservation

Data Centre, Mezzanine Floor - 395 Waterfront Crescent, Victoria, B.C., Canada VSW

9M2,; email: Leah.Ramsay@gov.bc.ca

The General Status Program arose from the Accord for the Protection of Species at

Risk (1996) and one of the resulting sections of the Species at Risk Act (SARA), which

include the requirement to assess and report regularly on the status of all wild species in

Canada. The resulting amassing of data provides the baseline for assessing the status of

all species in Canada—including invertebrates. A report is completed every five years

and the results are published. The first step in the assessment process is the refinement of

lists for Canada and each province and territory for the species group that is being

reported on. The lists are based on published literature, museum collections and personal

knowledge of experts. The next step involves pulling together whatever basic

information is available on the distribution, trends, habitat, threats and range extent using

the same sources as for the lists. These are some of the criteria that are used to determine

a conservation status rank or general status rank.

The conservation status assessments for the 2015 Wild Species report were done

following NatureServe methodology. This is the same process that is used for the status

ranks that are provided by the Conservation Data Centres and Heritage programs within

each of the jurisdictions. The factors, methods and the calculator are all found at

www.natureserve.org. In British Columbia the results, including the resulting lists are

held and maintained with in the B.C. Conservation Data Centre. The first report in 2000

assessed 1,670 species (mainly vertebrates) and the 2015 report will assess

approximately 30, 000 (final number to be determined), including capsular and non-

vascular plants and many groups of invertebrates. One of the focusses for the 2015 report

was assessment of as many of the pollinator groups as possible, including the bumble

bees, which. The bumble bees had been done initially in 2010, as well as the rest of the

bees were assessed as well as moths, bee flies, beetles and wasps, to name a few. This

presentation describes the assessment process and methodology, as well as discusses a

number of ways the General Status Program has been used to improve knowledge and

benefit species conservation in Canada.

The General Status program originated in order to fulfill the requirements of SARA

of enabling a metric to use to determine overall trends in the biodiversity of species in

Canada. Other beneficial offshoots include (but are not limited to), helping to inform

assessment priorities for the Committee on the Status of Wildlife in Canada (COSEWIC),

to establish a baseline of data for all species, identify knowledge gaps by highlighting

species where little is known and further inventory is required and provide taxonomic

lists for all of the jurisdictions and Canada. One can also get a snapshot of the diversity

of different groups in the different regions of Canada as well as see where across the

jurisdiction something may be declining or in good shape.

96 | J. ENTOMOL. SOC. BRIT. COLUMBIA 113, DECEMBER 2016

Pollination, pollinator diversity, and the determinants of plant reproduction

Jana Vamosi, Department of Biological Sciences, University of Calgary, 2500 University

Drive NW, Calgary, AB T2NIN4; email: vamosi@ucalgary.ca

Understanding the role of pollinators in determining plant reproduction is critical for

conserving and restoring ecosystems, as well as for maintaining food security. While

some crop plants are self-compatible and require little input from pollinators to produce

fruit and seeds, many plant species in natural systems require pollinators to effect pollen

transfer. Recent research reveals that when many plant species are in an area, they

receive inadequate pollination and produce suboptimal levels of fruit and seeds, yet the

mechanisms behind these observed patterns are unclear (Vamosi et al. 2006). Species in

species-rich areas may be more specialized, having traits that attract certain pollinators

and restrict access of others (Vamosi et al. 2014). While these traits may confer

advantages when a favoured pollinator is present, they also put the plant species at risk of

lower reproduction if their specialist group of pollinators is absent. Here, I summarize a

number of approaches how specialization may affect pollen delivery and conserve the

ecosystem function of pollination.

The initial approach to evaluating whether a plant species is receiving adequate

pollination service from pollinators is to estimate “pollen limitation”, typically through a

manipulative experimental design where the proportion of fruits or seeds set under

natural pollination conditions is compared to that under supplemental pollination

conditions (Knight et al. 2005). Thousands of pollen limitation experiments have now

been conducted throughout the world, with estimates that ~60% of species are pollen-

limited. In other words, these pollen-limited species would produce more fruits or seeds

if pollinators were optimally abundant. Reasons for this widespread pollen limitation are

currently unclear but some have posited that the pattern is a reflection of an impending

“pollination crisis” (Ingram et al. 2002), evidenced by the observations that pollinators

are declining in abundance and diversity at an alarming rate (Potts et a/. 2010). While

pollen limitation can be observed to increase with disturbance and loss of pollinators (Da

Silva et al. 2013 ), it is important to recognize that natural processes may also cause plant

species to exhibit pollen limitation as well. For example, observational studies comparing

the pollen limitation of species that were visited by diverse arrays of pollinators were

often no less pollen limited than those populations that received visits from few species

of pollinators (Davila et al. 2012), indicating that we do not fully understand the

functional role of pollinator diversity in communities. In examining the contribution that

differences in floral visitor composition make to increased selfing and seed production of

plant populations, we find that increased visitation by particular functional groups

ensures reproductive success of focal plant species more so than pollinator diversity

(Adderley and Vamosi 2015). Thus, while entire flowering communities may benefit

from functionally diverse pollinator communities, the reproductive success of a single

pollinator species is more contingent on a specialized subset of pollinators.

