J. ENTOMOL. SOC. BRIT. COLUMBIA 102, DECEMBER 2005 1

Journal of the

Entomological Society of British Columbia

Volume 102 Issued December 2005 ISSN #0071-0733

Directors of the Entomological Society of British Columbia, 2005-2006... 2

Naomi C. DeLury, Gary J.R. Judd and Mark G.T. Gardiner. Antennal detection of sex

pheromone by female Pandemis limitata (Robinson) (Lepidoptera: Tortricidae) and its

impact ontthein calling DEM aVIOUE. --.c.., cash ueost.n 0p vecdysqceeasactann cease aeuniad mapeereccayamet 3-12

Michael K. Bomford and Robert S. Vernon. Moisture tempers impairment of adult

Otiorhynchus sulcatus (Coleoptera: Curculionidae) climbing ability by fluoropolymer,

Pale dust, amd Tithiuim Crease oe csr ieee eterenasks eed cctsez no's eee aee es seaneeeene ines 13-20

Lawrence C. Wright, Wyatt W. Cone and David G. James. Sources of Spring and Fall

Hop Aphid, Phorodon humuli (Schrank), (Homoptera: Aphididae) Migrants in South

Cynthia L. Broberg and John H. Borden. Host preference by Saperda calcarata Say

(Coleoptera: C crab yC1dac) 2 ric. oes nce cord ska usadi ome. la tleglasenadnuen taco eosin ss ve tee eae ees 27-34

Peter J. Landolt, Alberto Pantoja and Daryl Green. Yellowjacket Wasps (Hymenoptera:

Vespidae) Trapped in Alaska with Heptyl Butyrate, Acetic Acid and Isobutanol 35-42

Rex D. Kenner. Redescription of Haliplus dorsomaculatus (Coleoptera: Haliplidae) with a

New Synonymy and Comments on Habitat and Distribution ............00.ccecceeee 43-56

Robert A. Cannings and John P. Simaika. Lestes disjunctus and L. forcipatus (Odonata:

Lestidae): An evaluation of status and distribution in British Columbia ............. 57-64

Glenn E. Haas, James R. Kucera, Amy M. Runck, Stephen O. MacDonald and Joseph A.

Cook. Mammal Fleas (Siphonaptera: Ceratophyllidae) New for Alaska and the South-

eastern Mainland Collected During Seven Years of a Field Survey of Small Mammals

See SABE Rect Rac Map crac teroee rinse Waar sstenieGe ei rede irae seca helene (hana heoeme nner pane eesiee 65-76

SCIENTIFIC NOTES

Sujaya Rao and Stephen C. Alderman. Infestation of Bent Grass by a New Seed Pest,

Chirothrips manicatus (Thysanoptera: Thripidae), in OregoN..............::ceeeeeeeeeees 77-78

Lawrence A. Lacey, Steven P. Arthurs and Heather Headrick. Comparative Activity of the

Codling Moth Granulovirus Against Grapholita molesta and Cydia pomonella

(Repidopterds i Ont Gide) 27 cen cere insets cna sesnceassneaeeun’ stent fas saaeonstaeebhasanaeeeses 79-80

Leland M. Humble. A novel host association for Monarthrum scutellare (Coleoptera:

Curculionidae: Scolytinae) in British Columbia.................cccccecescscssssceesseessseeeeees 81-82

Claudia R. Copley and Robert A. Cannings. Notes on the status of the Eurasian moths

Noctua pronuba and Noctua comes (Lepidoptera: Noctuidae) on Vancouver Island,

|e 1 ihci els C0) LD a0] otc ee nena mee cet ere nate Ota Eee DRGa URI cary Sannin aS amr enESt Te MOON ne Rt) 83-84

J. ENTOMOL. SOC. BRIT. COLUMBIA 102, DECEMBER 2005

DIRECTORS OF THE ENTOMOLOGICAL SOCIETY

OF BRITISH COLUMBIA FOR 2005-6

President:

Karen Needham

Spencer Entomological Museum

President-Elect:

Richard Ring

University of Victoria

Past-President:

Dave Raworth

AAFC-PARC Agassiz

Secretary/Treasurer:

Robb Bennett

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

Directors, first term:

Markus Clodius, Rob McGregor

Director, second term:

Jen Perry, Hugh Philip, Niki Hobischak

Regional Director of National Society:

Allan Carroll

Canadian Forest Service, Victoria

Editorial Committee, Journal:

Editor-in-Chief: Subject Editors:

Ward Strong Joan Cossentine (Agriculture)

BC Ministry of Forests and Range Lorraine MacLauchlan (Forestry)

ward.strong@gov.bc.ca Robb Bennett (Systematics/Morphology)

Co-Editors, Boreus:

Jennifer Heron, Suzie Lavallee

Jennifer. Heron@gov.bc.ca

Editor of Web Site:

Bill Riel

ESBC@harbour.com

Honourary Auditor:

Dave Raworth

AAFC-PARC Agassiz

Web Page: http://esbc.harbour.com/

J. ENTOMOL. SOC. BRIT. COLUMBIA 102, DECEMBER 2005 3

Antennal detection of sex pheromone by female

Pandemis limitata (Robinson) (Lepidoptera: Tortricidae)

and its impact on their calling behaviour

NAOMI C. DELURY’”, GARY J.R. JUDD!

and MARK G.T. GARDINER!

ABSTRACT

Previous observations lead us to believe that female Pandemis limitata (Robinson) (0 to

24 h old) are as attractive as their pheromone gland extract to males in clean air, but are

more attractive in an environment permeated with their major pheromone component

(Z)-11-tetradecenyl acetate. Therefore, in this study, we tested the hypothesis that fe-

males can detect and/or respond to their pheromone components. Using electroanten-

nographic detection, we found female P. /imitata able to perceive both of their known

pheromone components, (Z)-11-tetradecenyl acetate and (Z)-9-tetradecenyl acetate. Fe-

male antennal response was found to be 46.3% weaker than that of males, under identi-

cal conditions, with male antennae producing significantly higher deflections to the

higher pheromone doses tested and to the plant volatile, (£)-2-hexanal. Observations of

females in clean air versus (Z)-11-tetradecenyl acetate-permeated air showed no signifi-

cant differences with respect to onset time, frequency or duration of calling. Females

moved significantly less often in a (Z)-11-tetradecenyl acetate-permeated portion of a

flight tunnel than in the corresponding clean-air portion.

Key Words: (Z)-11-tetradeceny] acetate, (Z)-9-tetradecenyl acetate, female electroan-

tennography, flight tunnel, mating disruption, threelined leafroller, sprayable phero-

mone, microencapsulated pheromone, movement

INTRODUCTION

Although pheromone-based mating dis-

ruption of Lepidopteran species is widely

employed in some agricultural systems

(Thomson et al. 2001), questions about the

behaviour of female moths in the presence

of their own sex pheromone often remain

unanswered. In particular, while there is

evidence that some female moths perceive

(Michell et al 1972, Birch 1977,

Palaniswamy and Seabrook 1978, Light and

Birch 1979, Barnes et al. 1992) and even

modify their behaviour (Palaniswamy and

Seabrook 1978, 1985, Palaniswamy ef al.

1979, Sanders 1987, Weissling and Knight

1996, Evenden 1998) in response to their

own sex pheromone, others apparently do

not (El-Sayed and Suckling 2005) and in-

formation of this type is lacking for most

species where commercial use of mating

disruption is under study. During our own

studies of pheromone communication dis-

ruption in the threelined leafroller, Pan-

demis limitata (Robinson) (Lepidoptera:

Tortricidae), it appeared that males were

more responsive to “calling” females (0 to

24 h old) in a pheromone-permeated atmos-

phere than they were to female gland ex-

tracts, although no difference in male re-

sponse was detectable between these

sources in clean air (N.C.D., unpublished

data). One explanation for this observed

difference is that female P. /imitata can

detect their pheromone environment and

adjust their behaviour in a manner that

’ Agriculture and Agri-Food Canada, Pacific Agri-Food Research Centre, 4200 Hwy 97, Summerland, BC,

VOH 1Z0, Canada

* To whom correspondence should be addressed: Tel.: +1 250 494 7711; Fax: + 1 250 494 0755; E-mail:

deluryn@agr.gc.ca

4 J. ENTOMOL. SOC. BRIT. COLUMBIA 102, DECEMBER 2005

makes them more detectable to males than

the female gland extract alone. Using elec-

troantennograms (EAGs), we assessed the

ability of female P. limitata to perceive

their two known sex pheromone compo-

nents, (Z)-11-tetradecenyl acetate [Z11-

14:Ac] and (Z)-9-tetradeceny] acetate [Z9-

14:Ac] (Roelofs et al. 1976) and asked the

question, do females alter their calling be-

haviour in the presence of microencapsu-

lated (MEC) Z11-14:Ac applied in labora-

tory flight-tunnel assays (Judd et al. 2005)?

MATERIALS AND METHODS

Laboratory cultures. Experiments

were conducted with P. limitata from a

laboratory colony maintained at the Pacific

Agri-Food Research Centre, Summerland,

British Columbia (BC) since 1992 that

originated from wild populations collected

in the Similkameen Valley of BC. Pan-

demis limitata were maintained on a modi-

fied pinto bean-based diet (Shorey and Hale

1965) at 25 °C under a 16:8 h L:D photo-

regime. Pupae were removed from diet,

sexed and placed individually in 150 ml

plastic cups provided with a wet cotton

wick until adults eclosed. Male and female

moths were isolated from each other in

separate environmental chambers (25 °C,

65% r.h. with a 16:8 h L:D reversed photo-

regime).

Female and male perception of syn-

thetic pheromone. Perception of synthetic

pheromone by female and male P. /imitata

was measured using EAGs. Our EAG sys-

tem consisted of an IDAC-02 computer-

coupled data acquisition board, an INR-02

EAG-SSR system and AutoSpike software

(Syntech, Hilversum, The Netherlands).

Antennae excised from 0 to 24 h old fe-

males or males were mounted individually

inside 10 ul glass capillaries containing

silver-coated wire recording electrodes.

Glass tubes were pulled at 300 °C on a Nar-

ishige (Model PN-30, Tokyo, Japan) micro-

pipette puller. The distal antennal segment

was removed and that end of the remaining

antenna inserted into the indifferent elec-

trode capillary. As a result, each antenna

was suspended between two glass capillary

electrodes filled with insect Ringer’s solu-

tion (DeLury et al. 1999). Each antenna

was challenged with six doses, in ten-fold

increments from 0.1 ng to 10 pg, of Z11-

14:Ac (97.5% purity, Sigma Chemical

Company, St. Louis, MO), Z9-14:Ac (97%

purity, Regine Gries, Simon Fraser Univer-

sity, BC), and a 94:6 ratio blend of both,

respectively. The amount of Z11-14:Ac

remained equal between the treatments of

the main component and the blend; female

antennae were also exposed to 100 ug doses

from both Z11-14:Ac and the blend. Each

antenna was exposed to each dose of each

of the three compounds in order of increas-

ing concentration. Treatments were puffed

(200 ms; 10 ml per s) over each antenna,

beginning with the lowest dose (0.1 ng) of

each of the three compounds, which were

presented in random order at each incre-

mental dose. Baseline antennal responses

were established using HPLC-grade hexane

(two puffs) and a puff of the plant volatile,

(£)-2-hexanal in paraffin oil, before and

after each pheromone test concentration.

All stimulus puffs were applied at 30 s in-

tervals. All chemical stimuli were dis-

pensed in 10 ul aliquots onto folded

Whatman #5 filter paper (3 cm x 2 cm),

placed in pipette tubes and connected to the

puffer. The procedure was repeated using

six female antennae and four male anten-

nae.

Normalized percentage antennal deflec-

tions were calculated using the plant vola-

tile response as the standard. Data were

analyzed by one-way analysis of variance

(ANOVA) followed by Dunnett’s test to

compare each treatment mean to the hexane

control. All experimental error rates were

set at a = 0.05. Comparisons between fe-

male and male antennal responses at each

dose were conducted on individual antennal

deflections (mV) by first performing a two-

way ANOVA, where the factors were sex

and pheromone stimulus. A sex x phero-

mone stimulus interaction term was in-

J. ENTOMOL. Soc. BRIT. COLUMBIA 102, DECEMBER 2005

cluded in the model. Following a significant

ANOVA, linear contrasts using f-ratio sta-

tistics were used to make paired compari-

sons of mean antennal deflections for males

and females at each dose. Experiment-wise

error rates for these multiple-paired com-

parisons were fixed at 5% by adjusting a

values for individual comparisons using the

Bonferroni inequality. All statistical tests

were performed with JMP 5.1 (2003).

Female behaviour in a pheromone-

permeated environment. Female moths

were placed in clean air or a pheromone-

permeated environment and observed in

side-by-side comparisons. To accomplish

this, a sheet metal divider (73.3 cm x 37.4

cm) was installed vertically in the middle of

the upwind portion of a pulling-style flight

tunnel [75 cm wide =< 73.3 cm high =< 187

cm long flight section] covered with a re-

placeable 4 ml transparent polyester

‘skin’ (3M Canada Company, London, ON)

(Judd et al. 2005), creating two identical

isolated chambers (upwind area = 36.65

cm’). The upwind cross-sectional surface

was constructed of a set of replaceable hori-

zontal sheet metal panes (73.3 cm x 3.7 cm

with a 0.7 cm bend for stability) onto which

pheromone was applied, with an untreated

G200 Filtrete® with 0.5 oz Coverweb® (3M

Canada Company, London, ON) _ that

smoothed airflow, sealing the tunnel ca. 20

cm upwind of the panes. Smoke tests con-

firmed that air was drawn directly into each

chamber at the upwind end and immedi-

ately moved downwind in a linear fashion

through the tunnel. After observing females

in both chambers in clean air, one chamber

was randomly chosen to receive pheromone

(10 mg ai - m”), which was applied to hori-

zontal metal panes forming the upwind end

of the tunnel. As a precaution, pheromone

was not applied to a 1.5 cm strip immedi-

ately adjacent to the untreated chamber,

creating a buffer to ensure no pheromone

entered the clean-air portion of the tunnel.

Untreated panes were shielded during all

pheromone applications and the tunnel was

allowed to ventilate for a minimum of 18 h

between replicates. Pheromone treatment

consisted of Phase I MEC Z11-14:Ac (3M

Canada Company, London, ON) applied at

a rate equivalent to 10 mg ai - m~ to the

cross-sectional area of the tunnel as de-

scribed in detail by Judd et al. (2005).

Virgin female moths (0 to 24 h old)

were chilled for 5 min at 2 °C and trans-

ferred individually into stainless steel mesh

observation chambers (4.5 cm W x 3.5 cm

H x 5 cm D). Chambers were stacked verti-

cally to form a series of five individually-

caged females. Treated and control portions

of the tunnel each received one series of

five individually-caged females ca. 2.5 h

before scotophase, for a total of 15 females

in each treatment. Females were observed

for calling (raised wings with protruding

abdomen) and movement (physically walk-

ing in the chamber) every 15 m until the

initiation of scotophase, when they were

observed continually for 3 h. Data gathered

before scotophase was used solely for de-

termination of onset of calling behaviour

and was not included in any other analysis.

Frequencies, or the number of discrete

observations, of calling and movement

were analyzed separately using a two-factor

ANOVA where treatment (Z11-14:Ac- and

clean-air) and vertical position of female

were the factors (JMP 5.1 2003). Onset and

duration of calling were analyzed using a

two-sided Wilcoxon Signed Ranks Test

(JMP 5.1 2003), as the assumption of nor-

mality was not met for these data. The tun-

nel was assessed for positional bias be-

tween the two sides with respect to each

behaviour before the application of phero-

mone and resulting data were analyzed as

above.

RESULTS

Female and male detection of syn-

thetic pheromone. EAGs confirmed that

adult female P. limitata can perceive both

components of their sex pheromone (F> 274

= 19.40; P < 0.0001). However, antennal

deflections significantly differed from those

of the hexane control only for doses of 10

ug or greater for Z11-14:Ac and the 94:6

6 J. ENTOMOL. SOc. BRIT. COLUMBIA 102, DECEMBER 2005

ratio of Z11-14:Ac: Z9-14:Ac (Fig. 1A).

Response to Z9-14:Ac differed from the

hexane control at each of the extreme doses

tested, 0.1 ng and 10 ug, but did not differ

from any of the doses in between (Fig. 1A).

Antennae of male P. /imitata were more

sensitive to the pheromone components

than female antennae (with the exception of

0.1 ng Z9-14:Ac), responding with deflec-

tions significantly higher than those to hex-

ane at 1 ug for each individual compound

and as low as 100 ng for the blend (Fig.

1B). Male response to the plant volatile was

also significantly higher than their response

to hexane (Fig. 1B). Females had signifi-

cantly (Fy;9= 125.85; P < 0.0001) lower

antennal deflections (least squares mean +

SE: -6.13mV + 0.30mV) when compared to

males (least squares mean + SE: -11.41mV

+ 0.36mV). Comparisons of female and

male antennal responses to individual doses

showed that females had significantly

(F9.352 = 25.11; P < 0.0001) lower antennal

deflections for 10 ug Z9-14:Ac, and for

both | ug and 10 ug of Z11-14:Ac and the

94:6 ratio of Z11-14:Ac: Z9-14:Ac. Male

antennal response to the plant volatile was

also significantly higher than the female’s

(F 0.352 = 275.1] he P= 0.00134).

Female behaviour in a pheromone-

permeated environment. Observations of

females in clean air and Z11-14:Ac-

permeated environments showed that there

were no significant differences in onset of

calling (P > 0.934), frequency of calling

(F).4=0.6859; P => 0.4157), or duration of

calling (P > 0.890) (Fig. 2) among females.

However, females moved significantly less

in the Z11-14:Ac_ environment

(F) 1;=5.0999; P < 0.0452) than in clean air

(Fig. 2). There was no observed positional

bias in the tunnel for either chamber with

respect to any of the observed behaviours

[(P = 0.152), (F1,14=3.5383; P = 0.0809), (P

> 0.193) and (F;j9=1.1703; P = 0.3047)

respectively]. No vertical positional effects

were observed for frequency of calling or

movement [(F)24=0.9835; P > 0.4352) and

(F,.1;=0.8725; P = 0.5107) respectively].

DISCUSSION

We have found that adult female P. limi-

tata (0 to 24 h old) are able to perceive both

of their pheromone components; however,

significant responses to the main compo-

nent alone or in a blend were only detected

for doses of 10 ug or greater, in contrast to

males, which responded to | ug of the indi-

vidual compounds and to 100 ng of the

blend. Interestingly, female antennae gave a

significant response to only the lowest (0.1

ng) and the highest (10 ug) doses of the

minor component Z9-14:Ac. It is possible

that as treatments were presented randomly

in increasing concentrations, 0.1 ng Z9-

14:Ac would have contacted a relatively

‘fresh’ antenna compared to subsequent

doses, and as Z9-14:Ac is the minor phero-

mone component, occurring at approxi-

mately 6 to 9% in the female gland

(Roelofs et al. 1976), the antennae may be

able to perceive it at a lower level than Z11-

14:Ac; however, a similar phenomenon was

not observed for the blend or for the males.

The ability of female P. /imitata anten-

nae to perceive their own pheromone com-

ponents, albeit at high doses, is not surpris-

ing as this has been shown for other tortri-

cids. Palaniswamy and Seabrook (1978)

noted that female Choristoneura fumiferana

(Clemens) (Lepidoptera: Tortricidae) dis-

played threshold response levels corre-

sponding to pheromone concentrations

above which various behavioural patterns

became evident, such as increased walking,

extension of the ovipositor and antennal

grooming. Ross ef al. (1979) examined re-

sponse of female C. fumiferana antennae to

differing doses of pheromone and with in-

creasing age, finding that females have a

higher threshold for pheromone response

than males and that female antennae are at

their peak responsiveness in 3 to 6 d old

insects. Higher response thresholds may be

explained by the fact that antennae of fe-

male C. fumiferana have one-third to one-

half the number of sensilla trichodea as

J. ENTOMOL. SOc. BRIT. COLUMBIA 102, DECEMBER 2005 i

300

200

100 =]

Ses

Normalized Percentage Female Pandemis limitata Antennal Deflection

GQRoess— *

SAS NS

hexane PV

= |

ire}

p=]

iro}

oOo

(is)

(so)

{=}

Z9-14:0Ac 211-14:;0Ac 211:29-14: OAc (94:6)

Normalized Percentage Male Pandemis limitata Antennal Deflection

ie:

SEE |

hexane PV 0.1ng ing 10ng 100ng 14g 8 10ng pe 1ng 10ng 100ng 1g 104g O0.1ng ing 10ng 100ng ing § 10ng

Z29-14:0Ac Z211-14:0Ac Z211:Z9-14:O0Ac (94:6)

Pheromone Stimulus (10 ul volume)

Figure 1. Mean (+ SE) normalized electroantennogram responses of antennae from 0 to 24 h old A)

female (n = 6) or B) male (n = 4) Pandemis limitata to synthetic (Z)-11-tetradecenyl acetate (Z11-

14:0Ac) or (Z)-9-tetradecenyl acetate (Z9-14:O0Ac), or both in a ratio of 96:4. (E)-2-hexanal was

puffed over each antenna before and after each pheromone source for the purpose of normalization of

each antenna over time and HPLC-grade hexane was puffed over each antenna at the beginning and

end to establish a base response. All stimulus puffs were spaced by 30 s intervals. Normalized per-

centage deflections were calculated using the plant volatile (PV) response as the standard. Asterisks

indicate a significant difference (P < 0.05) from the hexane base response as determined by an

ANOVA followed by Dunnett’s test, treatment versus hexane (a = 0.05).

8 J. ENTOMOL. SOC. BRIT. COLUMBIA 102, DECEMBER 2005

= a a

g 8

(a)

3 aS

S D> 4

D> cS

= =

r= re)

re) a)

S = 3

oO @o

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5 ra)

£ ro

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E S

fm £

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= 9

Clean Air 211-14:Ac Clean Air 211-14:Ac

A B

180 a 3.5

ro) ex b

(29) (oe

s 160 =

{oe}

s 8 30

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oO Le)

® 140 =

< =

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w —

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0 s 0.0

Clean Air 211-14:Ac Clean Air 211-14:Ac

CG D

Atmospheric treatment

Figure 2. Mean (+ SE) duration (A and C) or frequency (B and D) of 0 to 24 h old female P.

limitata in either side of the divided permeation tunnel, exposed to either clean air or (Z)-11-

tetradecenyl acetate (Z11-14:Ac) released from 3M microcapsules applied to the upwind portion

of one side of the tunnel. Individually-caged females were placed in the tunnel (n = 15), immedi-

ately before pheromone application ca. 2 h before scotophase. Females were observed every 15

m from application until initiation of scotophase for onset of calling, and continuously once

scotophase began for frequency and duration of calling and frequency of movement. For each

behaviour observed, bars with the same letter are not significantly different (P > 0.05).