Conversely, the prevalence of certain plant traits within a community can influence

pollinator composition (Bruckman and Campbell 2014). In an investigation of the effects

of changes in plant composition that altered the prevalence of zygomorphy (1.e., floral

symmetry and the restrictiveness of flowers to certain pollinators), the conversion to

grazing pasturelands negatively impacted species richness and phylogenetic diversity.

Changes in community composition and structure had strong effects on the prevalence of

zygomorphic species, likely driven by nitrogen-fixing abilities of certain clades with

zygomorphic flowers (e.g., Fabaceae). Land conversion can thus have unexpected

impacts on trait distributions relevant for the functioning of the community in other

capacities (e.g., cascading effects to other trophic levels (i.e., pollinators) (Villalobos and

Vamosi 2016). These patterns indicate that we may be able to predict which pollinators

will be available to various crop plants by understanding their relationships with the

floral community and climate (Kerr et a/. 2015).

J. ENTOMOL. SOC. BRIT. COLUMBIA 113, DECEMBER 2016 97

REFERENCES

Adderley, L., and J. Vamosi. 2015. Species and phylogenetic heterogeneity in pollinator visitation affects

selfing and seed production in an island system. International Journal of Plant Sciences 176:186—

196.

Bruckman, D., and D. R. Campbell. 2014. Floral neighborhood influences pollinator assemblages and

effective pollination in a native plant. Oecologia 176(2):465-476.

Da Silva, E. M., V. M. King, J. L. Russell-Mercier, and R. Sargent. 2013 Evidence for pollen limitation

of a native plant in invaded communities. Oecologia 172:469-476.

Davila, Y., E. Elle, J. Vamosi, L. Hermanutz, J. Kerr, C. Lortie, A. Westwood, T. Woodcock, and A.

Worley. 2012. Ecosystem services of pollinator diversity: a review of the relationship with pollen

limitation of plant reproduction. Botany 90:535—543.

Ingram, M, S. Buchmann, G. Nabhan. 2002. Our forgotten pollinators: Protecting the birds and the bees.

pp 191-207. Im: A. Kimbrell ed. The Fatal Harvest Reader: The tragedy of industrial agriculture.

Washington, D.C.: Island Press.

Kerr, J. T., A. Pindar, P. Galpern, L. Packer, S. G. Potts, S. M. Roberts, P. Rasmont, O. Schweiger, S. R.

Colla, and L. L. Richardson. 2015. Climate change impacts on bumblebees converge across

continents. Science 349(6244):177-180.

Knight, T. M., J. A. Steets, J. C. Vamosi, S. J. Mazer, M. Burd, D. R. Campbell, M. R. Dudash, M. O.

Johnston, R. J. Mitchell, and T.-L. Ashman. 2005. Pollen limitation of plant reproduction: pattern

and process. Annual Review of Ecology, Evolution, and Systematics 36:467—-497.

Potts, S. G., J. C. Biesmeijer, C. Kremen, P. Neumann, O. Schweiger, and W. E. Kunin. 2010. Global

pollinator declines: trends, impacts and drivers. Trends in Ecology and Evolution 25(6):345-353.

Vamosi, J., T. Knight, J. Steets, S. Mazer, M. Burd, and T.-L. Ashman. 2006. Pollination decays in

biodiversity hotspots. Proceedings of the National Academy of Sciences 103:956—961.

Vamosi, J. C., C. Moray, N. Garcha, S. A. Chamberlain, and A. Mooers. 2014. Pollinators visit related

plant species across 29 plant-pollinator networks. Ecology and Evolution 4:2,303-2,315.

Villalobos, S., and J. C. Vamosi. 2016. Increasing land use drives changes in plant phylogenetic diversity

and prevalence of specialists. PeerJ 4: 1740 https://doi.org/1710.7717/peerj.1740.

Pollination in Agriculture: Insects and Ecological Intensification

Peter Kevan, Arthur Dobbs Institute, University of Guelph, Guelph, ON; email:

pkevan@uoguelph.ca

Agricultural expansion and intensification are central to the current demise of

biodiversity, affecting plants, animals (including insects) and fungi. The new buzz-phrase

“ecological intensification” goes beyond the restoration of biodiversity to the

reestablishment of ecosystem functionality. According to Food and Agriculture

Organization of the United Nations (FAO 2013) it is “a knowledge-intensive process that

requires optimal management of nature’s ecological functions and biodiversity to

improve agricultural system performance, efficiency, and farmers’ livelihoods”. Animal

pollination accounts for approximately one-third of our food supply, and new data from

around the world indicate that wild pollinators are far more important than managed ones

in many cropping systems. Yield drags of 20-30% have been documented for numerous

crops often stated to not require insect pollination, new cultivars are not tested for their

pollination requirements and their floral ecology is generally ignored, and scant attention

and little regard has been paid beyond bees to other insect groups essential to pollination

ecosystem services. Landscape management in agricultural and urban environments that

is focused on pollinator habitats has, at the same time, encouraged populations of

biocontrol agents, notably parasitoids and some predators that depend on floral resources

for part of their life cycles. The FAO’s call to “optimize management [as] a knowledge-

intensive process that requires optimal management of nature’s ecological functions and

biodiversity to improve agricultural system performance, efficiency and farmers’

livelihoods” strongly suggests intensive and well-informed human intervention. An

example may be the use of managed pollinators to disseminate microbial biological

control agents against crop pathogens and insects pests on crops. The multiple benefits of

better yields through pollination plus crop protection that are coupled to reduced uses of

chemical pesticides, conservation of water and less consumption of fossil fuels. The

98 J. ENTOMOL. SOC. BRIT. COLUMBIA 113, DECEMBER 2016

knowledge deficits, changing perspectives, and current emerging practices around

ecological intensification through biological control of pest diseases, IPM and pollination

are discussed.

Agriculture in the Okanagan: past, present and future

Kenna MacKenzie, Agriculture and Agri-Food Canada, Summerland, B.C., 4200

Highway #97 South, Summerland, B.C. VOH 1Z0; email: Kenna.MacKenzie@agr.gc.ca

The Okanagan is the second most important agricultural region in British Columbia.

In the late-1800s and early 1900s, agriculture began in the Okanagan with mixed

farming, in particular beef cattle and tree fruits. Over the years, various crops and

animals have been produced, such as vegetables, fruit, forage, beef and dairy, with tree

fruits becoming predominant. Changing climate, with higher winter temperatures and

longer growing seasons, has allowed a switch in crops. Today, tree fruits, particularly

sweet cherry and apples, and wine grapes, are the main agricultural crops in valley. These

trends are expected to continue in the near future.

Honeybee genetics and breeding a better honeybee

Brock Harpur and Amro Zayed, York University, Toronto, ON; email: Brock Harpur

harpur@yorku.ca; Amro Zayed zayed@yorku.ca

The genome contains the evolutionary history of a given species. The modestly sized

honey bee genome was sequenced in 2006, and since then many discoveries have been

made about the honey bee’s ancient history, its population expansions and adaptations,

and its genetic health. Here, we present the findings of several major studies from our

group that demonstrate a means through which both beekeepers and researchers can gain

valuable information about the genetic health and history of the honey bees with which

they work. First, we examine the evolution and genetic underpinnings of the honey bee

immune system and uncover valuable insights into how novel forms of social immunity

have evolved. Second, we demonstrate how, by applying genomic data such as this

within the beekeeping industry, we can make better, informed decisions about the genetic

health of our colonies.

Inventory and stewardship for pollinators in the Okanagan and Similkameen

valleys

Jennifer Heron, B.C. Ministry of Environment, Species Conservation Science Unit, Room

315, 2202 Main Mall, Vanceuver, BC Canada V6T IZ1; email:

Jennifer. Heron@gov.bc.ca

British Columbia has the highest bee diversity in Canada (at least 450 species) with

approximately one-third of the species within the Western Interior Basin in the south—

central area of the province not occurring anywhere else in Canada. Additional pollinator

groups, including butterflies and flies, are also diverse and endemic to this region.

However, this ecozone also coincides with some of the most desirable real estate in the

country, resulting in immense urban, rural and agricultural land-development pressure,

combined with threats from livestock overgrazing, wildfire suppression, natural

succession, and recreational use. Engaging land managers in stewardship actions for

pollinators is challenging, and first involves understanding the species richness and

distribution within the area of interest. To meet these objectives, a long-term project to

better understand the pollinator (primarily bee and butterfly) fauna in the Western

Interior Basin was started in 2010, initially to engage landowners, but also to contribute

to national knowledge of the bee diversity in Canada, and to try and prioritize species

that are priorities for assessment by the Committee on the Status of Endangered Wildlife

in Canada (COSEWIC). Ultimately, these data will be used to map bee species with plant

communities as a means to prioritize sites for pollinator protection. This talk highlights

some of the results from those and other invertebrate conservation work.