J. ENTOMOL. Soc. BRIT. COLUMBIA 102, DECEMBER 2005

male antennae (Albert and Seabrook 1973).

In Trichoplusia ni Hiibner (Lepidoptera:

Noctuidae), Light and Birch (1979) found

that antennal response in females was 25%

that of males. Our results show that anten-

nal responses in female P. /imitata over a

wide range of pheromone doses is 46.7%

less than that of males, with significant dif-

ferences observed at high doses for all com-

pounds tested. A difference was even ob-

served for the plant volatile.

Simultaneous observations of females in

clean and pheromone-permeated air, from

ca. 2 h before initiation of scotophase until

3 h into scotophase, showed no difference

in onset of female calling time (Fig. 2A).

However, a larger sample size may reveal

that females in environments permeated

with Z11-14:Ac do initiate calling earlier as

there was a large variance among females.

No differences were detected with respect

to duration or frequency of calling once

scotophase began (Fig. 2B, C). We did not

have the ability to measure female output of

pheromone directly in a background of

Z11-14:Ac, therefore we must rely on ob-

servations of behaviour to give us an indi-

cation of female response. We did not ob-

serve changes in frequency or duration of

calling, as one might expect if females were

altering concentrations or outputs of their

effluvia to compete with the level of com-

pounds perceived in the background

(N.C.D., unpublished data). Of course,

these changes, as well as a change in onset

of calling, may not be observable in the first

3 h of scotophase, only becoming apparent

after the female has been in the environ-

ment continuously for more than 24 h

(Palaniswamy and Seabrook 1985). In fact,

studies on female C. fumiferana have found

that female EAG responses to their sex

pheromone increase with age at least until

three days after emergence (Palaniswamy

and Seabrook 1978), indicating that females

may become more sensitive to their phero-

mone over time. As we only looked at re-

sponse in 0 to 24 h old P. limitata females

for a 5 h period, we would not have ob-

served such a phenomenon if one existed.

Our observations showed females

moved less often in air permeated with

Z11-14:Ac than females in the clean air

(Fig. 2D). It is possible that the tendency to

remain still in the pheromone background

may translate, in time, to increased calling,

as observed in C. fumiferana (Palaniswamy

and Seabrook 1985). Sanders (1987) found

that although pheromone in the background

clearly increased flight activity of both vir-

gin and mated females, virgin females re-

mained inactive for 48 h after emergence,

even in the presence of the pheromone. As

we observed increased movement of fe-

males in clean air, this does not appear to

be happening in P. /imitata, although obser-

vations over time would determine if the

level of activity increases with age.

Similar to our results in P. limitata, El-

Sayed and Suckling (2005) found that per-

manent exposure of female FEupoecillia

ambiguella (Htibner) (Lepidoptera: Tortri-

cidae), Lobesia botrana (Denis and Schif-

fermiiller) (Lepidoptera: Tortricidae), or

Spodoptera littoralis (Boisduval)

(Lepidoptera: Noctuidae) to their main

pheromone compounds did not alter the

timing, duration or frequency of calling. In

addition, Weissling and Knight (1996)

found that the temporal pattern of calling in

Cydia pomonella (L.) (Lepidoptera: Tortri-

cidae) was unaffected by the major compo-

nent of their pheromone; however, the like-

lihood of calling was increased. Neverthe-

less, other species have been found to initi-

ate calling earlier in pheromone environ-

ments, such as female C. fumiferana

(Palaniswamy and Seabrook 1985). At the

other extreme, Evenden (1998) observed a

delay in calling for female Choristoneura

rosaceana (Harris) (Lepidoptera: Tortrici-

dae) in field plots treated with their com-

plete pheromone blend, followed by a po-

tential reduction in time spent calling com-

pared to their counterparts in clean air. It is

important to remember that our experi-

ments, as well as those of El-Sayed and

Suckling (2005), did not use the complete

pheromone blend in the background, which

could have a greater impact on female re-

sponse.

While experiments described here do

10 J. ENTOMOL. SOC. BRIT. COLUMBIA 102, DECEMBER 2005

not encompass all of the potential factors

that may impact females in a pheromone

environment, such as background phero-

mone dose, completeness of pheromone

blend, age of females or changes in mating

status, our results are relevant to previous

work on disruption of pheromone commu-

nication in P. limitata (Judd et al. 2005;

N.C.D., unpublished data). The presence of

the major pheromone component (Z11-

14:Ac, applied at 10 mg ai - m7) does not

appear to cause female P. J/imitata to

change their calling behaviour, except with

respect to frequency of movement, during

the first 3 h of the first scotophase. As such,

alternative explanations, such as mode of

delivery of the gland extract, as well as the

use of nonchemical cues to attract males

like sound or vision (Castrovillo and Cardé

1980), or even chemical cues not associated

with the sex pheromone gland, need to be

explored to determine what cues become

important to males as they search for fe-

male moths in pheromone-permeated envi-

ronments (N.C.D., unpublished data).

ACKNOWLEDGEMENTS

We thank Brenda Lannard for modifica-

tions to the permeation flight tunnel, rearing

insects and other technical assistance;

Regine Gries for supplying chemicals;

Lynn Lashiuk for laboratory assistance;

Don Thomson and Kent Nielsen for techni-

cal advice on 3M MEC pheromone formu-

lations; Michael Roach for providing us

with Filtrete” material; Tom Lowery and

Maya Evenden for kindly reviewing an

earlier version of this manuscript. This re-

search was funded by 3M Canada and the

Agriculture and Agri-Food Canada Match-

ing Investment Initiative.

REFERENCES

Albert, P.J. and W.D. Seabrook. 1973. Morphology and histology of the antenna of the male eastern spruce

budworm, Choristoneura fumiferana (Clem.) (Lepidoptera: Tortricidae). Canadian Journal of Zoology

51: 443-448.

Barnes, M.M., J.G. Millar, P.A. Kirsch, and D.C. Hawks. 1992. Codling moth (Lepidoptera: Tortricidae)

control by dissemination of synthetic female sex pheromone. Journal of Economic Entomology 85: 1274-

E277.

Birch, M.C. 1977. Response of both sexes of Trichoplusia ni (Lepidoptera: Noctuidae) to virgin females

and to synthetic pheromone. Ecological Entomology 2: 99-104.

Castrovillo, P.J. and R.T. Cardé. 1980. Male codling moth (Laspeyresia pomonella) orientation to visual

cues in the presence of pheromone and sequences of courtship behaviours. Annals of the Entomological

Society of America 73: 100-105.

DeLury, N.C., G. Gries, R.Gries, G.J.R. Judd, and J.J. Brown. 1999. Sex pheromone of Ascogaster quadri-

dentata, a parasitoid of Cydia pomonella. Journal of Chemical Ecology 25: 2229-2245.

El-Sayed, A.M. and D.M. Suckling. 2005. Behavioural observations of mating disruption in three lepidop-

teran pests. Behaviour 142: 717-729.

Evenden, M.L. 1998. Semiochemical-based disruption of mate-finding behaviour in Choristoneura

rosaceana (Harris) and Pandemis limitata (Robinson) (Lepidoptera: Tortricidae) in British Columbia

Apple Orchards. PhD Thesis, Simon Fraser University, Burnaby, BC, Canada.

JMP 5.1. 2003. SAS Institute, Inc., Cary, NC, USA.

Judd, G.J.R., N.C. DeLury, and M.G.T. Gardiner. 2005. Examining disruption of pheromone communica-

tion in Choristoneura rosaceana and Pandemis limitata (Lepidoptera: Tortricidae) using microencapsu-

lated (Z)-11-tetradecenyl acetate applied in a laboratory flight tunnel. Entomologia Experimentalis et

Applicata 114: 35-45.

Light, D.M. and M.C. Birch. 1979. Electrophysiological basis for the behavioural response of male and

female Trichoplusia ni to synthetic female pheromone. Journal of Insect Physiology 25: 161-167.

Michell, E.R., J.R. Webb, and R.W. Hines. 1972. Capture of male and female cabbage loopers in field traps

baited with synthetic sex pheromone. Environmental Entomology 1: 525-526.

Palaniswamy, P. and W.D. Seabrook. 1978. Behavioural responses of the female eastern spruce budworm

Choristoneura fumiferana (Lepidoptera: Tortricidae) to the sex pheromone of her own species. Journal of

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Chemical Ecology 4: 649-655.

Palaniswamy, P. and W.D. Seabrook. 1985. The alteration of calling behaviour by female Choristoneura

fumiferana when exposed to synthetic sex pheromone. Entomologia Experimentalis et Applicata 37: 13-

LO:

Palaniswamy, P., P. Sivasubramanian, and W.D. Seabrook. 1979. Modulation of sex pheromone perception

in female moths of eastern spruce budworm, Choristoneura fumiferana by altosid. Journal of Insect

Physiology 25: 571-574.

Roelofs, W., A. Cardé, A. Hill, and R. Cardé. 1976. Sex pheromone of the threelined leafroller, Pandemis

limitata. Environmental Entomology 5: 649-652.

Ross, R.J., P. Palaniswamy, and W.D. Seabrook. 1979. Electroantennograms from spruce budworm moths

(Choristoneura fumiferana) (Lepidoptera: Tortricidae) of different ages for various pheromone concen-

trations. The Canadian Entomologist 111: 807-816.

Sanders, C.J. 1987. Flight and copulation of female spruce budworm in pheromone-permeated air. Journal

of Chemical Ecology 13: 1749-1758.

Shorey, H.H. and R.L. Hale. 1965. Mass-rearing of the larvae of nine noctuid species on a simple artificial

medium. Journal of Economic Entomology 58: 522-524.

Thomson, D., J. Brunner, L. Gut, G. Judd, and A. Knight. 2001. Ten years implementing codling moth

mating disruption in the orchards of Washington and British Columbia: starting nght and managing for

success! International Organization for Biological Control WPRS Bulletin 24: 23-30.

Weissling, T.J. and A.L. Knight. 1996. Oviposition and calling behavior of codling moth (Lepidoptera:

Tortricidae) in the presence of codlemone. Annals of the Entomological Society of America 89: 142-147.

J. ENTOMOL. SOc. BRIT. COLUMBIA 102, DECEMBER 2005

J. ENTOMOL. Soc. BRIT. COLUMBIA 102, DECEMBER 2005 13

Moisture tempers impairment of adult Otiorhynchus

sulcatus (Coleoptera: Curculionidae) climbing ability by

fluoropolymer, talc dust, and lithium grease

MICHAEL K. BOMFORD' and ROBERT S. VERNON"”

ABSTRACT

As part of a project to develop tools for the physical exclusion of flightless root

weevils, adult black vine weevils (BVW), Otiorhynchus sulcatus (F.), were placed

in open enclosures with smooth walls of glass, plastic or aluminum to test their abil-

ity to escape by climbing. Enclosure walls were left untreated or were treated with

substances known to reduce insect climbing ability: fluoropolymer, powdered talc

and lithium grease. No BVW escapes were observed under dry conditions, but all

treatments allowed some escapes under wet conditions, suggesting that moisture

helps BVW adults scale treated surfaces. The results help explain the ability of root

weevils to overcome physical barriers under field conditions.

Key Words: black vine weevil, insect barrier, physical control, root weevil

INTRODUCTION

Like other root weevils, the black vine

weevil (BVW), Otiorhynchus sulcatus (F.),

feeds on roots as a larva, leaves as an adult

and disperses by walking during the wing-

less adult phase. The biology and control of

BVW was reviewed by Moorhouse ef al.

(1992).

Flightless root weevils could be particu-

larly susceptible to physical control by ex-

clusion. While hardly a new strategy

(Feytaud 1918), physical control has re-

cently been the subject of some interest

(Vincent et a/. 2003). An aluminum fence

with a band of lithium grease (Cowles

1995, 1997) or fluoropolymer-coated tape

(Bomford and Vernon 2005) near the upper

edge can limit root weevil movement. Also

effective is a portable plastic trench, de-

signed to exclude Colorado potato beetle,

Leptinotarsa decemlineata (Say) (Hunt and

Vernon 2001). Both the fence and the

trench have reduced root weevil immigra-

tion into strawberry plots by about two-

thirds (Bomford and Vernon 2005). Sticky

bands and fluoropolymer-coated tape on

shrub stems are both recommended to re-

duce adult feeding on leaves (Antonelli and

Campbell 2001).

Like other insects, root weevils climb

using a combination of tarsal claws to hook

textured surfaces and adhesive pads on their

tarsomeres to attach to smooth surfaces.

These adhesive pads consist of densely

packed setae, each with a terminus a few

um in diameter that attaches to the surface

through weak van der Waals and capillary

forces (Arzt et al. 2003, Gao and Yao

2004). The sum of these weak forces can

support the insect only if a sufficient pro-

portion of the setae contact the surface.

Insect tarsi cannot adhere to surfaces

with sufficient micro texture to prevent a

large proportion of setae from making con-

tact, but insufficient macro texture for tarsal

claws to grip. Lithium grease is one such

surface, consisting of an open, fibrous crys-

tal matrix that holds tiny (~1 um) oil drop-

lets (Wilson 1964); fluoropolymers have

similar properties (Hougham 1999).

Smooth surfaces coated with fine, loose

' Pacific Agri-Food Research Centre, Agriculture and Agri-Food Canada, Box 1000, Agassiz, BC,

Canada, VOM 1A0

* To whom correspondence should be addressed

14 J. ENTOMOL. SOC. BRIT. COLUMBIA 102, DECEMBER 2005

dust particles are similarly difficult for in-

sects to climb because their tarsi adhere to

dust particles, which slip away from the

surface (Boiteau. and Vernon 2001).

Smooth dusted surfaces have shown poten-

tial as physical barriers to Colorado potato

beetle (Boiteau et al. 1994, Boiteau and

Osborn 1999), and root weevil (M.K.B.,

personal observation) movement.

This paper describes laboratory and

field studies testing the influence of surface

treatment and moisture on the ability of

adult BV W to climb materials that could be

used to construct physical barriers to root

weevil migration. The results are intended

to aid in the development of physical con-

trol tactics for root weevil management.

MATERIALS AND METHODS

Test insects. BVW adults were col-

lected from an apple rootstock nursery and

home garden near Vancouver, BC in late

summer and early fall. Weevils were held

for no more than 30 days at 20 °C under a

16:8 h L:D photoregime in clear plastic

cages containing potted strawberry, Fra-

garia x ananassa (Duchesne) plants as a

food source.

Glass surface treatments (dry). Eleven

250 mL glass Erlenmeyer flasks were

washed and dried. One end of a length of

surgical tubing was placed in each flask to

allow air to escape as it was dipped upside-

down in liquid fluoropolymer (Insect-A-

Slip, BioQuip, Rancho Dominguez, CA)

(four flasks), or powdered talc (four flasks),

evenly coating the top 3 cm of the neck

with the dip treatment. Excess talc and

fluoropolymer were shaken off, and the

fluoropolymer was allowed to dry to a hard,

smooth finish. Three remaining flasks were

left untreated as controls (unequal replica-

tion reflects flask availability). Flasks were

randomized, five BVW adults were placed

in each and all flasks were placed in an

incubator held at 20 °C and 20% RH under

a 16:8 h L:D photoregime. The number of

weevils in each flask was recorded after 0.5

h and all escapees were removed from the

incubator. The number of weevils remain-

ing in each flask was recorded again after

24 h when the experiment was terminated.

Data were analyzed by one-way ANOVA

for unequal number of replicates and treat-

ment means were separated by Tukey’s

honestly significant difference test (JMP

Version 4.0.4, SAS Institute 2001).

Outdoor plots. Three, one m square

enclosures, constructed from aluminum

gutters (75 mm deep by 120 mm wide)

sealed at all joints with hot glue, were sunk

into freshly-tilled soil so that the soil sur-

face was even with the upper lip of the gut-

ter. The soil inside each enclosure was cov-

ered with a square of landscape fabric with

its edges screwed to the inner gutter wall.

One litre of 1:1 water:dormant oil emulsion

was poured into each gutter.

Each enclosure was randomly assigned

to one of three treatments: The landscape

fabric pad was separated from the gutter by:

1) a 20 cm high aluminum fence with

fluoropolymer-coated tape (EnviroSafe,

Professional Ecological Services, Victoria,

BC) attached to the upper edge of the inner

surface (fence); 2) a portable plastic trench

(Hunt and Vernon 2001) coated inside with

dormant oil (trench); or 3) no barrier

(control).

Two days after plot setup, marked BVW

adults were released in the centre of each

enclosure at 2200 h, a time of high activity

among wild specimens observed in the area.

A flashlight was used to observe weevil

movement at five min intervals for one h

after release. Weevils that entered the alu-

minum gutter and became trapped in the

dormant oil emulsion (successful escapes)

were recorded during the first hour and

again the following morning at 1000 h. The

experiment was conducted in the same plots

three times (13, 18, and 20 August 1997),

with ten BVW per treatment in the first

replicate and 20 in the others. Hourly RH

readings recorded at the Vancouver Interna-

tional Airport (6 km from study site) during

each observation period were used to esti-

J. ENTOMOL. Soc. BRIT. COLUMBIA 102, DECEMBER 2005

mate the ambient RH range for each repli-

cate (Environment Canada 2005).

A two-way ANOVA was used to test for

treatment and replicate effects on weevil

escape rates after one and 12 h, and for in-

teraction between factors (JMP Version

4.0.4, SAS Institute 2001). Means were

separated by Tukey’s HSD test.

Plastic surface treatments (wet vs.

dry). Forty, 35 mL black plastic film canis-

ters (30 mm diameter by 50 mm deep) were

washed, dried, and randomly assigned to

one of four treatments: ten were untreated

controls; ten were dusted with powdered

talc; ten were coated with liquid fluoropoly-

mer; and ten had a 2.5 cm band of white

lithium grease applied to the inner top edge.

The following day, half of the canisters

from each group were rinsed with water and

then emptied, leaving droplets inside. These

were placed in a sealed plastic container

containing an open water source to create a

saturated environment. The remaining un-

rinsed canisters were placed in an identical

container without a water source (ambient

RH: 50-74%, Environment Canada 2005)

and left open to allow air circulation. Canis-

ter order was randomized within each con-

tainer.

Two BVW adults were placed at the

bottom of each canister. The number of

weevils remaining in each canister was re-

corded and escapees were removed at 0.5 h

intervals for 3.5 h. Canisters were not

treated on the outside, so re-entry was pos-

sible, but never observed. ANOVA was

used to test for treatment effects within

each container and means were separated

by Tukey’s HSD test (JMP, Version 4.0.4,

SAS Institute 2001). A t-test was used to

compare escape rates between containers

for each treatment.

Plastic surface treatments (saturated

vs. ambient RH). Eighteen, 290 mL plastic

cups (50 mm diameter at base, 70 mm di-

ameter at opening, 100 mm deep) were ran-

domly assigned to one of three treatments:

SIX were untreated controls; six had a 2.5

cm strip of white lithium grease applied

around the inner top edge; and six were

dusted with powdered talc.

Three BVW adults and a moist cotton

swab were placed in the bottom of each

cup. Cups from each treatment were evenly

divided into two identical plastic tubs, each

containng a damp cloth. One tub was

sealed to create a saturated environment in

which condensation formed on the plastic

cups; the other tub was left open to allow

air circulation and prevent condensation

(regional ambient RH: 67-95%, Environ-

ment Canada 2005). Tubs were held at 20 °

C for 20 h. Any weevils that escaped from

their cups were removed from the tubs at

hourly intervals for the first six hours and

then every other hour thereafter until the

study was terminated. The mean number of

escapes per cup was calculated for each

treatment in the open and sealed containers.

ANOVA was used to test for treatment ef-

fects within each container and means were

separated by Tukey’s HSD test (JMP Ver-

sion 4.0.4, SAS Institute 2001). A t-test was

used to compare escape rates between con-

tainers for each treatment. The time re-

quired to escape under each combination of

conditions was estimated by Kaplan-Meier

analysis and a Wilcoxon t-test was used to

test for differences in escape times between

treatments (JMP Version 4.0.4, SAS Insti-

tute 2001).

RESULTS

Glass surface treatments (dry). Al-

most all (93.3 + 3.3%, n = 3) weevils in the

control flasks escaped, but none (0.0 +

2.9%, n = 4) escaped from flasks treated

with talc dust or fluoropolymer, demon-

strating a strong treatment effect (Fr. =

285, P < 0.001). All escapes from the con-

trol flasks occurred within the first 30 min

of the 24 h observation period. Weevils in

the fluoropolymer treated flasks were fre-

quently observed walking up the glass to

the fluoropolymer strip and were occasion-

ally able to climb part-way over this strip

before falling. When the experiment was

16 J. ENTOMOL. Soc. BRIT. COLUMBIA 102, DECEMBER 2005

terminated, approximately half of the wee-

vils in the fluoropolymer treated flasks

were on the flask walls. Weevils in the talc

treated flasks showed much less ability to

scale the glass walls and were all at the

bottom of the flask at the end of the experi-

ment.

Outdoor plots. Treatment and replica-

tion both affected weevil escape rates (F> 14;

= 189 and 25, respectively; P < 0.001) and

an interaction was found between these

factors (F'414;= 12; P < 0.001). Almost all

weevils left control plots over the course of

all replications (Figure 1), but escapes from

plots surrounded by physical barriers only

occurred in the third replication, conducted

under light rain and high humidity condi-

tions. Under the drier conditions of the first

two replications weevils quickly climbed

the aluminum fence to the lower edge of the

fluoropolymer-coated tape and were unable

to climb further for the duration of the test.

Most weevils surrounded by plastic

—e— Control

dp) 10 —e— Fence

+l —<— Trench

xs 8

n°

(ok

14)

8 4

cD)

& 2

O O

trenches fell into the trenches and none

emerged. Under the wet conditions of the

third replication the first of 20 weevils was

able to walk onto the fluoropolymer within

5 min of its release. Within 20 min, four

more had achieved this feat, two had

reached the top of the aluminum fence and

one had crossed the trench. Statistical com-

parison of the replications showed a higher

escape rate from the fenced treatment in the

third repetition after 12 h (Fy47= 26; P <

0.001), but not from the trenched treatment

(F347 a 25: P= 0.09).

Plastic surface treatments (wet vs.

dry). Under dry conditions all weevils es-

caped from untreated canisters but none

escaped from those treated with talc,

fluoropolymer, or white lithium grease

(Table 1). Talc lost its dusty character un-

der wet conditions, allowing more escapes

(Table 1). The dried fluoropolymer reverted

to a liquid state in the presence of moisture,

clumping on tarsi and allowing only one

rao

Pao =o

720

Time after release (min)

Figure 1. Black vine weevil escapes from a one m square area surrounded by a 20 cm high

aluminum fence with fluoropolymer-coated tape attached inside (fence), a portable exclusion

trench (trench) or no barrier (control). Observations were made at 5 min intervals for 1 h after

insect release and 12 h after release. Lower error bars omitted from fence data points for clar-

ity. Final means labeled with the same letter do not differ significantly at a = 0.05 (Tukey’s

HSD test, n = 3).