J. ENTOMOL. SOC. BRIT. COLUMBIA 113, DECEMBER 2016 99

Symposium abstracts:

Urban Insects — They Live Among Us

Entomological Society of British Columbia

Annual General Meeting

Pacific Forestry Centre, Victoria, BC, October 15, 2016

Out of the cities and into the forest: Range expansions of non-indigenous

introductions in southwest British Columbia

Lee M. Humble, Natural Resources Canada, Canadian Forest Service, Victoria,

B.C., Canada

Since surveillance programs were established for the detection of non-indigenous

introductions in the mid 1990s, more than 25 introduced species of Coleoptera,

Hemiptera, Hymenoptera, and Lepidoptera have been discovered in the urban and

managed forests of British Columbia. The life histories and known distributions of

selected species are documented and used to infer likely pathways of introduction and to

illustrate the importance of anthropogenic influences on their range expansion. Future

research needs for species of potential importance to forestry in B.C. are briefly

discussed.

Preventing gypsy moth from establishing in British Columbia isn't fun

Tim Ebata, Resource Practices Branch, B.C. Ministry of Forests, Lands and Natural

Resource Operations, Victoria, B.C., Canada

My talk outlines how British Columbia has successfully remained "gypsy moth free"

and describes some of the difficulties faced in mounting eradication programs in an

urban environment.

Treading Carefully on Fire Ants in the Urban Landscape

Rob Higgins, Biological Sciences, Thompson Rivers University, Kamloops, B.C.,

Canada

Working quietly on ants while sitting in a forest, perhaps a hundred metres from a

colleague and a hundred kilometres from the nearest town allows you to develop a

specific set of research and social skills. Unfortunately, none of them provide much

guidance when you are talking to an angry homeowner who has recently retained a

lawyer because of the ants you are studying. Nor guidance when dealing with businesses

fearing major losses who need immediate advice you simply aren’t sure you have. Nor

dealing with the police and fire department that someone has called. Nor needing to keep

your data so confidential you cannot share it with your funding agency and certainly not

the media who keep asking. Working in a social environment as densely developed as the

condominiums you find yourself within is a uniquely challenging situation. Here we will

look at the invasive fire ants of BC in the urban landscape and reflect on those times we

spent sitting in that quiet forest while stuck in traffic.

180,000 bites later: The aggregation pheromone of the common bed bug is finally

identified.

Regine Gries', Robert Britton’, Michael Holmes? and Gerhard Gries', ‘Department

of Biological Sciences; *Chemistry Department, Simon Fraser University, Burnaby,

B.C., Canada

Drawing on our 2015 publication in Angewandte Chemie (International Edition), the

presentation describes our approach to collecting sufficient pheromone sources for

identifying the aggregation pheromone of the common bed bug (Cimex /ectularius;

100 a, ENTOMOL. Soc. BRIT. COLUMBIA 113, DECEMBER 2016

Hemiptera: Cimicidae), the analytical steps taken to identify the pheromone blend, the

pheromone components that mediate attraction and arrestment of bed bugs, and the

experiments we have run in the laboratory and in bed bug-infested apartments to test the

effect of synthetic pheromone as a trap lure. The presentation also highlights future

objectives, including the development of a commercial lure and trap.

Butterfly and moth conservation in urban and semi-urban habitats: Challenges and

reflections taken from species at risk recovery projects

Jennifer Heron, British Columbia Ministry of Environment, Vancouver, B.C., Canada

The order Lepidoptera is one of the largest and most studied orders of insects. This

group is ecologically and economically important, serving as pollinators of many plants

and pests for many others. Butterflies are considered by many to be the most charismatic

of the arthropods, and the public enjoys seeing these species in their gardens and

surrounding natural environments. Many species of Lepidoptera, especially pollinating

groups, are at risk. Although the butterflies are relatively well known, there are many

species of moths we know little about, and engaging the public in moth conservation

efforts is challenging. In this talk, we summarize the biology of these species and cite

examples of Lepidoptera conservation projects and the challenges encountered in urban

and semi-urban areas.

Beetles in the City: Carabid diversity in the urban environment

Rob McGregor and Veronica Wahl, Institute of Urban Ecology, Douglas College,

New Westminster, B.C., Canada

Ground beetle surveys (Coleoptera: Carabidae) have been widely used to assess

habitat quality and the influence of human disturbance on urban, agricultural, and

forested landscapes. Here, we describe carabid surveys conducted in urban forest

fragments in Coquitlam, B.C., where European carabid species predominate in disturbed

forests. In addition, we describe a citizen science program where homeowners and

community gardeners trap and identify carabids from urban gardens, in association with

insectary plants. Finally, we describe prelimimary work to document populations of a

threatened tiger beetle, Omus audouini, in coastal habitat in Delta, B.C.

Up on the Roof: Surveying Biodiversity in a Unique Urban Landscape

C.G. Ratzlaff and K.M. Needham, Beaty Biodiversity Museum, University of British

Columbia, Vancouver, B.C., Canada

We have been conducting a monthly survey of the insects making their home on the

"green roof" atop the Vancouver Convention Centre, West. The roof was planted almost a

decade ago with 23 species of plants and is nearly sox acres in size—the largest green

roof in Canada. It is left to grow throughout the year and is mowed only once in the fall.

Surrounded by tall buildings and concrete sidewalks, with no significant green spaces

nearby, we were curious about which insects might find this "meadow" a suitable habitat.

Beginning in April and ending in December, we will have visited the roof once a month,

including one black-light trapping event in September, and will have catalogued all of the

insects collected there. In this talk, we present our preliminary results and highlight some

of the finds we have made to date.

J. ENTOMOL. SOC. BRIT. COLUMBIA 113, DECEMBER 2016 101

Presentation Abstracts

Entomological Society of British Columbia

Annual General Meeting

Pacific Forestry Centre, Victoria, B.C., October 14, 2016

The uninvited dinner guests... and how to get rid of them

Joshua Pol, Regine Gries and Gerhard Gries, Department of Biological Sciences, Simon

Fraser University, Burnaby, B.C., Canada

German cockroaches are pests in human dwellings. We hypothesized that German

cockroaches are strongly attracted to human foods. In laboratory experiments, we

bioassayed the responses of the insects to many food types. In a field experiment, one

particularly attractive food type proved as appealing to the cockroaches as the leading

commercial bait.

Female black widow spiders respond to semiochemicals from conspecific females

Andreas Fischer', Manfred Ayasse*, and Maydianne Andrade’, 'Department of

Biological Sciences, Simon Fraser University, Burnaby, B.C., Canada; Institute of

Evolutionary Ecology and Conservation Genomics University of Ulm, Germany;

3Integrative Behaviour & Neuroscience University of Toronto Scarborough, Canada

Females of many web-building spiders produce long-range semiochemicals (message-

bearing chemicals). Males respond to them in their search for females, distinguishing

between prospective mates that differ in feeding status (well-fed or starved) and sexual

maturity (juvenile or adult).

Female spiders whose webs were destroyed often seek the presence of other females

when they rebuild their webs. We tested the hypothesis that females respond to

semiochemicals from other females when they select new web-building sites. We ran Y-

tube olfactometer experiments with adult virgin females of the Western black widow

spider (Latrodectus hesperus) and the redback spider (L. hasselti), offering them a choice

between a blank control stimulus and a treatment stimulus consisting of a conspecific

female. We found that L. hesperus females avoid sub-adult and adult conspecific females,

whereas L. hasselti females avoid only adult conspecific females. Females of both

species preferred starved to well-fed females. This is the first evidence for

semiochemical-guided decision-making by female spiders and for semiochemicals

produced by sub-adult females. Future work will focus on the identification of the

semiochemicals that mediate the avoidance of well-fed females.

Communication between yellowjacket wasps and symbiotic yeast

Tamara Babcock, Department of Biological Sciences, Simon Fraser University, Burnaby,

B.C., Canada

Recent studies suggest that yellowjackets may share a mutualistic relationship with

fermentative yeast, but little is known about how these organisms find each other. Our

research demonstrates that yeast from the digestive tract of yellowjackets produces

attractive volatiles when grown on grape juice-infused agar.

Mosquitoes: nectar thieves or pollinators?

Dan Peach and Gerhard Gries, Department of Biological Sciences, Simon Fraser

University, Burnaby, B.C., Canada

Outside of interactions with humans and pathogens, many aspects of mosquito ecology

have severe knowledge gaps. While feeding on floral nectar is important to mosquitoes,

they are traditionally thought to not pollinate the flowers they visit. Contradicting this,

we report evidence that mosquitoes pollinate common tansy and yarrow.

102 J. ENTOMOL. SOc. BRIT. COLUMBIA 113, DECEMBER 2016

A song of ants and fire: improving baiting methods for the European fire ant

Danielle Hoefele, Department of Biological Sciences, Simon Fraser University, Burnaby,

B.C., Canada

The invasive European fire ant (Myrmica rubra) defends nests aggressively, rendering

gardens, lawns and parks unusable. I tested various foods for foraging activity by

European fire ants to determine whether they prefer specific carbohydrates and proteins.

These results will be used in the future to develop a more effective insecticidal bait.

Evaluation of two passive horizontal transmission pathways for Metarhizium

brunneum in Agriotes obscurus click beetles

J. P. S. Leung, J. S. Cory, J. T Kabaluk, and A. F: Janmaat, Department of Biological

Sciences, Simon Fraser University, Burnaby, B.C., Canada

Fungal pathogens are unique among entomopathogens in that ingestion is not required

for transmission. They can be passively transferred to conspecifics through direct contact

or through contact with contaminated substrates. We discuss the relative importance of

these two pathways for Metarhizium brunneum in the control of Agriotes obscurus click

beetles.