J. ENTOMOL. SOC. BRIT. COLUMBIA 102, DECEMBER 2005

Table 1.

Mean percentage of adult black vine weevils that left plastic canisters or cups that were un-

treated (control) or coated inside with dried fluoropolymer, white lithium grease (grease) or

powdered talc (talc). Canisters and cups were placed in an open container (ambient RH) or a

closed container with an open water source (saturated RH). Canisters were rinsed immediately

before being placed in the closed container, leaving their surface wet.

Io 3 (cos Pea eka Oran Te Sas ee ee ee ees Se ee

C ys

anister escapes (%

n=10

Dry surface, Wet surface,

Treatment ambient RH saturated air

Cup escapes (%) ,

n=9

Dry surface, Dry surface,

ambient RH saturated air

Control 100 A 90 aA tg=2.3,P=0.15 89 aA 100 aA to=1.0,P=0.33

ae OA 10 bA hg=23,P=0.15 - :

Grease 0 0b O0bA 33 bA t=4.0, P= 0.06

Talc 0B 70 aA tig=73,P<0.0001 0b Oc

F339 = 30 Fong = 64 Fron, = 38

P< 0.0001 P<0.0001 P<0.0001

'Means followed by the same lower case letter within a column do not differ significantly

(Tukey’s test, a= 0.05); those followed by the same upper case letter within a study and row do

not differ significantly (t-test, a= 0.05).

escape. Weevils in fluoropolymer-treated

canisters largely ceased their activity until

the experiment ended. No weevils escaped

from moistened grease-treated canisters.

Plastic surface treatments (saturated

vs. ambient RH). Visible condensation

first appeared on cups in the saturated envi-

ronment 8 h after the test began and was

very heavy by the end of the test. No con-

densation was seen on cups in the lower

humidity environment. Almost all weevils

escaped from control cups within the first

hour of observation; the only weevil that

did not escape from a control cup in an hour

did not escape at all (Table 1). No weevils

escaped from cups treated with talc in either

container. One third of the weevils escaped

from grease-treated cups in the saturated

environment, but none escaped in the ambi-

ent RH environment (Table 1). On average,

escapes from greased cups took longer than

escapes from untreated cups in the sealed

container (16.7 + 0.7 versus 0.7 + 0.7 h,

respectively; y° = 27, df = 2; P < 0.001).

Mean escape times from untreated cups did

not differ between the open and sealed con-

tainers.

DISCUSSION

Under dry conditions talc dust, fluoro-

polymers and lithium grease treatments

rendered several smooth surfaces (glass,

plastic, and aluminum) unclimbable to

BVW adults for the duration of our tests.

Equivalent treatments were sometimes less

effective under wet conditions, or in satu-

rated environments. This may help explain

why physical barriers that would be ex-

pected to offer total exclusion, based on

observations under dry conditions, exclude

only two-thirds of root weevils in the field

(Bomford and Vernon 2005).

Most adult weevils quickly attempted to

leave the open containers we used for our

tests. Their success in exiting, and the

length of time they took to leave, were con-

sidered indicators of the difficulty they had

in scaling the barriers they faced. Under dry

conditions, surface treatments eliminated

escapes; under wet conditions they usually

reduced the proportion of insects able to

escape and lengthened escape times.

Cowles (1995) has suggested that root

18 J. ENTOMOL. SOC. BRIT. COLUMBIA 102, DECEMBER 2005

weevils are able to evade physical barriers

because natural bridges form over other-

wise unclimbable surfaces. He has seen

field debris, such as twigs, adhering to the

white lithium grease on his barriers, and

plant canopies touching across barriers

(R.S. Cowles, pers. comm., see Acknowl-

edgements). We have also seen natural

bridges that could allow root weevils to

cross portable trench barriers in field stud-

ies (Bomford and Vernon 2005), but these

were not a factor in the tests reported here.

We observed repeated instances of

BVW adults crossing vertical surfaces

treated with fluoropolymer, talc dust, and

lithium grease in the presence of moisture.

BVW adults scaled talc-dusted plastic that

had been lightly rinsed to mimic rainfall on

a dusted plastic exclusion trench. Similar

observations have been reported previously

for Colorado potato beetles challenged by

plastic-lined trenches after rainfall in field

studies (Boiteau eft a/. 1994). Rinsing did

not render greased surfaces climbable in

one test, reflecting field observations in

which greased aluminum barriers excluded

root weevils after irrigation (Cowles 1995).

We did, however, observe BVW scaling

greased plastic with visible surface conden-

sation in a high humidity environment and

scaling fluoropolymer-treated aluminum in

a light rain shower. We are unaware of

other reports of moisture enhancing an in-

sect’s ability to scale fluoropolymer or lith-

lum grease-coated surfaces. These observa-

tions lead us to suggest that the insects’

tarsal pads adhere to condensation on

treated surfaces. Essentially we hypothesize

that the insects can overcome physical bar-

riers by walking on water.

More rigorous tests of this hypothesis

are necessary. The studies reported here

reflect a variety of treatment combinations

observed under different conditions. Ex-

perimental factors were sometimes con-

founded. For example, BVW were unable

to scale a fluoropolymer treated fence under

dry conditions two and seven days after the

fence was erected, but scaled the same

fence in a light rain shower nine days after

setup. We attributed this difference to the

presence of moisture, but it might also have

been an effect of fence age. Similarly, our

analyses of interactions between surface

treatment and environment were con-

founded by the fact that surface treatments

were replicated within environments, but

only one instance of each environment was

tested in any study. Our observations sug-

gest intriguing avenues for further study,

not definitive conclusions.

ACKNOWLEDGEMENTS

We thank Murray Isman, University of

British Columbia, for providing laboratory

facilities and Lynne Guinet and John Traas

for access to weevil collection sites. Mark

Bomford, John Borden and Dave Raworth

gave valuable technical assistance, and

Sheila Fitzpatrick and Richard Cowles of-

fered comments on the manuscript. Richard

Cowles, The Connecticut Agricultural Ex-

periment Station, Windsor, CT, information

used with permission. Studies were sup-

ported in part by the H.R. MacCarthy,

Thelma Finlayson, and Simon Fraser Uni-

versity Graduate Fellowships and by a grant

from the BC Nursery Trades Association.

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J. ENTOMOL. Soc. BRIT. COLUMBIA 102, DECEMBER 2005

Sources of Spring and Fall Hop Aphid,

Phorodon humuli (Schrank), (Homoptera: Aphididae)

Migrants in South Central Washington

LAWRENCE C. WRIGHT'”, WYATT W. CONE!

and DAVID G. JAMES!

ABSTRACT

The hop aphid, Phorodon humuli (Schrank), flies from hop, Humulus lupulus L., to its

overwintering Prunus spp. hosts in the fall. The sources of these aphids were not known

because much of the aphid flight occurs after hop plants are removed from fields during

harvest. We found that the bottoms of hop plants remaining alive in harvested hop yards

averaged 1.7 to 5.8 hop aphids per leaf in three years of sampling. Unharvested hop

plants remaining after harvest averaged 32.8 to 127.1 aphids per leaf in two years. Feral

hops were also infested with hop aphids in late summer and early fall. Sources for the

spring aphid flight from Prunus spp. to hop included Prunus cerasifera Ehrhart, which

averaged 44.0 to 105.1 aphids per shoot in two years of sampling. Fruit-type Prunus

spp. trees growing on residential properties averaged 0.9 and 11.3 aphids per shoot in

the same years but few of these trees were found. Plum and prune orchards averaged 0

to 5.5 aphids per shoot in two years and estimates indicate that orchard trees are much

more numerous than other hop aphid host trees. Potential alternative management strate-

gies for hop aphid control are discussed.

Key Words: Homoptera, Phorodon humuli, hop, Humulus lupulus, Prunus, host plants

INTRODUCTION

The hop aphid, Phorodon humuli Hop is_ the aphid’s_ only

secondary

(Schrank), is an important pest of hop, Hu-

mulus lupulus L., in south central Washing-

ton state (WA) and in most hop-growing

areas of the Northern Hemisphere (Neve

1991). It is a holocyclic aphid that overwin-

ters in the egg stage on purple-leafed orna-

mental flowering plum, Prunus cerasifera

Ehrhart (also known as cherry plum or My-

robalan plum), Prunus divaricata Lede-

bour, Prunus domestica L., Prunus insititia

L., Prunus mahaleb L., and Prunus spinosa

L. (Blackman and Eastop 1994). Eggs hatch

in February and March followed by a vari-

able number of generations of parthenoge-

netic wingless females. The winged females

that fly to hop appear in WA in early to

mid-May and flight continues from mid-

July to early August (Wright et al. 1995).

(summer) host (Born 1968; Micinski and

Ruszkiewicz 1974; Eppler 1986). Partheno-

genetic, wingless females are produced on

hop during the summer (Campbell 1985;

Campbell and Tregidga 2005). In late Au-

gust, gynoparae (winged females) are pro-

duced on hop, which begin the flight back

to Prunus spp. Winged males that fly from

hop to Prunus spp. appear about mid-

September. Aphid flight often continues

into November and is terminated by foli-

age-killing frost (Wright ef a/. 1995). The

gynoparae give birth to a generation of

wingless females, the oviparae, which mate

with winged males and lay the overwinter-

ing eggs on Prunus spp. buds and stems.

Neither hop aphids nor their eggs have been

reported on hop during the winter. Further-

‘Irrigated Agriculture Research and Extension Center, Washington State University, 24106 N. Bunn

Road, Prosser, Washington 99350 USA

*To whom correspondence should be addressed

OB: J. ENTOMOL. Soc. BRIT. COLUMBIA 102, DECEMBER 2005

more, gynoparae do not settle on hop leaves

or reproduce on hop (Campbell and

Tregidga 2005).

The aerial parts of hop plants are killed

by fall frosts and only the hop roots, which

are several cm below the soil surface, sur-

vive the winter. In the spring, shoots grow

from the roots and they are trained to grow

up fiber strings which are tied to a trellis

that is about 5 m tall. During harvest (mid-

August to mid-September) hop plants are

cut at the top of the trellis and about 1 m

above ground, removed from the fields and

taken to stationary picking machines where

the cones are separated from the leaves and

stems. The cones are dried in large kilns at

60 °C and the waste leaves and stems are

chopped and spread on the fields soon after

harvest or after being stored in large piles.

It 1s considered unlikely that many aphids

could survive the picking process

(Campbell and Tregidga 2005). Following

harvest, about 1 m of basal foliage remains

alive in hop fields until it is killed by frost.

The amount of foliage remaining is quite

variable ranging from a few leaves to hun-

dreds of leaves per plant. Intact plants

growing up trellis poles remain in some hop

yards following harvest and feral (wild) hop

plants are also present in the hop-growing

region of WA (James ef a/. 2001). Approxi-

mately half of the gynoparae and very few

males have flown by the end of harvest

(Wright et al. 1995). One of our objectives

was to determine if harvested and unhar-

vested hop plants remaining alive in the

fields after harvest, as well as feral hop

plants, could be a source of fall migrants.

Another objective was to determine the

source of aphids that fly from Prunus spp.

to hop in the spring and summer. The hop-

growing area of Washington is an area of

diverse agriculture including a small num-

ber of plum or prune, Prunus domestica L.,

orchards. In addition, landowners have

planted ornamental and fruit Prunus spp.

near residences, businesses, and in parks.

Determining the sources of the spring and

fall migrants not only adds to our knowl-

edge of the aphid’s life cycle but also may

reveal alternatives to the traditional control

methods that are used on hops during the

growing season.

MATERIALS AND METHODS

Aphids in harvested hop yards. Hop

yards selected for sampling in three years

(1984, 1987, 1989) were in the Prosser -

Grandview area of the Yakima Valley, WA.

In 1984, plants in 11 harvested hop yards

were sampled between 25 September and

19 October. Apterae were identified in all

field studies described in this manuscript

with the aid of a 10X hand lens and the

descriptions in Blackman and _ Eastop

(1984). Hop aphids were counted in the

field on one leaf per piant from each of 200

randomly selected pla™:s in eight hop yards

and from 100 plants in three yards. One leaf

was sampled from each of 100 randomly

selected plants per yard: in 27 hop yards

(one yard had 94 samples) from 25 Septem-

ber to 6 October, 1987; and in 33 hop yards

(one yard had 89 samples) from 15 Septem-

ber to 9 October, 1989.

A small number of hop yards had vary-

ing numbers of unharvested, intact hop

plants growing up the trellis poles. Six to

100 (mean = 43.7) randomly selected un-

harvested plants were sampled in each of 11

yards between 25 September and 7 October,

1987 and 11 to 58 (mean = 27.2) unhar-

vested plants were sampled in each of nine

yards from 15 to 29 September 1989. One

leaf from about the 2 m height, which is a

representative sample (Wright et a/.1990),

was sampled per plant. The varieties sam-

pled in all years were Cascade, Ll

(Cluster), and Galena.

The mean aphids per leaf on harvested

plants was compared with the mean per leaf

on unharvested pole plants using the non-

parametric Wilcoxon Rank Sum test com-

puted by the NPARIWAY procedure of

SAS (SAS Institute 1988).

Aphids on feral hop plants. Six sites

with feral hop plants were located in the

J. ENTOMOL. SOC. BRIT. COLUMBIA 102, DECEMBER 2005

Yakima Valley of south central WA (James

et al. 2001). The plants grew on fences, or

poles, usually near roads. In 1999, the

plants were sampled on 7 to 8 September

and 11 to 12 October and in 2000, on 14 to

22 August and 18 to 19 September. Thirty

leaves were collected randomly per site and

the number of aphids per leaf were counted

under a stereomicroscope in the laboratory.

Survey of hop aphids on Prunus in the

spring. The survey area was divided into

two adjacent hop growing areas of WA: one

in western Benton County near Prosser, and

the other in eastern Yakima County near

Sunnyside, Grandview, and Mabton. Each

area was about 15,540 ha. Surveys were

conducted in 1990 (18 to 26 June) and 1991

(25 June to 5 July). In 1990 we drove the

roads in an unsystematic pattern and lo-

cated P. cerasifera and fruit varieties of P.

domestica by sight. Orchards were sampled

by selecting 10 trees at random and sam-

pling 10 shoots per tree. Hop aphids in

spring are concentrated on the new foliage

near the tips of the shoots (Wright ef ai.

1995). In addition to the hop aphid, we

found the mealy plum aphid, Hyalopterus

pruni (Geoffroy), and the leaf-curling plum

aphid, Brachycaudus_ helichrysi

(Kaltenbach). Ornamental and fruit trees at

residences and commercial properties that

were not orchards were sampled by exam-

ining 10 shoots per tree or shrub. Some

small trees did not have 10 shoots, so fewer

shoots were sampled on those trees. Aphid

numbers were expressed as the number per

shoot. Usually every tree at a site was sam-

pled but if a property had more than three

or four trees, a subsample of trees was se-

lected. In 1991, the survey was done sys-

tematically. Most of the roads in the sur-

veyed area are laid out in a grid of squares

that are 1.6 km on a side. Road sections of

1.6 km each were selected at random on a

map and 14 % of the roads in each area

were surveyed as in 1990. For orchards, the

number of trees per ha was calculated by

multiplying the number of orchards in the

surveyed area by 1,272, which was the av-

erage number of trees per plum and prune

farm in Benton and Yakima counties (the

counties of hop production) in 1992

(National Agricultural Statistics Service

1992) and dividing by the area surveyed.

The number of trees not in orchards was

determined by dividing the number of trees

in the survey by the hectares in the area

surveyed.

RESULTS

Aphids in harvested hop yards. We

found hop aphids on the bases of harvested

hop plants and on unharvested plants grow-

ing on trellis poles (Table 1). The unhar-

vested plants had significantly more aphids

per leaf than the harvested plants. Only two

yards in the three years of sampling had no

aphids in the samples.

Aphids on feral hop plants. In 1999,

we found a mean of 0.7 aphids per leaf on 7

to 8 September (range = 0 tol.6) and 20.9

on 11 to 12 October (range = 0 to 93.6). In

2000, there was a mean of 0.7 per leaf

(range = 0 to 1.7) on 14 to 22 August and

11.7 (range = 0 to 30.3) on 18 to 19 Sep-

tember.

Survey of aphids on Prunus in the

spring. In 1990, 14 commercial prune or-

chards were sampled and hop aphids were

found in four of them. The mean number of

aphids per shoot in all orchards was 5.5 but

most of the aphids were found in one or-

chard that averaged 81.0 aphids per shoot.

Fruit-type Prunus were found at three resi-

dences with one tree each and aphids were

found on two of the trees. The mean from

all three trees was 0.9 aphids per shoot.

Seventy-two purple-leafed ornamental plum

trees were sampled at 42 sites and hop

aphids were found on 50 trees at 32 sites.

The number of trees sampled per site

ranged from one to eight. The mean number

of hop aphids on all ornamental trees was

44.0 per shoot.

In 1991, we found four commercial

prune orchards and no hop aphids were

found in any of them. A total of seven fruit-

type plums were found at five residences

24 J. ENTOMOL. SOc. BRIT. COLUMBIA 102, DECEMBER 2005

Table 1.

Mean number of hop aphids per leaf on harvested and unharvested hop plants remaining in hop

yards in September and October. N, total number of leaves sampled (one leaf per plant). Z, test

statistic for Wilcoxon Rank Sum test.

Year Plant type Mean aphids (range) N Z

1984 harvested 4.4 (0.2 — 13.2) 1,900 na

1987 harvested 1.7 (0 — 18.9) 2,694 g2:

unharvested 32.8 (0.2 — 316.5) 481

1989 harvested 5.8 (0 — 56.4) 3,289 14.8!

unharvested 127.1 (0.2 — 481.6) 245

‘P<0.0001.

but aphids were found on only two trees at

one site with an average of 39.5 aphids per

shoot. The mean for all fruit trees at resi-

dences was 11.3 aphids per shoot. We sam-

pled 57 purple-leafed ornamental plum

trees at 37 sites and hop aphids were found

on 36 trees at 27 sites. The mean number of

hop aphids on all trees was 105.1 per shoot.

The estimated number of trees per ha was

1.16 for orchard trees, 0.017 for purple-leaf

ornamental flowering trees and 0.0016 for

fruit trees at residences.

DISCUSSION

Hop aphids were common in harvested

hop yards, indicating that harvested hop

yards were a major source of the aphids for

the fall flight to Prunus. Hop plants grow-

ing up the trellis poles had more leaves than

the bottoms of harvested plants and were

infested with more aphids per leaf (Table

1); however, unharvested plants were un-

common compared to the number of har-

vested plants, so they probably contribute a

small proportion of the hop aphids pro-

duced over the whole area.

Feral hop plants were infested with hop

aphids, occasionally with high numbers.

Hop is not native to the Pacific Northwest

(Hitchcock and Cronquist 1973) and only

female plants that produce seedless hop

cones are grown commercially in WA.

These factors may restrict the number of

feral hops growing in south central WA.

Wild hops may be an important source of

fall migrants in England (Campbell and

Tregidga 2005). Our observations indicate

that feral hops in south central Washington

are scarce compared to the number of com-

mercial hop plants but a more intensive

survey would be needed to determine the

population size of feral hops.

Our survey of Prunus spp. indicates that

purple-leafed ornamental flowering plums

were a major source of spring migrant hop

aphids. Only one commercial prune orchard

was heavily infested with hop aphids but,

because of the large number of trees in this

orchard, it could be a significant source of

aphids. Orchard trees are usually sprayed

with insecticides to control aphids and this

is probably the main reason aphid numbers

were generally low in orchards. Since this

survey was done, the plum and prune indus-

try has declined from 565 ha in Benton and

Yakima counties in 1992 to 311 ha in 2002

(National Agricultural Statistics Service

1992; 2002). The ornamental varieties were

much less abundant than orchard trees but

they were infested with higher densities and

they were well dispersed throughout the

survey area.

Knowing the sources of the spring and

fall migrating aphids and the timing of the

flights suggests some alternative aphid con-

trols. As gynoparae start flying before har-

vest is completed and males start flying

near the end of harvest in mid to late Sep-

J. ENTOMOL. SOC. BRIT. COLUMBIA 102, DECEMBER 2005

tember (Wright ef a/. 1995), controlling

aphids in harvested hop yards would reduce

the number of gynoparae but should be

more effective in reducing the number of

males. The desired result would be a reduc-

tion in the number of mated females and

eges on Prunus spp. Potential control of

aphids in harvested hop yards could involve

insecticide applications, destroying the folli-

age with cultivation, or defoliation with

herbicides. Because unharvested plants

contribute nothing to the harvest, perma-

nently removing them or cutting them off at

the base during harvest would be a good

field sanitation practice. A potential secon-

dary problem may be the disruption of in-

sect and mite natural enemies in hop yards

(Strong and Croft 1993; James et a/. 2001).

Successful control of hop aphids on

harvested hops would depend on hop grow-

ers over a large area cooperating in a fall

control program. Controls would have to be

applied as soon after harvest as possible and

would need to be extremely effective.

Workers in Idaho developed an area-wide

program to reduce potato leaf roll virus by

reducing the number of green peach aphids,

Myzus persicae (Sulzer), in the spring be-

fore the aphids flew to potatoes (Bishop

1967). They sprayed insecticides on intro-

duced flower and vegetable transplants and

home gardens, and removed the aphid’s

overwintering hosts, peach and apricot

trees. This program was successful in re-

ducing aphids and potato leaf roll virus

when spraying was thorough and well

timed. The small size and isolation of the

potato-growing areas were important fac-

tors in the program’s success.

The hop-growing region of Washington

is isolated from other hop-growing areas, so

perhaps a similar area-wide program could

be effective against the hop aphid. Control-

ling aphids in prune and plum orchards

would be essential. For ornamental trees,

one potential method would be the removal

of Prunus spp. host trees, especially P.

cerasifera. Dixon and Kindlmann (1990)

present theoretical evidence that aphid

abundance 1s directly related to host plant

abundance and the number of hop aphids

caught in suction traps in England and

Washington is related to the abundance of

host plants in the area (Taylor et al. 1979,

Wright et a/. 1995). This suggests that the

hop aphid populations may be susceptible

to manipulations of host plant abundance.

Hymenopterous parasitoids commonly at-

tack hop aphids on Prunus spp. in the

spring (Wright and James 2001). Perhaps

parasitoids and predators could be managed

to reduce the number of spring migrants

flying to hops. Spraying ornamental Prunus

spp. may be effective but could have nega-

tive impacts on natural enemies. Because

the hop aphid can migrate over long dis-

tances (Taylor ef al.1979), any area-wide

program would need to cover a large area to

be effective. To be successful, any alterna-

tive control would have to provide signifi-

cantly superior control, be safer to people

or the environment, or cost less than tradi-

tional methods.