Filling in the gaps of the IMD immune pathway of the kissing bug Rhodnius

prolixus

Nicolas Salcedo and Carl Lowenberger, Department of Biological Sciences, Simon

Fraser University

Rhodnius prolixus is a hemathophagous hemipteran vector of the parasite Trypanosoma

cruzi. Similar to other hemimetabolous insects, the genome of R. prolixus had no key

components of the highly conserved IMD pathway. However, IMD-related effector

immune genes are normally expressed. Using bioinformatics, I propose candidate genes

completing the IMD pathway.

Population dynamics of the western tent caterpillar: the roles of fecundity, disease,

and temperature

Paul MacDonald, Department of Biological Sciences, Simon Fraser University, Burnaby,

B.C., Canada

Many populations of forest Lepidoptera exhibit regular periodic cycling in abundance,

but mechanisms for such dynamics remain a subject of debate in ecology. I used annual

field data (1977-2015) from a cyclical species, the western tent caterpillar (Malacosoma

californicum pluviale), to elucidate how fecundity, viral disease, and temperature

contribute to the cyclical dynamics of five field populations.

A selfish X chromosome in a mushroom-feeding Drosophila

Graeme Keais and Steve Perlman, Department of Biology, University of Victoria,

Victoria, B.C., Canada

Selfish genetic elements are widespread and powerful forces in evolution. By increasing

their transmission relative to the rest of the genome for each generation, they spread

rapidly through populations, even if they carry a negative fitness cost. Driving X

chromosomes are selfish genetic elements that kill Y-bearing sperm in a number of

Dipterans. Because males that carry a driving X transmit almost exclusively X-bearing

gametes, they produce predominantly female offspring. We provide the first evidence for

a driving X chromosome in a common European mushroom-feeding Drosophila species.

Males carrying the driving X sire between 80—-100% female offspring, and most of their

sons (of which there are few) are sterile and appear to lack a Y chromosome. Sperm

bundles in driving X males develop abnormally, indicating that the driving X is acting

during male gametogenesis.

J. ENTOMOL. Soc. BRIT. COLUMBIA 113, DECEMBER 2016 103

Photosensitivity in developing mountain pine beetle (Dendroctonus ponderosae)

Debra Wertman, Department of Biology, University of Victoria, Victoria, B.C., Canada

This research explores the capacity for photosensitivity in developing mountain pine

beetles (Dendroctonus ponderosae). The identification of a long-wavelength opsin and

negative phototaxis in eyeless beetle larvae, as well as an effect of photoperiod on adult

emergence, suggest that light may function in survival and life-cycle coordination in this

species.

Earwigs (Forficula auricularia) as a biocontrol agent: deciphering a generalist

predator

Dennis Quach, Jenny Cory, Tamara Richardson and Margo Moore, Department of

Biological Sciences, Simon Fraser University, Burnaby, B.C., Canada

The European Earwig (Forficula auricularia) has garnered scientific interest and public

distrust due to its controversial status as both a beneficial predator and an urban pest. We

investigate the potential for its use as a biocontrol agent through dietary gut-content

analysis and the effects of temperature on its predation efficacy.

The role of pathogen diversity on the evolution of resistance in an insect

Leon Li, Department of Biological Sciences, Simon Fraser University, Burnaby, B.C.,

Canada

Our objective is to determine whether baculovirus diversity affects the rate and

magnitude at which resistance evolves. Using Trichoplusia ni as a host, changes in

resistance against single versus mixtures of ACMNPV were examined. We found

evidence of reduced resistance, as well as increased life-history costs in diverse

infections.

Response of epigaeic arthropods to riparian habitat enhancement trials in

Kinbasket Reservoir, British Columbia

Charlene M. Wood and Virgil C. Hawkes, LGL Limited Environmental Research

Associates

We monitored the response of ground-dwelling spider and beetle assemblages (over 200

species) to habitat enhancement trials in Kinbasket Reservoir, British Columbia, from

2014-2015. Species differed one year post-treatment, with treatment assemblages

initially dominated by bare-ground associated species. Monitoring is ongoing to evaluate

the turnover in species assemblages as vegetation establishes over time.

Role of toxins in insect defensive symbiosis

Steve Perlman, Finn Hamilton and Matt Ballinger, Department of Biology, University of

Victoria, Victoria, B.C., Canada

Insects are commonly infected with bacterial endosymbionts that are transmitted

primarily from mothers to their offspring, often in the egg cytoplasm. These inherited

symbionts play important roles in the ecology and evolution of their hosts, such as

protecting them against a wide range of natural enemies, including predators, parasites,

and pathogens. Little is known about the mechanism of protection by symbionts. Is there

specificity? How do defensive symbionts target enemies without harming their host? We

studied protection in a defensive symbiosis between the common mushroom-feeding fly,

Drosophila neotestacea, and its bacterial endosymbiont, Spiroplasma, which protects its

host against parasitic nematodes and wasps. We found that Spiroplasma encodes

ribosome-inactivating proteins, related to Shiga toxins, and that nematode ribosomes

show a strong signal of toxin-mediated attack. It is likely that symbiont-encoded toxins

are common and versatile tools in defensive symbiosis.