ACKNOWLEDGEMENTS

We thank the landowners for their gen-

erous cooperation and The Washington Hop

Commission for financial support. The sug-

gestions by two anonymous reviewers are

greatly appreciated.

REFERENCES

Bishop, G.W. 1967. A leaf roll virus program in Idaho’s seed potato areas. American Potato Journal 44:

305-308.

Blackman, R.L. and V.F. Eastop. 1984. Aphids on the World’s crops: An identification and information

guide. John Wiley & Sons, Chichester, UK.

Blackman, R.L. and V.F. Eastop. 1994. Aphids on the World’s trees: an identification and information

guide. CAB International, Wallingford, UK.

Born, M. 1968. Beitrage zur Bionomie von Phorodon humuli (Schrank, 1801). Archiv fiir Pflanzenschutz 4:

26 J. ENTOMOL. SOC. BRIT. COLUMBIA 102, DECEMBER 2005

37-52.

Campbell, C.A.M. 1985. Has the damson-hop aphid an alate alienicolous morph? Agriculture, Ecosystems

and Environment 12: 171-180.

Campbell, C.A.M. and E.L. Tregidga. 2005. Photoperiodic determination of gynoparae and males of dam-

son-hop aphid Phorodon humuli. Physiological Entomology 30: 189-194.

Dixon, A.F.G. and P. Kindlmann. 1990. Role of plant abundance in determining the abundance of herbivo-

rous insects. Oecologia 83:281-283.

Eppler, A. 1986. Untersuchungen zur Wirtswahl von Phorodon humuli Schrk. 1. Besiedelte Pflanzenarten.

Anzeiger fiir Schadlingskunde Pflanzenschutz Umweltschutz 59: 1-8.

Hitchcock, C.L. and A. Cronquist. 1973. Flora of the Pacific Northwest: an illustrated manual. University of

Washington Press, Seattle, WA.

James, D.G., T. Price, L.C. Wright, J. Coyle, and J. Perez. 2001. Mite abundance and phenology on com-

mercial and escaped hops in Washington state, USA. International Journal of Acarology 27: 151-156.

Micinski, B. and M. Ruszkiewicz. 1974. Biologia mszycy sliwowo-chmielowe} Phorodon humuli Schr.

Prace Naukowe Insytutu Ochrony Roslin 16: 79-101.

National Agricultural Statistics Service. 1992. 1992 Census of agriculture — county data. United State De-

partment of Agriculture, http://www.nass.usda.gov/.

National Agricultural Statistics Service. 2002. 2002 Census of agriculture — county data. United State De-

partment of Agriculture, http://www.nass.usda.gov/.

Neve, R. A. 1991. Hops. Chapman and Hall, London.

SAS Institute. 1988. SAS/STAT user’s guide, release 6.03. SAS Institute, Cary, NC.

Strong, W.B. and B.A. Croft. 1993. Phytoseiid mites associated with spider mites on hops in the Willamette

Valley, Oregon. Journal of the Entomological Society of British Columbia 90: 45-52.

Taylor, L.R., I.P. Woiwod, and R.A.J. Taylor. 1979. The migratory ambit of the hop aphid and its signifi-

cance in aphid population dynamics. Journal of Animal Ecology 48: 955-972.

Wright, L.C., W.W. Cone, G.W. Menzies, and T.E. Wildman. 1990. Numerical and binomial sequential

sampling plans for the hop aphid (Homoptera: Aphididae) on hop leaves. Journal of Economic Ento-

mology 83: 1388-1394.

Wright, L.C., K.S. Pike, D. Allison, and W.W. Cone. 1995. Seasonal occurrence of alate hop aphids

(Homoptera: Aphididae) in Washington state. Journal of Agricultural Entomology 12: 9-20.

Wright, L.C. and D.G. James. 2001. Parasitoids of the hop aphid (Homoptera: Aphididae) on Prunus during

the spring in Washington state. Journal of Agricultural and Urban Entomology 18: 141-147.

J. ENTOMOL. SOc. BRIT. COLUMBIA 102, DECEMBER 2005

Host preference by Saperda calcarata Say

(Coleoptera: Cerambycidae)

CYNTHIA L. BROBERG' and JOHN H. BORDEN”

ABSTRACT

We conducted five laboratory and one field experiments to examine potential host selec-

tion mechanisms of Saperda calcarata Say in British Columbia. Olfactory bioassays

indicated that female (and possibly male) beetles were attracted to volatiles from leafy

twigs of trembling aspen, Populus tremuloides Michaux. However, wounding of the

bole, ethanol baiting, or both, did not result in significant orientation toward or attack of

trembling aspens in the field. Feeding preferences for trembling aspen were strong for

both sexes in choice bioassays, but in no-choice bioassays, females did not discriminate

between trembling aspen and black cottonwood, P. trichocarpa Torrey & Gray.

Scouler’s willow, Salix scouleriana Barrat in Hooker, was fed upon the least by both

sexes. When diameter of bolts offered as oviposition hosts was equalized, frequency of

Oviposition was similar among the three hosts. Our data suggest that feeding preference

is the predominant mechanism of host selection by S. calcarata.

INTRODUCTION

The poplar borer, Saperda calcarata a

short period of maturation feeding

Say (Coleoptera: Cerambycidae: Lamiinae)

attacks living poplars from the sections

Populus (P. tremuloides Michaux, P. alba

L., P. grandidentata Michaux), Aigeiros (P.

deltoides Bartram, P. fremontii Watson, P.

nigra L. ‘Italica’), and Tacamahaca (P.

angustifolia James, P. balsamifera L., P.

trichocarpa Torrey & Gray) throughout

their range in North America (Hofer 1920,

Baker 1972, Drouin & Wong 1975, Ne-

beker ef al. 1985). The beetle also attacks

poplar hybrids (P. x acuminata) (Hofer

1920) and willows (Baker 1972). Populus

spp. are susceptible from approximately

three years of age (Abrahamson &

Newsome 1972), or 4-5 cm diameter at

breast height (dbh = 1.3 m) (Hofer 1920,

Drouin & Wong 1975, Nebeker ef al.

1985). Saperda calcarata adults reportedly

discriminate among poplar hybrids for feed-

ing (Garland & Worden 1969) and there are

differences in attack rates among P. del-

toides clones (Nebeker et al. 1985).

In British Columbia (BC), S. calcarata

adults emerge in late June to July, undergo

(Linsley 1959) and mate. Females oviposit

into oblong niches chewed in the bark of

host trees. Young larvae mine in the inner

bark and sapwood, then move deeper creat-

ing large, irregular galleries throughout the

sapwood and heartwood (Hofer 1920). Fre-

quently a single tree is repeatedly attacked

forming a ‘brood’ tree. Attacked trees are

identified by their deformed bole, oviposi-

tion scars, sap stains spreading down the

bark, and frass piles at their base. The life

cycle takes three to four years in Canada,

but is probably shorter in the south (Hofer

1920, Peterson 1947, Baker 1972).

Saperda calcarata 1s considered a major

pest of poplars (Solomon 1987) and fre-

quently becomes prevalent within stands

(Bird 1930, Nebeker et al. 1985). Physical

damage to the boles from larval galleries

makes trees susceptible to breakage. Open-

ings from oviposition niches and wood-

peckers lead to increased incidence of

pathogens like Hypoxylon mammatum

(Wahlenberg) Karsten (Graham & Harrison

1954) or Phellinus tremulae (Bondartsev)

' Department of Biological Sciences, Simon Fraser University, Burnaby, BC, V5A 186, Canada

* Current Address: Phero Tech Inc., 7572 Progress Way, Delta, BC, V4G 1E9

28 J. ENTOMOL. Soc. BRIT. COLUMBIA 102, DECEMBER 2005

Bondartsev & Borisov (Hofer 1920) which

girdle the bark or stain and rot the wood,

and attacked trees may be further damaged

by other insects, e.g., Agrilus anxius Gory

or Poecilonota cyanipes (Say) (Hofer

1920).

Because attacks by beetles may be

prevalent in poor sites, e.g., dry slopes

(Hofer 1920, Bird 1930, Morris 1963), or in

decadent hosts (Graham & Harrison 1954),

S. calcarata is assumed to prefer weakened

hosts that remain alive during attack (Hanks

1999). In agreement with this hypothesis,

less attack was observed on P. deltoides

clones from southern provenances which

grew most vigorously (Nebeker ef al.

1985). In contrast, Baker (1972) noted that

brood trees were larger and faster growing

than neighbouring trees, and Abrahamson

& Newsome (1972) concluded there was no

difference in attack level on different qual-

ity sites. Olfaction is generally believed to

play a large role in cerambycid host loca-

tion (Linsley 1961, Hanks 1999, Allison et

al. 2004).

We commonly observe S. calcarata

attack in trembling aspen, P. tremuloides, in

BC, but not in black cottonwood, P. tricho-

carpa, or Willow, Salix spp. Our objectives

were to determine: if olfactory attraction

occurs to trembling aspen, the apparent

preferred host, and if there are different

levels of feeding or oviposition among

these three hosts.

MATERIALS AND METHODS

Saperda Colonies. We collected ca. 2.5

m° S. calcarata-attacked trembling aspen

bolts from trees felled near 70 Mile House,

BC in April or May of 2002 to 2004. Adults

emerged from the caged bolts during June

and July for two successive years. A total of

76 and 101, 46 and 30, and 11 beetles

emerged each year from bolts harvested in

2002, 2003 and 2004, respectively. Timing

of emergence was in agreement with Gar-

land & Worden (1969). Adults were kept

on aspen branches in water in 1.2 x 1.8 x

0.6 m outdoor enclosures until used in bio-

assays. Beetles were used once in any one

type of bioassay, except for feeding bioas-

says in 2002, when tested beetles were re-

turned to the holding cage from which test

subjects were removed.

Plant material. Leafy aspen branches

were collected periodically, mostly from

various interior BC locations, but also from

Burnaby and Maple Ridge on the coast.

Branches were kept with the cut ends in

water at 4 °C, and used in bioassays within

one week. Both Scouler’s willow, Salix

scouleriana Barrat in Hooker, and black

cottonwood branches were collected in Bur-

naby, BC the same day bioassays were per-

formed. In total, four to five genotypes of

each species were tested.

Olfaction experiments. In 2003 and

2004, responses to volatiles were investi-

gated in the laboratory using a still air ol-

factometer (Figure 1). Randomly assigned

treatment and control jars contained either a

small jar of water with a small, leafy branch

of aspen, or just water, respectively. The

arena ceiling was a sheet of clear Lexan

(GE Polymershapes, Coquitlam, BC). A

video camera was positioned above the

arena. To induce the photopositive beetles

to approach the stimuli, a fluorescent light

was placed under the platform between the

two jars. The entire apparatus was covered

with a black cloth. Three to five adults of a

single sex taken directly from the holding

cage were placed into each arena. Presence

of beetles in the concentric rings above the

treatment or control jars were determined at

30 sec intervals for 2 h. Fourteen trials were

recorded. Feeding damage to the perforated

centres above control and treatment stimuli

was assessed for both recorded and unre-

corded trials. Jars were washed and the pa-

per covering replaced before each assay.

Assays commenced any time between 0800

and 2400 h.

A field experiment was set up in a trem-

bling aspen grove near Sabiston Lake,

northeast of Savona, BC, on 28 June, 2002.

Apparently healthy trees, spaced approxi-

mately 20 m apart, received one of four

treatments in a randomized block design (n

= 14): 4 axe cuts on opposite sides of the

J. ENTOMOL. Soc. BRIT. COLUMBIA 102, DECEMBER 2005

)

Figure 1. Schematic of olfactometer used to test olfactory responses of S. calcarata to trem-

bling aspen leaf volatiles. A 60 cm diameter circular arena consisted of a black cylinder 60 cm

high (A) placed on a white coroplast (GE Polymershapes, Coquitlam, BC) platform (C) with two

holes, 5.5 cm in diameter with centres 28 cm apart. This platform was covered with white paper

(B) perforated 41 times with a pin in a uniform, radial pattern above the openings. Three concen-

tric rings 5.5, 12.5, and 19.5 cm diameter were drawn on the paper (B) above each hole. The

platform (C) rested on glass cookie jars (aperture 11 cm diameter) (D), centred below each hole.

tree at ca. 1.5 m; ethanol bait stapled at ca.

2 m; 4 axe cuts plus ethanol bait; or no

treatment. Ethanol is a ubiquitous kairo-

monal indicator of stressed trees (Kelsey &

Joseph 1998 and references therein). The

basal 3 m of the trees were examined for

Oviposition in 2003 and 2004.

Feeding bioassays. Choice and no-

choice feeding bioassay experiments were

performed to investigate feeding prefer-

ences of S. calcarata among trembling as-

pen, black cottonwood and Scouler’s wil-

low. Three small branches, each with three

to five leaves, were placed in water-filled

vials inside 17x16x12 cm plexiglass boxes.

A single S. calcarata adult was allowed to

freely feed on plant material overnight. No-

choice bioassays contained one of the three

potential hosts, and choice bioassays con-

tained one branch of each. Before an assay,

the leaves were traced onto paper. After the

bioassay ended, leaves were attached to

their traced counterparts, scanned, and the

leaf area consumed was quantified using

Scion Image software (Scion Corporation,

Frederick, Maryland). A total of 17 choice

and 14 no-choice bioassays were performed

during July of 2002 and 2003.

Oviposition bioassays. In 2002 we

tested oviposition by S. calcarata in hold-

ing cages on eight freshly cut bolts of vary-

ing diameter of each of the above three spe-

cies. In 2003, two apparently healthy trem-

bling aspen, black cottonwood, and

30 J. ENTOMOL. SOC. BRIT. COLUMBIA 102, DECEMBER 2005

Scouler’s willow trees of similar diameter

were felled on 1 and 18 July 2003, bucked

and transported to SFU where they were

kept refrigerated until needed. Six to eight

beetle pairs were placed in 13 outdoor

cages, 90 x 90 x 90 cm, with one randomly

positioned bolt of each species and a central

water jar containing leafy trembling aspen

branches for seven days. Bolts were ca. 50

cm in length, and diameters taken from

their midpoint. Oviposition was determined

in both experiments by opening all niches

cut in the bark.

Statistical analyses. In all cases a =

0.05. For olfaction bioassays, one-tailed

paired t-tests were used to determine if the

frequency of observations in each of the

concentric rings was greater over treatment

than control jars. Chi-square tests were per-

formed for each sex to compare the fre-

quency of feeding damage above treatment

vs. control stimuli against the null hypothe-

sis of no discrimination between stimuli.

Data from feeding and oviposition bioas-

says were transformed by x” and log(x+1),

respectively, to correct for non-normality

and heteroskedasticity, then analyzed as

randomized complete blocks by ANOVA

with PROC GLM (SAS Institute 1990).

Because it was not possible to perform all

no-choice feeding assays at the same time,

the analysis included trial date and host

species effects. The final model did not

include interaction effects. Multiple com-

parisons were performed with REGWQ

(SAS Institute 1990).

RESULTS

Olfaction experiments. There were no

differences in the occurrence of beetles in

the middle and outside rings bordering the

perforated area above treatment (i.e. trem-

bling aspen leaves) and control stimuli for

both sexes (middle ring, females, t = 0.69,

P = 0.26, males, t = -0.80, P = 0.23; outer

ring, females, t = 0.76, P = 0.24, males, t =

0.95, P = 0.19). Females (but not males)

were present more frequently above the

treatment than the control stimulus (Table

1). Often, females and males fed on the

perforated paper directly above the treat-

ment stimulus, but in one instance females

fed above the control as well (Table 1).

Females also chewed curvilinear patterns in

the paper covering the arena floor that were

reminiscent of oviposition niches, but no

eggs were found.

Very few attacks were found on trem-

bling aspen treated to release host volatiles

(axe cuts) or baited with ethanol. A total of

8 oviposition chambers in two replicates

were found: 1 niche on an ethanol-treated

tree; | and 3 niches on two axe-cut trees; 2

and | niches on two trees with both treat-

ments and none on control trees. None of

these niches developed into successful lar-

val galleries.

Feeding bioassays. We observed feed-

ing on both the petioles and leaves as did

Garland & Worden (1969), but only quanti-

fied the more abundant foliar damage.

When given a choice (Figure 2), both sexes

clearly preferred trembling aspen over both

black cottonwood and Scouler’s willow

(females F,3, = 41.15, P < 0.0001; males

F339 = 42.68, P < 0.0001). There were also

significant differences in feeding in the no-

choice experiment (females F>34 = 20.53, P

< 0.0001; males F,33 = 26.96, P < 0.0001).

However, females accepted trembling aspen

and black cottonwood equally, and males

fed on black cottonwood more vigorously

than on Scouler’s willow (Figure 2).

Oviposition bioassays. When bolt di-

ameter was not controlled in 2002, S. cal-

carata females oviposited preferentially in

trembling aspen and black cottonwood, the

species with the largest diameter bolts

(Table 2). When bolt diameter was equal-

ized in 2003, there was no preference in

Oviposition among the three host species

(Table 2).

We observed some larvae feeding in the

bark of all three species, but they did not

survive long as bark quality deteriorated

rapidly because of infection by Cytospera

chrysosperma (Persoon: Fries) Fries, distin-

guished by characteristic orange conidial

tendrils (Callan 1998).

J. ENTOMOL. SOC. BRIT. COLUMBIA 102, DECEMBER 2005

Table 1.

Comparison of behavioural activity by male and female S. ca/carata within perforated centres

of arena floor above treatment (i.e. trembling aspen leaves) and control jars in the still-air ol-

factometer.

Observations Females Males

Observations of beetles within arena circle circumscribing perforated

area above treatment or control stimulus, 30 sec intervals for 2 h.

no. replicates 7 7

mean no. observations + SE

treatment stimulus 5544185 41.948.8

control stimulus 2032108 AWOaISS

t-value Lo? 0.00

Probability 0.047 0.50

Observations of feeding on perforated area of arena floor above

treatment or control stimulus

no. replicates 8 1]

no. times most feeding above treatment stimulus ‘i 6

no. times most feeding above control stimulus 0 0

no. times no feeding damage observed ] 5

Chi-square value 7.00 6.00

Probability 0.008 0.01

'In one trial feeding damage was observed above both stimuli, but damage was much greater

over the treatment stimulus.

DISCUSSION

Our results indicate that S. calcarata can

locate potential hosts by olfaction, can dis-

criminate among tree species through gusta-

tory cues, and may reject trees for oviposi-

tion if their diameter is too small.

Females were attracted to the volatiles

from leaf-bearing twigs of trembling aspen

in an arena olfactometer (Table 1). The fact

that males chewed the paper above treat-

ment but not control stimuli suggests they

too are attracted to host volatiles. In the

field experiment, we had hypothesized that

S. calcarata brood trees would produce

ethanol, and possibly other metabolites

caused by wounding, and that the combina-

tion would be attractive. The positive re-

sponse to leafy twigs, and the failure to

induce significant attack on trees that were

wounded, ethanol-baited, or both, suggests

that initial orientation is to volatiles from

leaves on which adults feed. Oviposition

tends to be located in the upper parts of the

bole beneath the canopy (Peterson 1947),

requiring littke movement by feeding bee-

tles. Similarly, in an unpublished study con-

ducted by C.L. Broberg and R. Gries

(SFU), 15 antennally-active volatiles from

the bark of trembling aspen were identified

by coupled gas chromatographic electro-

antennographic detection analysis. How-

ever, in six field-trapping experiments, no

S. calcarata were captured to various par-

tial or complete blends of these compounds.

These results may indicate that S. calcarata

use leaf volatiles in orientation toward suit-

able hosts during flight.

The lack of a strong olfactory response

by S. calcarata is not surprising. As a spe-

cialist of weakened hosts that may support

multiple generations on the same tree

(Hanks 1999), S. calcarata is not likely

subject to strong selection pressure to adapt

to finding new hosts. Furthermore, Populus

spp. are pioneer species and often occur in

locally abundant populations. Thus emer-

gent S. calcarata may not need to disperse

a2 J. ENTOMOL. SOc. BRIT. COLUMBIA 102, DECEMBER 2005

CHOICE EXPERIMENT

Females

Mean leaf area consumed (cm2) + SE

TA CW Wil

Host species

Figure 2. Leaf area consumed by female and male S. calcarata when presented with the three

potential hosts simultaneously (choice experiment) or separately (no-choice experiment). Bars

within an experiment and sex with the same letter are not significantly different, REGWQ test,

P <0.05. TA = trembling aspen, CW = black cottonwood, Wil = Scouler’s willow.

long distances to find a suitable host. In

contrast, stressed hosts which are moribund

and can only support one generation of bee-

tle (Hanks 1999) are often rare and/or

patchy in distribution. Cerambycid special-

ists on these hosts have evolved strong

long-distance response mechanisms _ that

often involve orientation to host volatiles

from recently downed or injured trees,

smoke volatiles from burned trees, or

pheromones produced by secondary bark

beetles pheromones (Allison et al. 2004).

Although these cerambycids mate and ovi-

posit on these newly found hosts, they en-

gage in maturation feeding on healthy trees

(Hanks 1999). Weakened host specialists

like S. calcarata can use a single individual

for all functions; thus even the malformed

emergent adults incapable of flight ob-

served by us and others (Peterson 1947;

Drouin & Wong 1975), can experience re-

productive success without long range dis-

persal and olfactory orientation to suitable

hosts.

Both sexes clearly preferred trembling

aspen in choice feeding bioassays and re-

J. ENTOMOL. SOC. BRIT. COLUMBIA 102, DECEMBER 2005 33

Table 2.

Comparison of oviposition by S. calcarata on bolts from three different hosts when bolt diame-

ters were unequal or similar.

Experimental re Bolt diameter (cm) Mean no.

description P Range Mean+SE' ovipositions + SE

Bolt diameters unequal Trembling aspen 6.3-13.2 954+0.8a 21029.6b

(2002) Black cottonwood 8.9-16.7 12.2+0.9a 27.5494a

Scouler’s willow 3.2-5.55 46+0.3b 0.4+0.2¢c

Bolt diameters similar Trembling aspen 7.8-17.1 13.5+0.6a 44+2.1a

(2003) Black cottonwood 8.1-17.3 13.14+0.6a 8.0+3.2a

Scouler’s willow 9.2-17.0 12.6+0.6a 8.342.8a

'Means within an experiment and column followed by the same letter are not significantly

different, REGWQ test, P < 0.05. ANOVA statistics as follows: 2002 bolt diameter F>). =

33.83, P < 0.0001 ; oviposition Fg = 21.17, P < 0.0001; 2003 bolt diameter Fy 54 = 2.57, P =

0.10; oviposition F254 = 1.17, P= 0.33.

jected other species, but in the no-choice

bioassays, females did not discriminate

between trembling aspen and black cotton-

wood, and males accepted black cotton-

wood more than Scouler’s willow (Figure

2). Females may have higher nutritive re-

quirements than males and therefore cannot

afford to be as selective. Preference for

trembling aspen could be a result of local

adaptation to a species that comprises 4.6

times more wood volume in BC than black

cottonwood (BC Ministry of Forests 1998).

Saperda calcarata is mobile enough to

sample numerous trees before finding one

that is suitable for feeding.

There are few records in BC of S. cal-

carata attack on Salix spp. In Saskatche-

wan, however, both Salix and Populus spp.

were reported to be “readily eaten” by

adults (Peterson 1947). Thus, there could be

host-related ecotypes in different regions of

North America.

The lack of discrimination between

hosts for oviposition when trembling aspen

leaves were available for maturation feed-

ing, and diameters of bolts from the three

species were equalized, indicates that host

volume is more important than species for

larval feeding and development (Table 2).

Oviposition in large-diameter hosts would

be adaptive in ensuring that most hosts did

not suffer breakage during the three to four

years required for larval development.

In conclusion, lack of evidence for long-

range olfactory orientation to new hosts,

correlation between S. calcarata incidence

in BC and gustatory preferences, high mo-

bility, and lack of discrimination between

hosts for oviposition, suggest that feeding

preference constitutes the predominant

mechanism of host selection by S. calcarata

in BC.

ACKNOWLEDGEMENTS

We thank Nicole Vander Wal, Ashley

Mohle, and James Inkster for assistance;

and Leland Humble for review of this

manuscript. The research was supported by

Abitibi Consolidated Inc., Ainsworth Lum-

ber Co. Ltd., B.C. Hydro and Power Au-

thority, Bugbusters Pest Management Inc.,

Canadian Forest Products Ltd., Gorman

Bros. Ltd., International Forest Products

Ltd., Louisiana-Pacific Canada Ltd., Man-

ning Diversified Forest Products Ltd., Muil-

lar-Western Forest Products Ltd., Phero

Tech Inc., Scott Paper Ltd., Slocan Forest

Products Ltd., Tembec Forest Industries

Ltd., TimberWest Forest Ltd., Tolko Indus-

tries Ltd., West Fraser Mills Ltd., Western

Forest Products Ltd., and Weyerhaeuser

Canada Ltd.

34 J. ENTOMOL. SOc. BRIT. COLUMBIA 102, DECEMBER 2005

REFERENCES

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tations. Forest Science 18: 231-232.

Allison, J.D., J.H. Borden, and S.J. Seybold. 2004. A review of the chemical ecology of the Cerambycidae

(Coleoptera). Chemoecology 14: 123-150.

Baker, W.L. 1972. Saperda calcarata, pp. 185-187. /n Eastern Forest Insects, USDA Forest Service Miscel-

laneous Publication No. 1175. USDA Forest Service, Washington, DC.

BC Ministry of Forests. 1998. Forest Inventory Reporting, Provincial Summary 1998. http://

srmwww.gov.be.ca/tib/reports/index.html.

Bird, R.D. 1930. Biotic communities of the aspen parkland of central Canada. Ecology 11: 356-442.

Callan, B.E. 1998. Diseases of Populus in British Columbia: A diagnostic manual. Natural Resources Can-

ada, Canadian Forest Service, Pacific Forestry Centre, Victoria, BC.

Drouin, J.A. and H.R. Wong. 1975. Biology, damage and chemical control of the poplar borer (Saperda

calcarata) in the junction of the root and stem of balsam poplar in western Canada. Canadian Journal of

Forest Research 5: 433-439.

Garland, J.A. and H.A. Worden. 1969. Feeding and mating of the longhorn beetle, Saperda calcarata Say.

(Coleoptera: Cerambycidae). The Manitoba Entomologist 3: 81-84.

Graham, S.A. and R.P. Harrison. 1954. Insect attacks and hypoxylon infections in aspen. Journal of Forestry

52: 741-743.

Hanks, L.M. 1999. Influence of the larval host plant on reproductive strategies of cerambycid beetles. An-

nual Review of Entomology 44: 483-505.

Hofer, G. 1920. The aspen borer and how to control it. USDA Farmer's Bulletin 1154. US Department of

Agriculture, Washington, DC.

Kelsey, R.G. and G. Joseph. 1998. Ethanol in Douglas-fir with black-stain root disease (Leptographium

wageneri). Canadian Journal of Forest Research 28: 1207-1212.

Linsley, E.G. 1959. Ecology of Cerambycidae. Annual Review of Entomology 4: 99-138.

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

Publications in Entomology 18: 1-135.

Morris, R.C. 1963. Trunk borers in cottonwood. Mississippi Farm Research 26: 8.

Nebeker, T.E., J.J. Schmitt, J.D. Solomon, and C.R. Honea. 1985. Clonal resistance to and incidence of the

poplar borer in southern cottonwood plantations, pp. 247-251. Jn Shoulders, E. (ed.), Proceedings of the

Third Biennial Southern Silvicultural Research Conference (1984), USDA Forest Service General Tech-

nical Report SO-54. US Department of Agriculture, Atlanta, GA.

Peterson, L.O.T. 1947. Some aspects of poplar borer, Saperda calcarata Say, (Cerambycidae) infestations

under parkbelt conditions. Report of the Entomological Society of Ontario 78: 56-61.

SAS Institute Inc. 1990. SAS/STAT user's guide, version 6. SAS Institute Inc., Cary, NC.

Solomon, J.D. 1987. Management of insect pests in cottonwood nurseries and plantations, pp. 41-44. Jn

Proceedings, Poplar Councils of the United States and Canada Joint Meeting (1987) New York/Ontario,

USDA Forest Service Southern Forest Experiment Station. New Orleans, LA.

J. ENTOMOL. Soc. BRIT. COLUMBIA 102, DECEMBER 2005

Yellowjacket Wasps (Hymenoptera: Vespidae) Trapped in

Alaska with Heptyl Butyrate, Acetic Acid and Isobutanol

PETER J. LANDOLT', ALBERTO PANTOJA

and DARYL GREEN!

ABSTRACT

Eight species of vespine wasps were captured in traps near Fairbanks, Delta Junction

and Palmer, Alaska, during 2003 and 2004. These were Vespula vulgaris L., V. acadica

(Sladen), V. consobrina

(Saussure), V. rufa

(L.)(=intermedia [Buysson]),

Dolichovespula maculata (L.), D. arenaria (F.), D. norwegica (F.)(=albida [Sladen]),

and D. norvegicoides (Sladen). Workers and males of V. vu/garis were captured primar-

ily in traps baited with the combination of acetic acid and isobutanol. Workers of V.

acadica, V. consobrina, and V. rufa were captured primarily in traps baited with heptyl

butyrate. Queens and workers of D. maculata were captured primarily in traps baited

with acetic acid, or acetic acid plus isobutanol. The small numbers of D. arenaria, D.

norvegicoides, and D. norwegica captured did not permit treatment comparisons. Sea-

son-long trapping indicated a presence of V. acadica, V. consobrina, and V. rufa work-

ers from late June through July, D. maculata from early July into early August, and V.

vulgaris from late July to early September. The earliest wasps captured were queens of

V. vulgaris and D. maculata in late May, while the latest wasp captured was a worker of

V. vulgaris the first week of October, in Palmer.

Key Words: social wasps, Vespinae, Vespula, Dolichovespula, trapping, attractant

INTRODUCTION

There is little information available on

the abundance, distribution, or seasonality

of social wasps (Vespidae, Vespinae) in

Alaska, despite their likely widespread and

recurring pest status. Many species of yel-

lowjackets, which belong to the genera

Vespula and Dolichovespula (Greene and

Caron 1980), are often stinging hazards to

people, pets, and livestock. An early report

from the Harnman Alaska Expedition

(Kincaid 1900) listed only two species:

Dolichovespula norwegica (F.) (as Vespa

marginata Kirby) from Kukak Bay, and D.

arenaria (F.) (as Vespa borealis Kirby)

collected in Sitka. Distributions given for

vespid wasps of North America by Miller

(1961), Wagner (1978), Akre et al. (1980),

Eck (1984), and Carpenter and Kojima

(1997) indicate that D. norwegica (= albida

Sladen), D. adulterina (Buysson) (= arctica

Rohwer), D. arenaria, D. norvegicoides

(Sladen), D. alpicola Wagner, D. maculata

(L.), Vespula vulgaris (L.), V. acadica

(Sladen), V. austriaca (Panzer), and V. rufa

L. [= intermedia (Buysson)]| are present in

Alaska. The pest status of V. vulgaris, the

common yellowjacket, in Alaska is indi-

cated by Shippey (1994) (cited in Barnes er

al. 1996).

Chemical attractants useful in trapping

and monitoring yellowjacket wasps include

heptyl butyrate (Davis et al. 1969), and

acetic acid plus isobutanol (Landolt 1998).

Heptyl butyrate is a strong attractant for

Vespula pensylvanica (Saussure) (Davis ef

al. 1973), and also attracts significant num-

bers of V. squamosa (Drury) and some

members of the V. rufa species group: V.

atropilosa (Sladen), V. acadica, V. conso-

brina (Saussure), and V. vidua (Saussure)

(Grothaus et al. 1973, MacDonald ef al.

1973, Howell et al. 1974, Reed and Landolt

‘USDA-ARS, Yakima Agricultural Research Station, 5230 Konnowac Pass Road, Wapato, WA 98951

? USDA-ARS Subarctic Agricultural Research Unit, Fairbanks, AK 99775

36 J. ENTOMOL. SOc. BRIT. COLUMBIA 102, DECEMBER 2005

2002, Landolt et a/. 2003). Acetic acid plus

isobutanol is attractive to some members of

the V. vulgaris species group: V. germanica

(F.), V. pensylvanica, V. vulgaris, and V.

maculifrons (Buysson), as well as to V.

squamosa, Vespa crabro L. and several

species of Polistes (Landolt 1999; Landolt

et al. 2001). Acetic acid or isobutanol alone

are also weakly attractive to some species

of social wasps (Landolt et a/. 1999, Reed

and Landolt 2002), and acetic acid was co-

attractive with heptyl butyrate for trapping

V. pensylvanica (Landolt 1998).

In this study, we sought to determine

and compare the responses of species of

social wasps in Alaska to heptyl butyrate,

acetic acid, and isobutanol, particularly to

test the hypothesis that yellowjackets in the

genus Dolichovespula and the V. vulgaris

species group are primarily attracted to ace-

tic acid and isobutanol, while yellowjackets

in the V. rufa species group are primarily

attracted to heptyl butyrate. We also sought

to confirm information on the species of

social wasps that are present in Alaska and

determine the seasonal pattern of abun-

dance of species that are likely to be pestif-

erous. We report here the results of trapping

tests that provide significant information on

yellowjacket wasp responses to chemical

lures, and on the seasonality of several spe-

cies of Vespinae in Alaska.

MATERIALS AND METHODS

Dome or Trappitt® traps (Gempler’s,

Belleville, Wisconsin, USA) were used to

capture attracted wasps. These traps are

pear-shaped with clear plastic tops and

opaque yellow bottoms within which is

placed a drowning solution. Wasps enter

the trap through the invaginated bottom of

the trap. Attractants were dispensed from

polypropylene vials with holes in the lid for

chemical release. Each vial contained

chemical attractant on cotton balls. Vial

sizes, active ingredient load amounts, and

hole diameters selected were based on re-

sults of previous studies of wasp responses,

as well as chemical release rates from vials

under laboratory conditions. At ambient

laboratory temperature (22.5 °C), estimated

rate of release of compounds from vials

with 3 mm diam holes is 200 ug heptyl bu-

tyrate per hour (Landolt ef a/. 2003), 8.2 mg

acetic acid per hour (Landolt and Alfaro

2001), and 10 mg isobutanol per hour

(using gravimetric methods reported in

Landolt and Alfaro 2001). Vials were sus-

pended at the top of the inside of the trap.

Traps also contained 200 to 300 ml of a

drowning solution which was 0.125% un-

scented detergent and 2% boric acid in wa-

ter. Traps were placed a minimum of 20 m

apart, and were placed at a height of 1.0 to

1.5 m on vegetation or on fences. Traps

were checked once per week, at which time

the drowning solution was replaced, and

lures were replaced every month, which

would be before the attractant in the dis-

penser was depleted.

2003 Trapping Test. Five sets of traps

were placed on the campus of the Univer-

sity of Alaska, Fairbanks North Star Bor-

ough, Alaska, during the second week of

July, 2003. Trap treatments at each location

were: 1) an unbaited trap as a control, and

traps baited with 2) acetic acid, 3) isobu-

tanol, 4) acetic acid plus isobutanol, 5) hep-

tyl butyrate, and 6) acetic acid plus heptyl

butyrate. Each chemical (10 ml load) was

dispensed from its own 15 ml vial with a 3

mm diam hole. Trap sites were in the vicin-

ity of forested tracts, agricultural land, and

a horticultural garden.

2004 Trapping Test. Traps were set up

in early May at three locations, as pairs of

traps baited with two chemical attractants:

heptyl butyrate and acetic acid plus isobu-

tanol. Heptyl butyrate (10 ml load) was

dispensed from a 15 ml vial with a 3 mm

hole and acetic acid plus isobutanol was

provided as a mixture of the two com-

pounds (10 ml load) in a single 15 ml vial

with a 6 mm diameter hole. Four pairs of

traps were placed on the main campus of

the University of Alaska, Fairbanks, five

pairs of traps were placed at the University

of Alaska field site at Delta Junction,

J. ENTOMOL. SOC. BRIT. COLUMBIA 102, DECEMBER 2005

Southeast Fairbanks Borough, and four

pairs of traps were placed at the University

of Alaska field site at Palmer, Matanuska-

Susitna Borough. Trap sites were near both

forested and agricultural lands. Traps at

Fairbanks and Palmer were maintained until

the third week of September and traps at

Delta Junction were maintained until the

first week of October.

Insects captured in traps were placed in

pre-labeled plastic locking freezer bags. A

separate bag was used for each trap and for

each day the trap was checked. These were

stored in a freezer until bag contents were

analyzed. Descriptions, illustrations, and

keys in Miller (1961), Wagner (1978), Akre

et al. (1980), and Eck (1984) were used to

identify captured vespine wasps. Taxonomy

used here follows that of Carpenter and

Kojima (1997). Voucher specimens are

deposited in the James Entomological Col-

lection at Washington State University,

Pullman, WA, and with the USDA, ARS

Subarctic Agricultural Experiment Station

in Fairbanks, AK.

For each species, means for wasps cap-

tured per trap per week in 2003 were com-

pared between chemical attractant treat-

ments using ANOVA and Tukey’s test

(DataMost 1995) to determine differences

among means. Similar data for 2004 were

compared using a paired ¢-test. For develop-

ing seasonality profiles for each species,

numbers of wasps captured in each trap per

week were averaged for the traps at each

site. Unless stated otherwise, data analyses

and results are for worker wasps.

RESULTS

The most abundantly trapped wasp in

both years and at all three study locations

was V. vulgaris. In 2003, 508 worker V.

vulgaris were captured in traps. In 2004, 4

queens, 1825 workers, and 36 males of V.

vulgaris were captured. In 2003, V. vulgaris

workers were primarily in traps baited with

the combination of acetic acid and isobu-

tanol, with no or few wasps in unbaited

traps and traps baited with acetic acid, iso-

butanol, heptyl butyrate, or acetic acid plus

heptyl butyrate (Table 1). In 2004, V. vul-

garis workers again were primarily in traps

baited with acetic acid plus isobutanol, with

nearly none in traps baited with heptyl bu-

tyrate (Table 2). The same pattern was seen

for V. vulgaris males in traps (Table 2). The

four V. vulgaris queens were captured in

traps (in late May and early June) baited

with acetic acid plus isobutanol. Workers

were captured between mid June and early

October, and males from late July into late

September. Workers were most abundantly

trapped from mid July into late August

2004 (Figure 1A).

In 2003, 9 worker V. acadica were cap-

tured in traps, primarily in traps baited with

heptyl butyrate (Table 1). In 2004, the 42

worker and 4 queen V. acadica captured

were all in traps baited with heptyl butyrate

Table 1.

Mean + SE numbers of wasps captured per trap, for unbaited traps (CONTROL), and for traps

baited with acetic acid (AA), isobutanol (IB), heptyl butyrate (HB), acetic acid plus isobutanol

(AAIB), and acetic acid plus heptyl butyrate (AAHB). Fairbanks, Alaska, 2003.

een CONTROL AA AAIB HB AAHB

V. vulgaris 5.0+1.8a 8.0+43a 11.24£5.6a 60.0432.9b 16+14a 15.8+10.0a

D. maculata 0.0+0.0a 12.2+7.1lc 1.640.7ab 7.6+43.1lbc 0.6+0.6a 4.8+42.3ab

V.acadica 0.0£0.0a 0.0+0.0a 0.0+0.0a 0.0+0.0a 1.6+0.8b 0.2+0.2a

'For each species, means followed by the same letter are not significantly different at P<0.05

by Tukey’s test. N= 5.

38 J. ENTOMOL. Soc. BRIT. COLUMBIA 102, DECEMBER 2005

Table 2.

Mean + SE numbers of wasps captured per trap, for traps baited with heptyl butyrate (HB) and

for traps baited with acetic acid plus isobutanol (AAIB). Fairbanks, Delta Junction and Palmer,

Alaska, 2004.

Wasps’ HB AA/IB

V. vulgaris workers 0.46 + 0.27a 140.18 + 46.10b

V. vulgaris males 0.07 + 0.07a 3.08 + 0.87b

D. maculata workers 0312 0.l7a 33.92 215.176

D. maculata queens 0.00 + 0.00a 2.022 0772b

V. acadica workers 3.54+0.91b 0.00 + 0.00a

V. consobrina workers 1.380.536 0.00 + 0.00a

V. rufa workers 1.54 + 0.42b 0.00 + 0.00a

'For each species, means followed by the same letter are not significantly different at P<0.05

by a paired /-test. N = 13.

(Table 2). Three of the queens were cap-

tured in late June and one in mid Septem-

ber. Workers were captured from mid June

to mid September 2004 (Figure 1B).

In 2003, only three V. consobrina and

one V. rufa were captured; all were workers

in traps baited with heptyl butyrate. In

2004, 17 V. consobrina and 20 V. rufa

workers were captured, all in traps baited

with heptyl butyrate (Table 2). Vespula

consobrina workers were captured in traps

from 7 July to 3 August in Delta Junction

and from 12 June to 19 July in Fairbanks.

Worker V. rufa were captured in traps from

29 June to 27 July in Delta Junction, from

12 June to 19 July in Fairbanks, and two

were in traps on 5 August in Palmer.

In 2003, 134 worker D. maculata were

captured. Numbers of bald-faced hornets in

traps baited with acetic acid, and with ace-

tic acid plus isobutanol, were significantly

greater than in unbaited traps (Table 1). In

2004, 445 worker and 27 queen D. macu-

lata were captured. Most were captured in

traps baited with acetic acid plus isobu-

tanol. In both years, very few were captured

in traps baited with heptyl butyrate (Table

2). No male D. maculata were captured in

these traps. Queens were captured from mid

May to early June, and workers from late

June into late August (Figure 1C).

In this study, too few D. arenaria (9) D.

norvegicoides (2), or D. norwegica (2)

wasps were captured for any statistical

analysis, while no D. alpicola, D. adul-

terina, or V. austriaca wasps were captured.

DISCUSSION

The species captured in traps during this

study in the vicinities of Fairbanks, Delta

Junction, and Palmer, Alaska, vary some-

what from those species reported by Miller

(1961), Akre et al. (1980), Eck (1984) and

Carpenter and Kojima (1997). Vespula con-

sobrina were captured in traps at two of the

three sites (Fairbanks and Delta Junction),

despite its indicated absence from most of

Alaska by both Miller (1961) and Akre ef

al. (1980). The nearest collection locations

in those references are in the southernmost

Alaska panhandle, northern British Colum-

bia, and southeastern Yukon. The absence

of D. adulterina, D. alpicola, and V. aus-

triaca in traps could have been the result of

their absence in the areas trapped, or a lack

of response to the lures.

The seasonal patterns of wasp captures

in traps indicate a broad period during

which they could be pestiferous; from early

July to early September. Of most interest as

a pest is the common wasp V. vulgaris, be-

cause of its abundance and its scavenging

J. ENTOMOL. SOC. BRIT. COLUMBIA 102, DECEMBER 2005 39

140

2 -

VESPULA VULGARIS

84 -

56 +

WASPS/TRAP/WEEK

28 OF

ae / : L)

Tl a -5=6 gen

OE eta —s

10 17 24 31 7 14 21 28 5 12 19 26 2 9 16 23 30 6 13 20 27 4

MAY JUNE JULY AUG. SEPT.

4.00

3.20

VESPULA ACADICA

2.40

1.60

WASPS/TRAP/WEEK

0.80

0.00

10 17 24 31 7 14 21 28 5 12 19 26 2 9 16 23 30 6 13 20 27 4

MAY JUNE JULY AUG. SEPT.

40 F-

DOLICHOVESPULA MACULATA

30 |

WASPS/TRAP/WEEK

10 17 24 31 7 14 21 28 5 12 19 26 2 9 16 23 30 6 13 20 27 4

MAY JUNE JULY AUG. SEPT.

Figure 1. Mean + SE numbers of female V. vulgaris (A), V. acadica (B) and D . maculata (C)

in traps baited with acetic acid and isobutanol, through the 2004 field season. Lines do not

imply dependence of data. Sites are Fairbanks (open triangles), Delta Junction (filled squares),

and Palmer (open circles).

40 J. ENTOMOL. SOC. BRIT. COLUMBIA 102, DECEMBER 2005

behavior which bring it into frequent con-

tact with people (Akre eft a/. 1980). The

bald-faced hornet, D. maculata, may also

be pestiferous in Alaska due to its abun-

dance during July and August. Other spe-

cies of wasps captured, such as V. acadica,

and V. consobrina, are less likely to be pes-

tiferous because they are not known for

scavenging habits, occur in smaller colo-

nies, and do not occur in high densities,

compared to species such as V. vulgaris

(Akre ef al. 1980).

The patterns of responses of wasps in

Alaska to different chemical attractants are

consistent with results from trapping studies

in other areas of North America. The three

members of the V. rufa species group (V.

acadica, V. consobrina, and V. rufa) were

attracted to heptyl butyrate, and were not

captured in significant numbers in traps

baited with acetic acid plus isobutanol. This

pattern was seen in Washington State,

where V. atropilosa, a V. rufa species group

member, was attracted to heptyl butyrate

and not to acetic acid plus isobutanol

(Landolt 1998), and in Michigan, where V.

consobrina and V. vidua, both V. rufa spe-

cies group members, exhibited the same

response pattern (Reed and Landolt 2002).