104 J. ENTOMOL. SOC. BRIT. COLUMBIA 113, DECEMBER 2016

3D images = public understanding

Scott Montague, Blackhole Collections

Latin words alienate the general public from engaging with the most accessible and

fascinating animals on the planet. Learn how new technology effortlessly bridges the

vocabulary gap that used to stop readers in their tracks. Public support is a factor for

government funding; now, the layperson can understand and agree.

J. ENTOMOL. Soc. BRIT. COLUMBIA 113, DECEMBER 2016 105

OBITUARY

Thelma Finlayson

(29 June 1914 - 15 September 2016)

Thelma Finlayson passed away September 15, 2016. She was 102 years old, and a

beloved teacher, mentor, and colleague. Born on 29 June, 1914, Thelma grew up in

Trenton, Ontario. She earned her B.A. (Honours, Biology) from the University of

Toronto in 1936, a certification in Taxonomy and Biological Control from ARPE in 1971,

and a Doctor of Laws (Honoris causa) from Simon Fraser University in 1996. She was

named to the Order of Canada in 2005 as “a trailblazing entomologist and a beloved

teacher and advisor.” Thelma was a past president of the Entomological Society of BC

and an Honorary Member and a Fellow of the Entomological Society of Canada. She was

a Life Member of the Entomological Society of British Columbia and a Fellow of the

Entomological Society of Ontario. Two insect species, Anisota finlaysoni Ruiotte

(Lepidoptera: Saturniidae) and Mesopolobus finlaysoni Dogenlar (Hymenoptera:

Pteromalidae), have been named in her honour.

Thelma had taken an entomology course from E. M. Walker and began her

professional career in 1937 in the Dominion Parasite Laboratory in Belleville, Ontario,

eight miles from her home, as a Technical Officer. She obtained her position by patiently

sitting in the office of the laboratory until someone needed an extra pair of hands. Even

in her early career, she was an implacable force; her first project was to rear “millions of

sawflies” (her words) in the newly built quarantine facility, searching for parasites to

control European spruce sawfly that were decimating Quebec and New Brunswick

forests. This led to her lasting interest in the taxonomy of parasitic larvae. She was, in

fact, one of the first women scientists in the federal research service, but in 1946, as a

married woman, she was asked to resign because men were returning from the war.

However, at that time, her husband Roy had become very ill and would soon be

compelled to retire, making it necessary for Thelma to work. She challenged the request

to resign by threatening to ensure that every other married woman in the Civil Service

would be fired for the same reason. In due course, the Assistant Deputy Minister of

Agriculture verified her right to work, saving her own job and establishing an important

human rights precedent for the Federal Civil Service. In 1959, she was promoted to

Research Officer (levels 1, then 2) and later Research Scientist.

In 1967, Thelma joined seven other scientists, led by Bryan Beirne, who left the

Belleville Research Institute for Biological Control to expand the Department of

Biological Sciences at the newly established Simon Fraser University, in Burnaby,

British Columbia. There, Thelma was appointed Assistant Professor and Curator of

Entomology. She helped found SFU’s Pestology Centre, later renamed the Centre for

Pest Management, one of the first of its kind. The Department showed its regard for her

teaching and research by promoting her to Full Professor in 1976, despite her having

only an Honours-level B.Sc.—an unheard-of advancement in the Faculty of Science. She

held the position of Professor Emerita from 1979 and was the University’s first Emerita.

While at SFU, she officially mentored seven Masters and Ph.D. candidates, and

unofficially, countless others. Graduate and undergraduate students sought Thelma’s

advice for decades. In addition to the time she devoted to students, Thelma was an

advocate of entomology and education through significant financial contributions toward

Pest Management fellowships. Furthermore, she financed an endowment to establish the

Finlayson Chair in Biological Control in the Department of Biological Sciences,

currently held by Jenny Cory.

106 J. ENTOMOL. SOC. BRIT. COLUMBIA 113, DECEMBER 2016

Thelma taught courses in introductory biology and, of course, insect biology. The

insect biology course was encyclopaedic, and students typically left each lecture with 30

to 40 pages of detailed notes and sketches. This course inspired many biology students to

careers in entomology.

In 1971, Thelma was persuaded to be the first advisor for science in the newly formed

Academic Advice Centre and, in 1983, she was appointed as Special Advisor, mostly

advising students in academic difficulty and retaining that position until 2012. During

this time, she advised more than 7,000 students. At age 97, she may well have been the

oldest student advisor on the planet. In 2012, the centre was renamed the Thelma

Finlayson Centre for Student Engagement.

Thelma’s primary interest was in the taxonomy of natural enemies with application to

Biological Control. She published approximately 40 papers, memoirs, and book chapters

during her career. She was particularly successful in her work on the taxonomy of

immature Hymenoptera, which in many ways foreshadowed the use of DNA to identify

remains of parasitoids within hosts. She was also consulted on entomological questions

by the RCMP, which led to the development and establishment of the internationally

recognized Forensic Entomology Laboratory in SFU’s Department of Criminology by

her faithful friend, Dr Gail Anderson.

In the first lecture of every course Thelma taught, she told students “My door is

always open for you.” Those who had the privilege of being her students and of being

mentored and supported by this singular woman found that statement to be true always.

Written by

BERNIE ROITBERG, DAVE GILLESPIE, and PETER BELTON

J. ENTOMOL. SOC. BRIT. COLUMBIA 113, DECEMBER 2016 107

Correction:

Gelling, Lea. 2015. Habitat associations of adult Oregon Branded Skipper at Cordova

Shore, Vancouver Island, British Columbia. Journal of the Entomological Society of

British Columbia 112:57-68.

In Vol. 112 (2015) the species name, Grindelia oregonia (Oregon gumweed), is

incorrect in the Abstract and Table 3; the correct species is Grindelia stricta.d

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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.entsocbc.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.

Review and forum articles — Please submit ideas for review or forum articles for consideration to the editor

at journal@entsocbe.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. Print back issues of many volumes of

the journal are available at $15 each.

Membership in the Society is open to anyone with an interest in entomology. Dues are $20 per year 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://blogs.sfu.ca/groups/esbc/membership/.

3 9088 01976 7821

Journal of the

Entomological Society of British Columbia

Volume 113 December 2016 ISSN#0071-0733

Directors of the Entomological Society of British Columbia 2016-2017......... Shak tie eer ageit 2

Development and oviposition preference of Xenotemna pallorana Robinson (Lepidoptera:

Tortricidac) on. alfalfa.and fruit tree foliagesse..sic cist ue ede ae ee ee 3

Efficacy of SPLAT® Verb for protecting individual Pinus contorta, Pinus ponderosa, and

Pinus lambertiana from mortality attributed to Dendroctonus ponderosae

wi sab rue ale winsnitre vals ie oeiRiwi ap biars oe cdl abd oe Pep a ee Te 11

History of the balsam woolly adelgid, Adelges piceae (Ratzeburg), in British Columbia,

with notes.on @ recent rahge expansion. jie sds Gs vueasse ear eaters emote eee a Zt

Pollen preferences of two species of Andrena in British Columbia’s oak-savannah

COSY BUSI odie ce discs ecaceranlin walb'y's a 410 aualptealy UOiSaNar ats glo AE i aie ella tt meee ee 39

Relative efficacies of sticky yellow rectangles against three Rhagoletis fly species (Diptera:

Tephritidae) in Washington State and possible role of adhesives ..................cc0ceee eee ees 49

SCIENTIFIC NOTES

Pacific Flatheaded Borer, Chrysobothris mali Horn (Coleoptera: Buprestidae), found

attacking apple saplings in the Southern Interior of British Columbia ........................ a

NATURAL HISTORY AND OBSERVATIONS

First Canadian records for two invasive seed-feeding bugs, Arocatus melanocephalus

(Fabricius, 1798) and Raglius alboacuminatus (Goeze, 1778), and a range extension for a

third species, Rhyparochromus vulgaris(Schilling, 1829) (Hemiptera: Heteroptera)........ 74

Notes on insects recently introduced to Metro Vancouver and other newly recorded species

+ from rites Cola se oia:susdus'sie.sapdoina od darsscadies pose ree ee 79

Rhyparochromus vulgaris (Schilling) (Hemiptera: Heteroptera: Rhyparochromidae): newly

discovered in the interiorot British Colwinbia: 3.55.2. 435.00 gaan emia meres oa, 90

SYMPOSIUM ABSTRACTS

Proceedings of the Pollination Science and Stewardship Symposium.....................008. 92

Symposium abstracts: Urban Insects — They Live Among US ...............ceesceeeee cence 99

Presentation Abstracts: ESIC Ag. 0.55 sinc van siomadse eves pa uae nena eee tech gsi: 101

OBITUARY: Thelma Finlayson (29 June 1914 - 15 September 2016) .............0..000. 105

BRAT A isa sesci o's orev te nln bvie Moved saya bg nig gee a Onli 11 ge rater eR TTS hi 107

NOTICE TO CONT RIBU PORS ori lie did iods sl ou ae ee Inside Back Cover