In the present study, the only member of the

V. vulgaris species group present was V.

vulgaris. Unlike species in the V. rufa

group, it was attracted to acetic acid plus

isobutanol and not to heptyl butyrate. This

pattern matches results of earlier studies

(Landolt 1998, Landolt et al. 1999, Reed

and Landolt 2002), where V. flavopilosa

Jacobson, V. germanica, V. maculifrons,

and V. vulgaris, all V. vulgaris species

group members, were trapped with acetic

acid plus isobutanol and not with heptyl

butyrate. Vespula pensylvanica (another V.

vulgaris group member), however, is

clearly attracted to both lures (Landolt

1998). The response by D. maculata to ace-

tic acid plus isobutanol and lack of a re-

sponse to heptyl butyrate is consistent with

earlier studies. In Maryland and in western

Washington (Landolt et al. 2001), as well

as in Michigan (Reed and Landolt 2003), D.

maculata workers were trapped with acetic

acid, but more so to acetic acid plus isobu-

tanol. In this study, numbers of workers of

D. maculata captured in traps baited with

acetic acid plus isobutanol were not signifi-

cantly higher than with acetic acid alone.

The small numbers of workers of D.

arenaria, D norvegicoides, and D. norwe-

gica trapped here are not suitable for statis-

tical analyses, and indicate either a very

weak response to the lures, or very low

population densities.

ACKNOWLEDGEMENTS

Technical support was provided by T.

Adams, L. Defoliart, D. Hall, K. Maher, J.

Malapanis, and R. Torgerson in Alaska and

by J. MacKenzie in Yakima, Washington.

This work was supported in part by a Coop-

erative Research and Development Agree-

ment with Sterling International, Inc. of

Spokane, Washington.

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Barnes, B.M., J.L. Barger, J. Seares, P.C. Tacquard, and G.L. Zuercher. 1996. Overwintering in yellow-

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J. ENTOMOL. Soc. BRIT. COLUMBIA 102, DECEMBER 2005

Redescription of Haliplus dorsomaculatus

(Coleoptera: Haliplidae) with a New Synonymy

and Comments on Habitat and Distribution

REX D. KENNER!

ABSTRACT

Adults of Haliplus dorsomaculatus Zimmermann are redescribed including a discussion

of male and female morphological characters. The species ranges from southern British

Columbia east to western Montana and south to northeastern Utah and northern Califor-

nia; there is little geographic variation. Its preferred habitat appears to be emergent

vegetation near the margins of slowly flowing water. Haliplus allisonae Brigham is a

junior synonym of H. dorsomaculatus.

Key Words: Haliplus allisonae, Nearctic, wing venation, mandible, female genital

sclerites

INTRODUCTION

Haliplidae is a small family of aquatic

adephagan beetles most obviously charac-

terized by the greatly expanded metacoxal

plates. There are about 65 species in North

America; these are placed in four genera,

the largest of which is Haliplus Latreille.

Wallis (1933) revised the Nearctic Haliplus

species; six species have subsequently been

described (Mank 1940; Leech 1948; Brig-

ham & Sanderson 1972, 1973; Brigham

1977; Wells 1989). The Nearctic members

of Haliplus have been assigned to three

subgenera, Haliplus sensu stricto, Liaphlus

Guignot and Paraliaphlus Wallis.

Haliplus dorsomaculatus Zimmermann

(1924) was described from a single male

specimen from “Boreal America”. It be-

longs to the nominotypical subgenus

(Guignot 1928, see discussion in Holmen

1987, p. 90), characterized by the presence

of pronotal plicae and penultimate segments

of the labial palps produced on the me-

dioapical angle. Wallis (1933) did not ex-

amine the type specimen for this species

and uncertainly referred to a male from CA

and a female from CO as “? dorsomacula-

tus’. Based on these specimens and his in-

terpretation of Zimmermann’s description,

Wallis provided a new description and in-

cluded H. dorsomaculatus in his key to

species. Certain characters in his key do not

apply to the H. dorsomaculatus holotype

and his illustration of the male genitalia is

incorrect for that species. These errors have

long been suspected; a specimen correctly

identified by H.B. Leech in 1946 has a note

attached: “Can this be the true H. dor-

somaculatus Zimm?”

More recently, Brigham (1977) de-

scribed a new species, Haliplus allisonae,

based on a series of specimens from BC

similar to those he previously correctly

identified as H. dorsomaculatus. Brigham

discussed separating his new species from

H. distinctus Wallis but mentioned H. dor-

somaculatus only in his amended version of

Wallis’s key. Here it is shown that H. alli-

sonae iS a junior synonym of H. dor-

somaculatus.

Apart from Zimmermann’s brief de-

scription, little has been published about H.

dorsomaculatus. No habitat information is

available and, due to the high level of mis-

identifications, all published distribution

information should be considered suspect.

Here, the habitat preference for H. dor-

Spencer Entomological Museum, Department of Zoology, University of British Columbia, Vancouver

BC V6T 124

44 J. ENTOMOL. SOC. BRIT. COLUMBIA 102, DECEMBER 2005

somaculatus 1s described and its currently

known distribution, based on examined

specimens, is mapped.

In water beetles, as with most organisms

with male intromittent organs (Eberhard

1985), male genitalia frequently provide

unequivocal characters for the identification

of species. Female characters are often

overlooked and in many cases only males

can be identified with certainty. Galewski

(1972a, 1972b) showed that European

haliplid females can be identified by man-

dibular and external genital characters and

more recent guides to these species have

included illustrations of the _ latter

(Franciscolo 1979, Holmen 1987). Here,

these characters and the metathoracic wing

are illustrated for H. dorsomaculatus and a

brief description of the internal structures of

the female reproductive system is given.

MATERIALS AND METHODS

In addition to the type specimens of H.

dorsomaculatus and H. allisonae, 718 H.

dorsomaculatus from the museums and

private collections listed in Table 1 were

examined. Collection information for all H.

dorsomaculatus specimens examined is

listed in the text (type specimens) or Ap-

pendix 1. Standard postal abbreviations for

states and provinces are used throughout.

Using a calibrated ocular micrometer on

a Wild M5 stereomicroscope with 10x eye-

pieces, the following measurements were

taken: L, distance from the front of the head

to the apices of the elytra in lateral view;

W, maximum width across the elytra in

dorsal view; IO, minimum distance be-

tween the eyes in dorsal view; and H, maxi-

mum width of the head measured across the

eyes in dorsal view. Five to eight specimens

of each sex were measured for a given lo-

cality; means and standard errors are re-

ported. Where insufficient numbers of

specimens from a single locality were avail-

able, those from localities in the same

county were combined. The normalized

interocular distance, Rel. IO, was calculated

using Rel. IO=IO/H.

Many male specimens were dissected in

order to examine the genitalia. For freshly

collected specimens, the genitalia were sim-

ply extended, spread and allowed to dry in

place. Older specimens were relaxed in hot

water and the genital capsule removed us-

ing fine forceps. The capsule was cleared of

non-sclerotized tissue in hot 10% aqueous

KOH and the genitalia were teased out of

the capsule. After examination, the genitalia

and capsule were stored in glycerin in geni-

talia vials on the same pin as the specimen.

Some females were dissected in a similar

manner to characterize the external genita-

lia. Other females were dissected following

a procedure similar to Mazzoldi (1996) and

Miller (2001a), using 0.2% aqueous Tolu-

idine Blue to stain the preparation before

microscopic examination. Mandibles_ or

metathoracic wings were removed from

several specimens and examined in tempo-

rary mounts on microscope slides. Nomen-

clature for wing venation follows Ward

(1979). Drawings were made using a draw-

ing tube on a Wild M5 stereomicroscope.

RESULTS

Haliplus dorsomaculatus Zimmermann

1924.

Holotype: 4, “Amer. bor.”, no date, no

collector, ZSMC.

Haliplus allisonae Brigham 1977. Holo-

type: 5, CANADA, BC, Creston, King Cr.,

22 Sept 1955, G. Stace-Smith, INHS; Allo-

type: 9, same data, INHS; Paratypes: 4 4,

4 2, same data; INHS; 1 3, 1 9, same data,

CAS. NEW SYNONYM Y.

For habitus drawings, see Brigham

(1977, Fig. 1) and Hatch (1953, Plate

XXXIII Fig. 3 - note the figure is incor-

rectly labeled as H. Jongulus LeConte and

is broader (L/W = 1.75), has a larger Rel.

IO (=0.63) and shorter pronotal plicae than

J. ENTOMOL. Soc. BRIT. COLUMBIA 102, DECEMBER 2005

Table 1.

Sources of Specimens examined in this study with abbreviations and contact person.

Abbr.

AMNH

BCPM

BYU

CAS

CBBC

CNC

EMEC

INHS

JBWM

MTEC

NSNH

OSAC

RDKC

ROME

UASM

UBCZ

WSU

ZSMC

Source

American Museum of Natural History, New York, New York

Royal British Columbia Museum, Victoria, BC

Monte L. Bean Life Science Museum, Brigham Young Univer-

sity, Provo, UT

California Academy of Sciences, San Francisco, CA

Chery! Barr Collection, c/o Essig Museum,

University of California, Berkeley, CA

Canadian National Collection of Insects,

Agriculture and Agri-Food Canada, Ottawa, Ontario

Essig Museum of Entomology, University of California,

Berkeley, CA

Illinois Natural History Survey, Champaign, Illinois

J.B. Wallis Museum, University of Manitoba, Winnipeg, Mani-

toba

Montana Entomological Collection, Montana State University,

Bozeman, MT

Nova Scotia Museum of Natural History, Halifax, Nova Scotia

Oregon State Arthropod Collection, Oregon State University,

Corvallis, OR

Rex D. Kenner Collection, Vancouver, BC

Royal Ontario Museum, Toronto, Ontario

Strickland Museum, University of Alberta, Edmonton, AB

Spencer Entomological Museum, University of British Colum-

bia, Vancouver, BC

Maurice T. James Entomological Collection,

Washington State University, Pullman, WA

Zoologische Staatssammlung Miinchen, Miinchen, Germany

Contact

L. H. Herman

R. A. Cannings

R. W. Baumann

D.H. Kavanaugh

C. B. Barr

Y. Bousquet

C. B. Barr

C. Favret

R. E. Roughley

M. A. Ivie

C. Majka

D. Judd

R.D. Kenner

D. Currie

D. Shpeley

K. M. Needham

R. S. Zack

M. Baehr

normal for H. dorsomaculatus).

Holotype. Male on point with labels:

white rectangular “Amer. bor.”; circular

white “Typop” handwritten; red rectangular

“Typus” printed; white rectangular

“Samml. A. Zimmermann”; red rectangular

“Holotype < Haliplus dorsomaculatus

Zimmermann 1924”, species handwritten;

blue rectangular “Zool. Staatsslg.

Miinchen”. Genitalia on point on separate

pin with labels: white rectangular “penis H.

dorsomaculatus” handwritten; white rectan-

gular “Samml. A. Zimmermann’; red rec-

tangular “Holotype Haliplus dorsomacula-

tus Zimmermann 1924”, species handwrit-

ten; blue rectangular “Zool. Staatsslg.

Miinchen”.

L = 3.32 mm, W = 1.62 mm, L/W =

2.05, Rel. IO = 0.51. Elongate oval, fairly

pointed posteriorly; not very convex, maxi-

mum near midpoint. Head amber with

darker post-ocular band extending to mid-

dle of compound eyes; labrum medially

emarginate with dense fringe of setae, me-

dial dark mark extending to clypeus, micro-

punctation sparser medially and anteriorly;

clypeus coarsely punctured, narrow im-

punctate area extending onto frons; intero-

cular area coarsely punctured with inverted

U-shaped impunctate area; dense punctures

along ocular margin; palpi same colour as

head, not infuscate apically, labial palpi

with penultimate segment produced medi-

ally; vertical carina behind eye on side of

46 J. ENTOMOL. SOC. BRIT. COLUMBIA 102, DECEMBER 2005

head. Pronotum yellow, paler than head;

narrow medial impunctate area extending

approximately 2/3 along midline from pos-

terior margin, ovate impunctate area medi-

ally each side of the midline, puncturing

less dense anterior to these impunctate ar-

eas; plicae deeply impressed with brown

lateral edges, impression steep on lateral

side, sloping more gradually medially; pli-

cae approximately 1/3 length of pronotum

measured along plical line; pronotal lateral

bead ends just anterior to posterolateral

comer; pronotum wider than elytral base by

width of lateral bead; hind margin sinuate

with point of inflection slightly medial to

midpoint of each side; anterior part of hy-

pomeron visible in lateral view; lateral mar-

gin of pronotum evenly curved ventrally

from posterior to anterior; anterolateral

corner about midpoint of eye. Elytra same

colour as pronotum with brown maculation

as follows: sutural blotch extending to stria

3 anteriorly, stria 4 posteriorly; medial dis-

cal blotch between striae 3 and 4 with pos-

teromedial corner connected to sutural

blotch; postmedial discal blotch between

striae 5 and 7; weak indications of preme-

dial discal blotch between striae 5 and 6,

most obvious along stria 6 where 4 or 5

punctures have merging “brown halos”.

Punctures of striae 1-4 large and blackened,

basal punctures of striae 2 and/or 3

enlarged; punctures of striae 5-10 decreas-

ing in size laterally, those of stria 9 similar

in size to interstrial punctures, stria 10 with

even smaller, barely blackened punctures.

Sutural interstrial row with punctures nearly

linear and single with occasional doubled or

misplaced punctures; subsequent interstrial

rows with punctures more widely spread,

especially apically where some rows obso-

lete; no micropunctures visible at 50x be-

tween larger punctures; elytral apical mar-

gin very weakly sinuate. Prosternal process

with sides converging to minimum of con-

striction, which occurs at about anterior

margin of procoxae, subsequently widening

posteriorly to just short of apex, then nearly

parallel to apex; not channeled anteriorly,

very shallowly channeled from anterior end

of constriction to apex, flat between mar-

gins; coarsely punctured with dense micro-

puncturing from near bottom of declivity to

posterior margin;. Metasternum mostly

impunctate, a few larger punctures laterally;

in ventral view, weakly depressed behind

mesocoxae. Ventral surface similar in col-

our to dorsal surface, legs somewhat redder;

micropuncturing as described in Brigham

(1977). Genitalia as in Brigham (1977,

Figs. 2-4) except aedeagus with distal end

of “dorsal hump” somewhat farther from

apex (fragments of tissue on dorsal edge

suggest “hump” originally in exact agree-

ment). Protarsi only slightly produced, with

specialized setae; protarsal claws about

equal in length but anterior claw more

sharply bent near base and somewhat

broader and thicker than posterior claw.

Mesotarsi slightly produced, with special-

ized setae; mesotarsal claws equal, longer

and more gently curved than _protarsal

claws.

Males. L = 3.16 + 0.02 mm, W = 1.65 +

0.01 mm, L/W = 1.92 + 0.01, Rel. IO =

0.53 + 0.002 (n=50; no significant geo-

graphical variation observed in these char-

acters, see Table 2); as in holotype except

as follows: medial discal blotch usually

merged with sutural blotch to give triangu-

lar sutural blotch with apex pointed posteri-

orly; one to three lateral blotches on each

elytron (no geographic pattern to variation

in maculation); in a minority of specimens,

apex of aedeagus slightly more tapered to

form a narrower tip (no geographic pattern

except tapered condition more prevalent in

CA specimens). Mandibles and metatho-

racic wing as in females (see below).

Females. L = 3.13 + 0.02 mm, W = 1.65

+ 0.01 mm, L/W = 1.89 + 0.01, Rel. IO =

0.54 + 0.003 (n=49; no significant geo-

graphical variation observed in these char-

acters, see Table 2); similar to males except

as follows: protarsal claws slender, evenly

curved, equal; pro- and mesotarsi not pro-

duced and without specialized setae. Exter-

nal genital sclerites, mandibles and

metathoracic wing as shown in Figs. 1-3.

Internal structures of female reproductive

system weakly sclerotized, spermathecal

duct short, such that spermatheca ventral to

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the prosternal process is essentially flat in

apex of vagina; bursa copulatrix connected

cross-section between the margins with

ventrally to vagina between spermatheca

and apex of vagina.

cross-section 1s convex with sparse coarse

dant larger punctures. In H. /ongulus, the

punctures and no micropunctures.

dense micropuncturing between the abun-

Diagnosis. Among the Nearctic mem-

bers of Haliplus s. str., the adults of H. dor-

Other external characters that help to

separate the adults of these species are the

somaculatus are most similar to those of H.

longulus; these two species share a similar

elongate shape and size. They may most

easily be separated by the characters of the

pronotal plicae and the elytral maculation.

In H. dorsomaculatus, each plica is asym-

prosternal process. In H. dorsomaculatus,

48 J. ENTOMOL. SOc. BRIT. COLUMBIA 102, DECEMBER 2005

Figure 1. Female genital sclerites of Haliplus dorsomaculatus, posterior up; (A) gonocoxae,

(B) tergal halves [X, (C) gonocoxosternites. USA, ID, Benewah Co., E Fork of Charles Creek,

27 July 1987, R.S. Zack, WSU.

Figure 2. Mandibles of Haliplus dorsomaculatus, female, anterior up; (A) dorsal view, (B)

ventral view. Canada, BC, Wynndel, Head of Lizard Creek, 07 April 1946, G. Stace-Smith,

SEM #3142.

J. ENTOMOL. SOc. BRIT. COLUMBIA 102, DECEMBER 2005

Figure 3. Right metathoracic wing of Haliplus dorsomaculatus, ventral view, female, O: ob-

longum cell; SAC: anterior sector cell, M: medial veins, A: anal veins, nomenclature according

to Ward (1979). Canada, BC, Surrey, 110 Ave. right-of-way N of 168 St., 03 September 2004,

R.D. Kenner, RDKC.

metrically impressed, usually with an infus-

cate lateral margin. In H. /ongulus, each

pronotal plica is symmetrically impressed

and is not infuscate. In H. dorsomaculatus,

the sutural blotch is roughly triangular with

the apex pointed posteriorly and there are

one to three lateral blotches on each

elytron. In H. /ongulus, the sutural blotch is

not roughly triangular and may be almost

obsolete; there are zero to one lateral

blotches on each elytron.

Males of H. dorsomaculatus and H.

longulus can also be separated by differ-

ences in their aedeagi. In H. dorsomacula-

tus, the aedeagus is shaped like “an inverted

boot” (Zimmermann 1924), see Brigham

(1977, Fig. 3). In H. Jongulus, the main axis

of the aedeagus is gently curved and the

apical quarter is narrowed, see Wallis

(1933, Fig. 9d), Hilsenhoff & Brigham

(1978, Fig. 5T), Gundersen & Otremba

(1988, Fig. 66), Durfee, Jasper &

Kondratieff (2005, Fig. 3).

Distribution. Haliplus dorsomaculatus

ranges along the coastal mountains from

northern CA to southern BC, east to west-

ern Montana and south along the Rocky

Mountains to northeastern UT (Fig. 4). No

records are known from south or east of the

Great Divide Basin in WY. This may be an

artifact as relatively little material from

either WY or CO was available for this

study. Durfee, Jasper & Kondratieff (2005)

list H. dorsomaculatus as “unconfirmed” in

CO. All examined specimens from CO and

CA previously determined as H. dor-

somaculatus were misidentified. A long

series of H. dorsomaculatus from Lassen

Co., CA was found in the unidentified ma-

terial from the CAS. This series is the only

valid record for CA known to the author.

Habitat. Many of the collection loca-

tions for H. dorsomaculatus imply lotic

habitats although some specimens appear to

have been collected in lentic habitats. De-

scriptions of habitats at some collection

sites have been provided by R. S. Zack

(pers. comm.), and R. W. Baumann (pers.

comm.). Zack reported taking haliplids by

digging or kicking into emergent vegetation

along the margins of generally slowly flow-

ing water. Baumann reported that all of his

sites were associated with spring-fed

creeks.

In 2004, H. dorsomaculatus was col-

lected from three locations in the Lower

Fraser Valley, BC: an apparently perma-

nently flooded drainage ditch beside a high-

way, a very shallow, apparently spring-fed

pool beside an industrial parking lot and a

small creek draining an apparently spring-

fed swamp at the base of bluffs on the edge

of the Fraser River flood plain. Only two

and three specimens, respectively, were

50 J. ENTOMOL. SOC. BRIT. COLUMBIA 102, DECEMBER 2005

Figure 4. Distribution of Haliplus dorsomaculatus based on specimens examined by the au-

thor. Collection data given in Appendix 1.

collected from the first two sites. The creek

site yielded a good series, including prob-

able teneral specimens, all from dense

emergent vegetation near the creek margin;

no H. dorsomaculatus were found in other

microhabitats in the creek.

DISCUSSION

The confusion surrounding the identifi-

cation of H. dorsomaculatus stems chiefly

from errors in Wallis (1933). It is likely that

Wallis had no H. dorsomaculatus speci-

mens in the material he examined. The ap-

parent CA specimen is possibly H. robertsi

Zimmermann although the species limits

and status of that taxon are not clear. The

CO specimen has not been found.

Two couplets in Wallis’s key can give

problems in determining H. dorsomacula-

tus: those concerning: (i) channeling of the

prosternal process and (11) the characters of

the mid-metasternum and male protarsal

claws. The first couplet requires a subjec-

tive choice between “evidently” and “very

feebly or not” channeled. The prosternal

process of H. dorsomaculatus can easily be

described as “very feebly channeled”. That

choice leads to H. Jongulus and many previ-

ous determinations have reached that con-

clusion. The second couplet requires a mid-

metasternum with lateral longitudinal im-

pressions and male protarsal claws equal.

J. ENTOMOL. Soc. BRIT. COLUMBIA 102, DECEMBER 2005

The mid-metasternum of H. dorsomacula-

tus has coarse punctures laterally and, in

some specimens, these overlap to produce

what could be described as a longitudinal

impression. However, the male protarsal

claws are never equal. The other half of this

couplet leads to H. distinctus Wallis. See

Brigham (1977) for a discussion of the dif-

ferences between H. dorsomaculatus (as H.

allisonae) and H. distinctus.

Comparison of the holotypes of H. dor-

somaculatus and H. allisonae makes it clear

that these are the same species. The only

significant difference between the two

specimens 1s in the base colour: yellow for

H. dorsomaculatus and reddish brown for

H. allisonae. The colour of the H. allisonae

type series 1s likely an artifact of specimen

preparation or storage because other speci-

mens collected by Stace-Smith at the H.

allisonae type locality over a two-week

period (which includes the collection of the

type series) are the “normal” yellow colour

and similar darker specimens occasionally

occur in preserved specimens of other spe-

cies.

Female reproductive characters.

Galewski (1972a) showed that, although

interspecific differences among the charac-

ters of the external female genitalia can be

subtle, there are larger differences at higher

taxonomic levels. Drawings of these

sclerites showing greater detail have subse-

quently been published for the European

species (Franciscolo 1979; Holmen 1987).

The external genitalia of H. dorsomaculatus

(Fig. 1) are, most similar to those of H.

wehnckei Gerhardt (=H. © sibiricus

Motschulsky, see Lundmark, Drotz & Nils-

son (2001)) illustrated in Holmen (1987,

Fig. 251). Characters of the gonocoxae

should be used with caution, since these

structures tend to collapse or distort upon

drying and are difficult to characterize in

drawings. Further study of the external fe-

male genital sclerites will need to be done

before their diagnostic utility for the North

American species can be determined.

Characters of the internal female repro-

ductive system, especially the spermatheca,

have proven useful both in the determina-

tion of phylogenetic relationships and in

species determinations (Burmeister 1976,

Ordish 1985, Mazzoldi 1996, Miller 2001a,

2001b). It is anticipated that these charac-

ters may be similarly useful in haliplids

(Holmen 1987). Burmeister (1976) investi-

gated the internal parts of the female repro-

ductive system in Haliplus (Neohaliplus)

lineatocollis (Marsham) although he did not

investigate the spermatheca in detail. His

findings differ from what was observed

here for H. dorsomaculatus; they are, how-

ever quite similar to what was found for

Haliplus (Liaphlus) gracilis Roberts (RDK,

unpublished data). In H. lineatocollis, the

spermathecal duct 1s relatively long and the

bursa copulatrix is connected to the anterior

end of the vagina (Burmeister 1976, Fig.

44b) whereas in H. dorsomaculatus, the

spermathecal duct is short and the bursa

copulatrix is connected ventrally to the va-

gina. With no other species for comparison,

it is hard to draw conclusions at this point

but these results do suggest that these char-

acters deserve further investigation.

Mandibular and metathoracic wing

characters. Galewski (1972b) suggested

the use of mandibular characters for the

identification of haliplid females. However,

in this work no difference in mandibular

characters was found between conspecific

males and females suggesting that these

characters may also be useful for the identi-

fication of males. Although the differences

in these characters between species can be

subtle, they are potentially useful at the

species and higher taxonomic levels

(Galewski 1972b, RDK, unpublished data).

Within the Haliplus s. str. species shown by

Galewski, the relative size of the apical

tooth of the right mandible appears to be a

useful character. Comparing the mandibles

of H. dorsomaculatus (Fig. 2) with those

shown in Galewski (1972b, Figs. 1-5, 7—

10), H. dorsomaculatus is, as above, most

similar to H. wehnckei. Until information is

available for more Nearctic species, it 1s

hard to determine how useful mandibular

characters will be for species identification

in North America.

Characters of the metathoracic wings

52 J. ENTOMOL. SOc. BRIT. COLUMBIA 102, DECEMBER 2005

are little used in the identification of bee-

tles, in part, because venation shows little

variation between species in many groups

of beetles. Ward (1979) discussed three

wing venation characters that appeared to

be useful in adephagan beetles: (i) the

shape of the oblongum cell, (11) the posi-

tion of the distal segment of M, relative to

M; and Cu, and (111) the position of the SA

vein which separates the SA cell from the

3R cell.

There appear to be few published ex-

amples of haliplid metathoracic wing vena-

tion comparable to that shown in Fig. 3.

Balfour-Browne (1943) examined 13 spe-

cies of Haliplidae but illustrated only the

area around the oblongum cell for Pe/to-

dytes caesus (Duftschmidt) (Balfour-

Browne 1943, Fig. 18). He discussed gen-

eral trends in the venational characters of

adephagan beetles, including haliplids, but

it is difficult to make detailed comparisons

in the absence of drawings. Illustrations of

the wings of three haliplid species have

been published: Haliplus (Haliplus) rufi-

collis De Geer (Franciscolo 1979), Pelto-

dytes muticus (LeConte) (Wallace & Fox

1980) and AHaliplus (Liaphlus) fulvus

(Fabricius) (Holmen 1987). In addition, a

Master's thesis (Mousseau 2004) includes

drawings for the three Nearctic Brvchius

species.

Even within a single genus, the overall

shape of the metathoracic wing can vary

(Mousseau 2004). In the Haliplus figures

referenced above, the posterior margin of

the wing is smooth and continuous, or

nearly so. In H. dorsomaculatus the poste-

rior margin is relatively deeply emarginate

at the position of the anal _ fold

(approximately aligned with the distal end

of the 1A, vein) and, to a lesser extent, at

the end of the 1A, vein (Fig. 3).

The wing venation is identifiably dif-

ferent in the figures for each of the seven

species listed above. The three Haliplus

species differ in the shape of the oblongum

cell and the relative position of the distal

portions of M; and M4. The position of the

SA vein appears to be similar in all three

species. These results suggest that in

Haliplidae, wing venation may provide

diagnostic characters at the species level,

although more work is needed to determine

the limits of intraspecifi: variation.

Distribution and iiabitat. Haliplus

dorsomaculatus appears to be a strictly

western Nearctic species (Fig. 4). A record

from NF (Larson 1987, Roughley 1991) is

almost certainly an error. No voucher

specimen has been found to support that

record and the original source is unknown.

Haliplus dorsomaculatus should be re-

moved from the NF list until a verifiable

voucher specimen has been found.

In western North America, the distribu-

tion for H. dorsomaculatus corresponds to

the mountainous areas. This correlation is

particularly apparent in WA where, in

eight decades of collecting, there are there

are records from the Olympic Mountains

and almost every county overlapping the

Cascade Mountains but none from eastern

WA . Haliplus dorsomaculatus may even-

tually be found in extreme southeastern

WA as there are records from the

neighbouring Blue Mountains in northeast-

ern OR. The correlation with mountainous

terrain fits well with a preferred habitat

associated with relatively permanent flow-

ing water, as springs or creeks need

sources of water at higher elevations.

Further collecting is needed to clarify

the distribution in several areas. Given the

known distribution in southern BC and

western MT, it seems likely that H. dor-

somaculatus will be collected in south-

western AB. More collecting is needed in

OR to determine if the apparent gap in the

distribution between northern CA and

northern OR is real. The southern limits,

both in CA and the Rocky Mountains, need

further investigation. In UT, H. dor-

somaculatus appears limited to the north-

eastern corner; its status in the Uinta

Mountains is uncertain. With the habitat

information given here, it should be possi-

ble to conduct directed searches to clarify

the status of this species in all of these ar-

eas.

J. ENTOMOL. SOC. BRIT. COLUMBIA 102, DECEMBER 2005

ACKNOWLEDGEMENTS

I thank Martin Baehr of the Zoolo-

gische Staatssammlung Miinchen for al-

lowing me to borrow the holotype of H.

dorsomaculatus and Colin Favret of the

Illinois Natural History Survey for loan of

the type series of H. allisonae and other

specimens determined by Brigham. I also

thank all of the listed curators and institu-

tions for loan of material, Richard Zack

(Washington State University, Pullman,

WA) and Richard Baumann (Brigham

Young University, Provo, UT) for permit-

ting the use of their information about

habitats, Mogens Holmen for help in locat-

ing the type specimen for H. dorsomacula-

tus and helpful comments, and Launi Lu-

cas for help in producing the distribution

map and inking the final copies of the fig-

ures. Finally, I thank Rob Roughley for

useful discussions, copies of references

and two wonderful field trips to try to add

records in CA, OR and AB.

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Wallace, F.L. and R.C. Fox. 1980. A comparative morphological study of the hindwing venation of the

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Kafer 20: 65-80.

APPENDIX 1

Material examined. The number preceding

the repository is the number of specimens; the

following abbreviations are used for names of

some of the collectors: DLG (D. L. Gustaf-

son), GSS (G. Stace-Smith), HBL (Hugh B.

Leech), MHH (Melvin H. Hatch), nc (no col-

lector), RDK (Rex D. Kenner), and RSZ

(Richard S. Zack).

CANADA.

BC: Abbotsford, roadside ditch, 14-1x-45,

HBL, 3 CAS, 1 CNC; Chilliwack, 23, 27, 28-

v-63, D.J. Farish, 5 UBCZ; Coquitlam, ditch

behind warehouses at W end of Rocket Way,

27-1x-2004, RDK, 2 RDKC; Creston, Goat

Mt. Lake, 5000 ft, 2-vii-33, GSS, 1 UBCZ;

Creston, Goat River, 25-vii-46, GSS, 1

UBCZ; Creston, King Creek, 27-vii-48, GSS,

1 UBCZ; same loc., 8-1x-48, GSS, 2 UBCZ;

same loc., 7, 15-vii-49, GSS, 3 UBCZ; same

loc., 18-1x-55, GSS, 2 UBCZ; same loc., 19-

ix-55, GSS, 1 INHS, 1 UBCZ; same loc., 21-

1x-55, GSS, 1 UBCZ; same loc., 2-x-55, GSS,

1 UBCZ; Creston, 24-ix-55, GSS, 1 CAS, 1

CNC; Fernie, 31-viii-35, HBL, 3 CAS; Kitch-

ener, roadside pond, 1-x-55, GSS, 1 UBCZ;

Mission City, 18, 20-vi-53, G.J. Spencer, 2

CNC; Salmon Arm, 2-1x-29, HBL, 1 CAS;

Surrey, shallow pond at 116 Avenue x 136

Street, 2-vi-2004, RDK, 3 RDKC; Surrey,

extension of 110 Avenue N of 168 Street, 2-

vi-2004, RDK, 2 RDKC; same loc., 3-1ix-

2004, RDK, 24 RDKC; Wynndel, head of

Lizard Creek, 7-x-45, GSS, 3 UBCZ; same

loc., 7-iv-46, GSS, 1 OSAC, 8 UBCZ; same

loc., 23-vi-46, GSS, 1 UBCZ; same loc., 4-v-

47, 1 UBCZ; same loc., 11-v-47, GSS, 14

CAS, 1 CNC, 8 UBCZ; same loc., 28-viu-47,

GSS, 1 CNC, 2 UBCZ.

USA.

CA: LASSEN Co.: Norval Flats, 5500 ft,

15-ix-20, J.O. Martin, 90 CAS.

ID: BEAR LAKE Co.: Little Spring, be-

low Davis Canyon campground, 18-viii-2004,

S.M. Clark & R.W. Baumann, 5 BYU.

BENEWAH Co.: Charlies Creek, 4-5 mi SE

of Emida, 2-ix-86, RSZ, 1 WSU; E Fork of

Charlies Creek, ca. 6 mi SE of Emida, 29-vii-

87, RSZ, 51 WSU. BONNER Co.: Pack

River, 8 mi N of Sandpoint, 22-ix-69, J.

Schuh, 2 AMNH; Sagle, 4-vii-49, N.M.

Downie, | OSAC. BUTTE Co.: Little Lost

River, 10.6 mi N of Howe, 25-ix-91, RSZ, 1

WSU. CLEARWATER Co.: Badger Mead-

ows ca. 7 mi E of Bovill, 28-vii-87, RSZ &

V.L. Zack, 45 WSU. LATAH Co.: Big

Meadow Creek Recreation Area, 5 mi. NW of

Troy, 9-iv-87, RSZ, 1 WSU; E Fork Emerald

Creek, ca 20 mi E of Harvard on Rt 447, 6-1x-

90, RSZ, 1 WSU; N Fork of Palause River ca

11 mi NE of Harvard, 19-v-87, 2 WSU; pond

by Rte 3, 7 m SSW of Clarkia, 28-ix-90.

LEMHI Co.: Canyon Creek, Railroad Can-

yon, 2.5 mi on Rte 29 below top of Bannock

Pass, 7075 ft, 24-viii-69, HBL, 5 CAS. VAL-

J. ENTOMOL. SOC. BRIT. COLUMBIA 102, DECEMBER 2005

LEY Co.: Trail Creek, FS Rte 22, 22.4 mi

NNE of Cascade, 22-ix-91, RSZ, 3 WSU.

MT: DEER LODGE Co.: Jct Hwy 10A &

Hwy 10, 7-vii-63, R.D. Anderson, 1 BYU.

FLATHEAD Co.: Thompson R., Rte 56 ca

6.6 mi S of Rte 2, 25-1x-90, 1 WSU.

GALLATIN Co.: no locality, 21-iv-24, nc, 3

MTEC; pond 2 mi from Bozeman, 17-v-55,

nc, 2 MTEC; side ponds of Bridger Creek,

4800 ft, 28-vil-87, DLG, 3 MTEC; pond by

Bridger Creek, 19-1x-87, DLG, 5 MTEC;

Bridger Creek, 4800 ft, 26-11-87, DLG, 3

MTEC; same loc., 18-v-87, DLG, 1 MTEC;

same loc., 20-vi-87, DLG, 2 MTEC; Bridger

Cr. 2 mi NE of Bozeman, 4800 ft, 18-v-86,

DLG, 1 MTEC; Bridger Warm Springs, 3 mi

NE of Bozeman on Hwy 86, 23-vii-89, C.B.

Barr, 1 CBBC; E Gallatin River, 4600 ft, 22-

vi-87, DLG, 1 MTEC; Gallatin River, 8 mi W

of Bozeman, 23-vi1i-86, DLG, 3 MTEC; same

loc., 1-ix-86, DLG, 3 MTEC; same loc., 13-x-

86, DLG, 1 MTEC; Gallatin River, Bozeman,

v-1x-88, DLG, pitfall trap, 2 MTEC; Gallatin

River, weedy side pond, 4700 ft, 6-1x-87,

DLG, 2 MTEC; same loc., 20-x-87, DLG, 1

MTEC; same loc. 10-xi-87, DLG, 4 MTEC;

Gallatin R., 4700 ft, 12-11-87, DLG, 7 MTEC;

same loc., 25-v-87, DLG, 2 MTEC; same loc.,

15-vu-87, DLG, 2 MTEC; same loc., viti-87,

DLG, pitfall trap, 1 MTEC; same loc., 10-xi-

87, DLG, 3 MTEC; same loc., 30-1x-88—12-

iv-1989, DLG, pitfall trap, 3 MTEC; same

loc., 12-iv—18-vii-89, DLG, pitfall trap, 2

MTEC. LAKE Co.: wet area just E of Swan

River, ca 3 mi S of Swan L. at N.F. Road 129,

4-vii-89, C.B. Barr, 2 CBBC. LEWIS &

CLARK Co.: Beaver Creek, 3-x-86, DLG, 1

MTEC; Beaver Creek near mouth, 3-x-86,

DLG, 1 MTEC. LINCOLN Co.: Libby Fish

Hatchery, 19-ix-86, DLG, 4 MTEC; same

loc., 20-11-87, DLG, 2 MTEC; same loc., 7-

iv-88, DLG, 4 MTEC. MEAGHER Co::

Sphagnum bog & beaver pond with ice, 26.8

mi N of White Sulphur, 27-x-73, R.E. Rough-

ley & M.L. Roughley, 1 JBWM. RAVALLI

Co.: Lee Metcalf N.W.R., Pond 2, 9-viii-94,

DLG, USFWS bottle trap, 1 MTEC; same

loc., Pond 3, 9-viui-94, DLG, USFWS bottle

trap, 3 MTEC; same loc., 8-ix-94, DLG,

USFWS bottle trap, 2 MTEC; same loc., 30-

ix-94, DLG, USFWS bottle trap, 2 MTEC.

OR: BAKER Co.: Beecher Creek, 7 mi N

of Halfway, 4250 ft, 14-x-69, D. Gray, K.

Gray, R. Rosenstiel, J. Schuh & D. Johnson, 1

AMNH:; Richland “Env.”, 2000 ft, 1-vi-45,

H.P. Chandler, 4 EMEC. GRANT Co.: pond 9

mi W of Seneca, 15-x-71, J. Schuh, 1 AMNH;

Rte 395 ca 9 mi N of Mt. Vernon, beaver

pond, 17-vii-90, RSZ, 4 WSU; spring at SE

Corner, 14-x-67, J. Schuh, 1 AMNH. MULT-

NOMAH Co.: marsh near Ainsworth State

Park, 10-x-88, RSZ & R.D. Akre, 13 WSU.

UNION Co.: Elgin, 29-vii-32, MHH, 1

OSAC.

UT: CACHE Co., A.J. Park, Blacksmith

Fork Canyon, 14-vi-2003, M.J. Peterson, 3

BYU; Logan, 27-iv-63, E. Drake, 1 BYU;

Spring Hollow, Logan Canyon, 16-iv-82, E.

Coombs, 3 BYU; Spring Hollow pond, Logan

Canyon, 5-v-89, nc, 6 BYU.

WA: CHELAN Co.: Rte 207 ca 3 mi N of

Coles Corner, 13-1x-90, RSZ, 1 WSU. CLAL-

LAM Co.: Matriotti Creek, Rte 101, 2.5 mi W

of Sequim, 24-vi-92, RSZ, 2 WSU. CLARK

Co.: E Fork of Lewis River ca 6 mi E of Heis-

son, 13-viii-89, J. Back, 1 WSU; Heisson ca 5

mi NE of Battle Ground, pools, 18-x1-88, J.

Back, 5 WSU. GRAYS HARBOR Co.: Co-

palis Lagoon, 20-v1i-33, T. Kincaid, 1 OSAC.

JEFFERSON Co.: Quilcene, 26-vii-36, nc, 6

OSAC. KING Co.: Canyon Park, Bothell, 17-

v-28, T. Kincaid, 2 OASC; pool ca 1.5 mi E

of Redmond, 24-vii-89, RSZ, 1 WSU; Cedar

Mt., 18-v-37, nc, 1 OSAC; same loc., 12-v-

39, MHH, 4 OSAC; same loc., 6-vii-39, I.M.,

2 OSAC; same loc., 9-v-40, MHH, 3 OSAC;

same loc. & date, R.H. Foster, 2 OSAC; same

loc., 10-vi-40, R.H. Foster, 2 OSAC; same

loc., 22-v-41, MHH, 2 OSAC; same loc. &

date, Thomas, 3 OSAC; same loc. & date,

D.R. Orcutt, 3 OSAC; same loc., 26-111-44, nc,

2 OSAC; same loc., 29-v-45, MHH, 2 OSAC;

same loc., 16-v-46, MHH, 3 OSAC; same

loc., 15-v-47, MHH, 3 OSAC, nc, 2 OSAC;

Evans Creek, 30-vi-29, MHH, 1 OSAC;

Malony’s Grove, 20-iv-32, nc, 2 OSAC;

North Bend, Malony’s Grove, 20-ix-29,

MHH, 14 OSAC; same loc., 10-v-30, MHH, 3

OSAC; same loc., 16-v-30, P. Ludy, 1 OSAC;

same loc., 16-v-31, MHH, 3 OSAC; Red-

mond, 4-vi-67, D. Frechin, 1 WSU; Renton,

22-v-41, Campell, 1 OSAC; Renton, Cedar

River, 22-v-41, Campell, 1 OSAC; same loc.,

29-x-45, H.J. Jensen, 5 EMEC; pond in Fors-

gren Park, Seattle, 23-iv-92, K.A. Rosema, 4

WSU; Seattle, viii-28, nc, 1 OSAC; Snoqual-

mie Falls, 10-v-30, nc, 1 OSAC; Snoqualmie

R., Malony’s Grove, 13-v-28, MHH, 48

56 J. ENTOMOL. Soc. BRIT. COLUMBIA 102, DECEMBER 2005

OSAC; Stillwater, 11-11-34, nc, 1 OSAC.

KITTITAS Co.: Rte US 10, 2.9 mi NW of

Ellensburg, 8-x-77, R. Thorne, 1 WSU;

Easton, 28-iv-39, MHH, 1 OSAC; Ellensburg,

19-vii-32, MHH, 2 OSAC; Jungle Cr., FS Rte

19 ca 11 mi N of Cliffdell, 31-viii-90, RSZ, 1

WSU; Manastash Canyon, ca 10 mi WSW of

Ellensburg, beaver pond, 3-x1-88, RSZ, 1

WSU; Roslyn Ponds, 7-v-2000, E. Sugden, 1

BYU. KLICKITAT Co.: pool by Klickitat

River ca 5.5 mi N of Lyle, 17-vii-89, RSZ, 1

WSU; Outlet Creek ca 1.5 mi S of Glenwood,

17-vii-89, RSZ, 2 WSU. MASON Co.: irriga-

tion ditch 3.5 mi SW of Kamilche on Rte 108,

19-1-94, RSZ, 1 WSU. OKANOGAN Co.:

Granite Creek ca 6 mi W of Republic on Rte

20, 11-ix-90, RSZ, 1 WSU; small pool 12 mi

N of Nespelem on RTE 155, 30-ix-87, R.D.

Akre, 23 WSU. PIERCE Co.: Mt. Rainier

National Park, Longmire, Fish Creek Valley,

spring-fed pond, 3000 ft, 15-vi-69, I. Smith,

36 ROME; Mt. Rainier National Park, Long-

mire, Fish Creek Valley, shallow pond in

flood plain, 3000 ft, 15-vi-69, I. Smith, 10

ROME; Mt. Rainier National Park, Tahoma

Creek, 2300 ft, 12-vii-73, A. Smetana, Z.

Smetana & D. Smetana, 2 CNC; WSU Res &

Ext. Center, Puyallup, pond, 24-1x-92, RSZ, 2

WSU. SKAMANIA Co.: Moffett Creek ca 2

mi W of North Bonneville, 12-x-88, RSZ &

R.D. Akre, 16 WSU. SNOHOMISH Co::

Canyon Park, 15-x-28, nc, 2 OSAC; same

loc., 6-v-30, nc, 2 OSAC; same loc., 8-vii-32,

nc, 1 OSAC; Martha Lake, 5-v-31, nc, 1

OSAC; Monte Cristo, in stomach of rainbow

trout, 987 m, 9-vu-38, Kiser, 1 OSAC; Sultan,

1-vi-53, B. Malkin, 1 OSAC; Thomas Lake,

5-vii-32, nc, 1 OSAC; same loc., 31-v-34, nc,

1 OSAC; same loc., 10-vii-34, nc, 3 OSAC.

WHATCOM Co.: Kendall Creek, 14-viii-32,

nc, | OSAC; Lynden, 6-v1i-64, nc, 3 OSAC;

same loc., 7-vil-64, nc, 3 OSAC; same loc.,

2-v1i-65, L. Russell, 4 OSAC; N Fork Nook-

sack River, 17-vii-32, nc, 1 OSAC; Silver

Lake, 14-viii-32, nc, 1 OSAC. YAKIMA Co.:

Milk Creek, FS Rte 12, 1-viii-90, RSZ, 2

WSU.

WY: TETON Co.: Snake River, 15.8 mi S

of Jackson on Rte 26/89, 24-ix-91, RSZ, 1

WSU.

J. ENTOMOL. Soc. BRIT. COLUMBIA 102, DECEMBER 2005

Lestes disjunctus and L. forcipatus (Odonata: Lestidae):

An evaluation of status and distribution in British Columbia

ROBERT A. CANNINGS' and JOHN P. SIMAIKA’

ABSTRACT

Of the five species of the damselfly genus Lesves that live in British Columbia, Lestes

forcipatus Rambur and L. disjunctus Selys are the most difficult to separate morphologi-

cally. Females can be readily distinguished by the size of the ovipositor, but males are

difficult to separate. In British Columbia, L. disjunctus is more common, widespread

and familiar. Before 1998, when it was first reported in BC, specimens of L. forcipatus

were misidentified as L. disjunctus because the former is known mainly from eastern

North America and most Lesfes species are usually most readily identified using male

characters. The identities of museum specimens of the two species were checked and

corrected by us as necessary. Ecological and behavioural observations and up-dated

distribution maps of the species are presented. Throughout its range in BC, L. forcipatus

is mostly sympatric with L. disjunctus but lives in a narrower range of habitats and lo-

calities — mostly cool sedge marshes and fens. The two species show some temporal and

behavioural separation.

Key Words: Odonata, Lestes disjunctus, Lestes forcipatus, British Columbia, distribu-

tion, habitat preference, plant associations, temporal separation, oviposition

INTRODUCTION

Five species of Lestes occur in British

Columbia (BC): JL. congener Hagen

(spotted spreadwing), L. disjunctus Selys

(northern spreadwing), L. dryas Kirby

(emerald spreadwing), L. forcipatus Ram-

bur (sweetflag spreadwing), and L. ungui-

culatus Hagen (lyre-tipped spreadwing).

Lestes disjunctus is the most common,

widespread and familiar of these in the

province, and one of the most abundant

odonates in Canada where it ranges as far

north as the Arctic treeline (Cannings

2002). It inhabits many types of standing

water habitats with abundant aquatic vege-

tation and, in southern BC, adults have been

recorded from mid-June to mid-October

(Cannings 2002).

Lestes forcipatus, although as abundant

as L. disjunctus in some cold fen habitats, is

generally much less common; both species

often occur at the same site. Lestes forci-

patus does not range as far north as L. dis-

Junctus but, although not known from much

of BC’s north, it has been collected in

south-eastern Yukon (S.G. Cannings, pers.

comm.). In the western Canadian Cordil-

lera, it 1s most common in sedge fens

(Cannings 2002). Walker (1953) described

L. forcipatus habitat in Ontario as "ponds,

both temporary and permanent, marshy

lakes, and slow, weedy streams". In BC,

adults of L. forcipatus have been reported

from mid-May to mid-September

(Cannings 2002).

Lestes forcipatus was first recorded in

BC in 1998, when it was collected in the

Rocky Mountain Trench north of Golden

and subsequently found in many other lo-

calities in the south-eastern part of the prov-

ince (Cannings eft al. 2005). However, un-

doubtedly it has long been a resident of the

province but was overlooked because of its

close resemblance to L. disjunctus. Before

1998, L. forcipatus was unknown west of

Saskatchewan in Canada (Walker 1953,

Westfall and May 1996), although in 1997

Royal British Columbia Museum, 675 Belleville St., Victoria, British Columbia, Canada V8W 9W2

* #323-3969 Shelbourne St., Victoria, British Columbia, Canada V8N 6J5

58 J. ENTOMOL. SOC. BRIT. COLUMBIA 102, DECEMBER 2005

it was found in Washington State, the first

record west of Montana in the United States

(Cannings ef al. 2005). The species is now

known from seven counties in Washington

and one in Idaho (Paulson 2005). Subse-

quently, collectors found L. forcipatus at

several other BC locations farther south and

west in 1999; by 2000 it had been collected

on Vancouver Island. Inventories in north-

erm BC (2000-2003) have extended its

range to about 55°N latitude (Cannings ef

al. 2005), although records in south-eastern

Yukon in 2004 and 2005 (S.G. Cannings,

pers. comm.) indicate it probably ranges

through much of north-eastern BC, at least.

In addition, Catling et al. (2004) reported

the species from southern Northwest Terri-

tories near Fort Smith. A map of the distri-

bution of L. forcipatus in North America

was published by Donnelly (2004).

Catling (2002), Donnelly (2003) and

Simaika and Cannings (2004) provided

practical information on characters for the

identification of the two species. Simaika

and Cannings (2004) particularly empha-

sized the usefulness of pruinosity patterns

in western North American populations. A

recent check of some specimens in the

Royal BC Museum (RBCM) (Victoria) and

the Spencer Entomological Museum, Uni-

versity of BC (SEM) (Vancouver) by one of

us (JPS) revealed that a signficant number

identified as L. disjunctus (even among

those collected after 1998) are misidentified

L. forcipatus (Cannings eft al. 2005). The

discovery of misidentified specimens indi-

cates that other museum collections across

western Canada probably contain such

specimens. Herein we report the results of

an identification check of all L. disjunctus

specimens in the RBCM and the SEM, the

two collections holding the majority of

Odonata specimens from BC.

This paper establishes accurate distribu-

tions for both species in BC by publishing

up-dated distribution maps. We also pro-

vide revised information on habitat prefer-

ences and life histories.

MATERIALS AND METHODS

Specimens. We examined 1853 speci-

mens previously identified as L. disjunctus

in the RBCM and SEM collections and

separated the two species using characters

documented in Simaika and Cannings

(2004). Females were identified by the rela-

tive lengths of the ovipositor. The most

useful character for separating males is the

amount of pruinescence on the thorax and,

in L. forcipatus, the presence of a bare, non-

pruinose patch on the posterior third of the

dorsum of the second abdominal segment.

Habitat and life-history data. Informa-

tion on habitat preferences, emergence

times, flight period and breeding behaviour

was extracted from the RBCM and SEM

databases. The wetland site association

classification used is that of MacKenzie and

Moran (2004); site associations noted are

listed and defined in Table 1.

Distribution maps. Maps (Figs. 1 and

2) were produced electronically from the

databases of the RBCM and SEM by Clo-

ver Point Cartographics Ltd. (Victoria, BC)

using Microsoft Visual Basic version 6 and

Environmental Systems Research Institute

(ESRI) Arc Info Workstation version 9.0

(ESRI 2005). The base line features are

from Terrain Resource Information Map-

ping (TRIM) 1: 2,000,000 and the surface

model is based on Clover Point and TRIM

Digital Elevation Model data. TRIM data

are used under license from the BC Minis-

try of Environment.

RESULTS AND DISCUSSION

Changes in specimen identification. In

the RBCM collection, 38 specimens (236,

152) from 3ritish Columbia collected be-

fore 2004 and previously identified as L.

disjunctus were re-identified as L. forci-

patus; five (33, 29) identifications were

changed in the SEM collection. These

changes represent 2.3% of the sample. This

material came from ten localities over the

southern two-thirds of the province in the

J. ENTOMOL. SOC. BRIT. COLUMBIA 102, DECEMBER 2005 59

Table 1.

Main wetland habitat types used by Lestes disjunctus (L. dis.) and L. forcipatus (L. for.) in Brit-

ish Columbia. Wetland site associations and codes are taken from MacKenzie and Moran

(2004). Site association names indicate dominant plant species used to define the habitat type;

codes are used in the discussion of Lestes habitat in the text. A, absent; C, common; R, rare; U,

uncommon.

a: ee Site Association Name ee eee

Saline Gs01 Distichlis spicata var. stricta (Alkali saltgrass) C A

associations Gs02 Puccinellia nuttalliana — Hordeum jubatum (Nuttall’s alkahi- = C A

at grassland grass - Foxtail barley)

ponds Gs03 Carex praegracilis (Field sedge) C A

aaa ieee ee nee Erne sar Saaaainein sae iil

Peat-moss)

Wb13 Carex limosa — Menyanthes trifoliata — Sphagnum spp. U A

(Shore sedge - Buckbean - Peat-moss)

Wb50 Ledum groenlandicum — Kalmia microphylla — Sphagnum Cc A

Bogs spp. (Labrador Tea — Bog-laurel - Peat-moss)

WbS1 Pinus contorta — Empetrum nigrum — Sphagnum austinii C A

(Shore pine—Black crowberry—Tough peat-moss)

Wb52 Juniperus communis — Trichophorum cespitosum — c A

Rhacomitrium lanuginosum (Common juniper —

Tufted clubrush — Hoary rock-moss)

~ Wf01 = Carex aquatilis — Carex utriculata (Water sedge-~Beaked CC U_

Sedge)

Wf02 Betula nana — Carex aquatilis (Scrub birch — Water sedge) C U

Wf03 Carex aquatilis — Sphagnum (Water Sedge — Peat-moss) R R

Wf04 Salix barclayi — Carex aquatilis — Aulacomnium palustre 10) R

(Barclay’s willow — Water sedge — Glow moss)

Wf05 Carex lasiocarpa — Drepanocladus aduncus (Slender sedge = -C U

— Common hook-moss)

Wf06 Carex lasiocarpa — Menyanthes trifoliata (Slender sedge — C C

ae Buckbean)

Wf07 Betula nana — Menyanthes trifoliata — Carex limosa fens C C

(Scrub birch — Buckbean — Shore sedge)

Wf08 Carex limosa — Menvyanthes trifoliata — Drepanocladus spp. = C C

(Shore sedge — Buckbean — Hook moss)

Wf09 Eleocharis quinqueflora — Drepanocladus (Few-flowered U A

spike-rush — Hook moss)

Wfl0 Trichophorum alpinum — Scorpidium revolvens (Hudson C C

Bay clubrush — Red hook-moss)

Wfl2 Eriophorum angustifolium — Caltha leptosepala (Narrow- C A

leaved cotton-grass — Marsh-marigold)

er eee at oe Cee ae oi. ain

sedge)

Wm02 Equisetum fluviatile - Carex utriculata (Swamp horsetail — C U

Beaked sedge)

Marshes Wm04 Eleocharis palustris (Common spike-rush)

Wm05 Typha latifolia (Cattail)

Wm06 Schoenoplectus acutus (Great bulrush)

Cy Cro)

> AAA

Wm07 Juncus balticus (Baltic rush)

60 J. ENTOMOL. Soc. BRIT. COLUMBIA 102, DECEMBER 2005

Figure 1. Distribution of records of Lestes disjunctus in British Columbia to 2004. Data repre-

sent specimen records only from the Royal BC Museum, Victoria, and the Spencer Entomo-

vicinities of Bowser, Cawston, Duncan,

Fort St. James, Germansen Landing, Horse-

fly, Mackenzie, Qualicum Beach, Rogers

Pass, and Wells Gray Provincial Park.

Distribution and status. The newly

plotted range maps for L. disjunctus (Fig. 1)

and L. forcipatus (Fig. 2) update our know!l-

edge of the BC distributions of the species

to the 2004 collecting season. The distribu-

tion of L. disjunctus in the province remains

unchanged; it 1s so common and wide-

spread that the subtraction of specimens

from ten localities made little difference to

the updated range map (Fig. 1). Indeed, in

all but a very few localities, ZL. disjunctus

has been collected wherever L. forcipatus 1s

found. Lestes disjunctus is second only to

Enallagma boreale Selys as the most fre-

quently collected odonate in BC; it occurs

over the entire province where dragonflies

are able to live; the only major gaps in its

known distribution are those areas difficult

to access by road.

Although our reidentifications show that

L. forcipatus has lived in the region since

long before 1998, all but five of the 43

known localities were recorded from 1998

to 2004. Thus, in terms of its known status

in BC, the species has gone from anonym-

ity to being a widespread and fairly com-

mon taxon in only seven years. Although it

was collected at two localities in the year of

its discovery and 12 the year after, it was

retained on the provincial Blue List of spe-

cies of concern until 2004 (Ramsay and

Cannings 2005). Despite this rapid change

in its status, L. forcipatus is clearly less

common and widespread than L. disjunctus

J. ENTOMOL. SOC. BRIT. COLUMBIA 102, DECEMBER 2005 61

in BC. In the Okanagan-Similkameen basin,

probably the most thoroughly collected area

of the province, it has been found at only

one locality (near Cawston). It has yet to be

collected in the Shuswap region or in the

Lower Mainland of south-eastern BC and

still must be considered rare on the coast in

general. It was not found in intensive sur-

veys in the Peace River and Fort Nelson

regions in 1997, nor along Highway 37 or

in the Atlin area in 2003, although in 2004

and 2005 it was collected in south-eastern

Yukon (S.G. Cannings, pers. comm.). Fur-

ther study will likely fill some of these

gaps; nevertheless, the narrower range of its

preferred habitats will continue to make it

harder to find than L. disjunctus in most

places in BC.

Habitat Requirements. See Table 1. A

ce 200 km

Figure 2. Distribution of records of Lestes forcipatus in British Columbia to 2004. Data repre-

sent specimen records only from the Royal BC Museum, Victoria, and the Spencer Entomo-

logical Museum, University of BC, Vancouver.

major basis for the greater abundance and

wider distribution of L. disjunctus com-

pared with L. forcipatus is the ability of the

former to use a wider range of habitats.

This is especially true of warmer habitats in

the southern parts of the province such as

eutrophic marshes (Wm04-07) and saline

ponds (Gs01-03), where L. disjunctus is

common and L. forcipatus is rare or absent.

Lestes forcipatus has yet to be found in the

widespread bogs of the outer coast, from

the Queen Charlotte Islands and Prince

Rupert regions to Vancouver Island and the

Fraser River delta (Wb13, Wb50-52). Both

species are found in Carex and Equisetum

marshes (e.g. Wm01-02), but L. disjunctus

is more common in these places. Lestes

forcipatus appears to be most frequent in

fens or bogs dominated by Carex,

62 J. ENTOMOL. SOc. BRIT. COLUMBIA 102, DECEMBER 2005

Trichophorum, Menyanthes, Comarum and

mosses such as Sphagnum, Drepanocladus

and Scorpidium (Wb12, Wf01-10, 12).

However, L. disjunctus is usually more

abundant in these habitats and, apparently,

L. forcipatus 1s absent from many localities

with such habitat types, especially in the

North, that superficially appear ideal for its

development. Walker (1953) notes that, in

eastern Canada, L. forcipatus is also more

locally distributed than L. disjunctus. In

summary, L. forcipatus 1s most common in

cold sedge and moss fens and uncommon,

rare or absent in warmer habitats such as

eutrophic marshes.

Life histories. Lestes disjunctus adult

records range from 15 May to 9 October.

The bulk of them fall between late June and

early August with a peak in the last half of

July. For example, on 22 July 1996 hun-

dreds of teneral adults were observed at

Burs Bog in the Fraser River delta. Re-

cords of mating pairs range from 12 July to

5 October and oviposition dates range from

12 July to 17 September. Adult records of

L. forcipatus range from 18 May to 4 Sep-

tember; about 85% of these are from late

June through late July, with the peak in the

last half of July. Mating has been observed

from 28 June to 14 August and oviposition

from 12 July to 14 August. Although there

is strong overlap of the flight periods of the

two species in BC, there is some evidence

that adult L. forcipatus emerge earlier than

L. disjunctus where they co-occur. At Nahl-

beelah wetlands near Kitimat on 10 July

2005, fully mature adult L. forcipatus were

common but the population of L. disjunctus

was just beginning to emerge. At Hamilton

Marsh near Qualicum Beach, adult L. forci-

patus were flying on 18 May 2004; adult L.

disjunctus appeared on 23 June and sexu-

ally mature specimens were not observed

until 3 July. This suggests a difference of

about two to four weeks in emergence times

of the two species and is similar to the

amount of time between the first emergence

of the two species in eastern Canada noted

by Walker (1953).

In addition to a possible temporal shift

in the flight period and, consequently, the

mating times of the two species, there may

be some interspecific differences in ovi-

position behaviour. Simaika (2005) and

Simaika and Cannings (2006) reported that

at Hamilton Marsh, near Qualicum Beach,

ovipositing females of L. disjunctus in-

serted eggs into only two species, Carex

lanuginosa Michaux and Juncus arcticus

Willdenow. Females on C. lanuginosa ovi-

posited into fresh stems, just above the wa-

ter surface; on J. arcticus they laid eggs in

dead stem tissue, about 10 cm from the tip

of the stem. Lestes forcipatus will also ovi-

posit on C. /anuginosa and J. articus but,

unlike L. disjunctus, it appears to prefer the

living stems of J. arcticus and will also

utilize Menyanthes trifoliata L.

These observations suggest that there

may be some niche separation of L. disjunc-

tus and L. forcipatus in BC. More research

is required to elucidate the ecological and

behavioural differences between these two

closely related, sympatric damselflies.

ACKNOWLEDGEMENTS

Karen Needham and Rex Kenner

(Spencer Entomological Museum, UBC,

Vancouver) loaned specimens. Mike

Shasko of Cloverpoint Cartographics

(Victoria) produced the maps as part of a

larger project mapping the results of drag-

onfly surveys in northern BC, funded in

part by the Habitat Conservation Trust

Fund. The personal communications with

Syd Cannings (NatureServe Yukon, Yukon

Territorial Government, Whitehorse, YT)

are used with permission.

REFERENCES

Cannings, R.A. 2002. Introducing the Dragonflies of British Columbia and the Yukon. Royal British Co-

lumbia Museum, Victoria, BC.

Cannings, R.A., S.G. Cannings, L-.R. Ramsay and G.E. Hutchings. 2005. Four species of Odonata new to

British Columbia, Canada. Notulae odonatologicae 6: 45-49.

J. ENTOMOL. SOC. BRIT. COLUMBIA 102, DECEMBER 2005 63

Catling, P.M. 2002. An evaluation of some characters separating male Lestes disjunctus and Lestes forci-

patus in Ontario. Ontario Odonata 3: 51-58.

Catling, P.M., S. Carriere, D. Johnson and M. Fournier. 2004. Dragonflies of the Northwest Territories,

Canada: New records, ecological observations and a checklist. Argia 16(1): 9-13.

Donnelly, T.W. 2003. Lestes disjunctus, forcipatus and australis: a confusing complex of North American

damselflies. Argia 15: 10-13.

Donnelly, T.W. 2004. Distribution of North American Odonata, Part II: Calopterygidae, Lestidae, Coe-

nagrionidae, Protoneuridae, Platystictidae. Bulletin of American Odonatology 8: 33-39.

ESRI (Environmental Systems Research Institute, Inc.). 2005. Arc/Info Workstation version 9.0. Redlands,

CA.

MacKenzie, W.H. and J.R. Moran. 2004. Wetlands of British Columbia: a guide to identification. Land

Management Handbook No. 52. Research Branch, BC Ministry of Forests, Victoria, BC.

Paulson, D.R. 2005. Western US Odonata range maps: Lestes forcipatus. http://www.ups.edu/biology/

museum/westernOD.html.

Ramsay, L.R. and R.A. Cannings. 2005. Determining the status of British Columbia's dragonflies, pp. 1-12.

In T.D. Hooper (Ed.), Proceedings, Species at Risk 2004 Pathways to Recovery Conference, March 2-6,

2004, Victoria, British Columbia. http://www.speciesatrisk2004.ca/pdf/ramsay_edited_final_feb_28.pdf

Simaika, J.P. 2005. Diversity and behaviour of dragonflies (Insecta: Odonata) at Hamilton Marsh, Vancou-

ver Island, British Columbia. BSc Honours thesis. University of Victoria, Victoria, BC.

Simaika, J.P. and R.A. Cannings. 2004. Lestes disjunctus Selys and L. forcipatus Rambur (Odonata: Lesti-

dae): Some solutions for identification. Journal of the Entomological Society of British Columbia 101:

131-140.

Simaika, J.P. and R.A. Cannings. 2006. The Odonata of Hamilton Marsh, Vancouver Island, British Colum-

bia, Canada. Notulae odonatologicae 6(7): in press.

Walker, E.M. 1953. The Odonata of Canada and Alaska. Vol. 1. University of Toronto Press, Toronto, ON.

Westfall, M.J. Jr., and M.L. May. 1996. Damselflies of North America. Scientific Publishers, Inc. Gaines-

ville, FL.

J. ENTOMOL. Soc. BRIT. COLUMBIA 102, DECEMBER 2005

J. ENTOMOL. SOc. BRIT. COLUMBIA 102, DECEMBER 2005 65

Mammal Fleas (Siphonaptera: Ceratophyllidae)

New for Alaska and the Southeastern Mainland Collected

During Seven Years of a Field Survey of Small Mammals

GLENN E. HAAS’, JAMES R. KUCERA’, AMY M. RUNCK’,

STEPHEN O. MACDONALD?‘ and JOSEPH A. COOK’

ABSTRACT

Ten taxa of mammal fleas were among 124 collection records from 12 host species (one

shrew, nine rodents and two carnivores), at 72 localities on the southeastern Alaska

mainland in 1989 and during an extensive survey of mammals in 1992-1995 and 1997-

1999. Megabothris asio megacolpus (Jordan) ex Microtus pennsylvanicus (Ord), Mala-

raeus telchinus (Rothschild) ex Peromyscus keeni (Rhoads) and Clethrionomys gapperi

(Vigors) are new fleas for Alaska. Orchopeas caedens (Jordan) ex Tamiasciurus hud-

sonicus (Erxleben) is a new flea for southeastern Alaska. Syvnaptomys borealis

(Richardson) is a new host record for Opisodasys k. keeni (Baker). The other six taxa of

fleas collected were Hystrichopsylla dippiei spinata Holland, H. o. occidentalis Holland,

Catallagia charlottensis (Baker), Ceratophyllus ciliatus protinus Jordan, Megabothris

abantis (Rothschild) and Opisodasys vesperalis (Jordan). Of these, H. 0. occidentalis, C.

charlottensis and M. abantis have seven new host records for the southeastern Alaska

mainland. Distribution patterns of the fleas and their host relationships in North America

are discussed.

Key Words: fleas, Siphonaptera, mammals, Alaska

INTRODUCTION

The advancement in knowledge of the

fleas of southeastern Alaskan mammals has

lagged behind that of Alaska west of the

Yukon Territory in part due to difficulties

of travel in the fragmented, rugged coastal

to montane topography. An extensive sur-

vey by the University of Alaska Museum,

Fairbanks, of shrews, mice, voles, lem-

mings and some larger mammals, such as

arboreal squirrels during 1992-1995

(MacDonald and Cook 1996) and 1997-

1999 included the collection of fleas. This

survey produced 124 collection records

(including three from an earlier study of

marten) at 72, mostly new, localities. Two

fleas of mice and voles new for Alaska, one

squirrel flea new for southeastern Alaska,

seven new host records for three other fleas

for the southeastern Alaska mainland, and

one new lemming host record for a mouse

flea were added (Table 1).

MATERIALS AND METHODS

The fieldwork for the mammal survey

was conducted as described by Murrell ef

al. (2003) on ticks collected from some of

the same mammal specimens that produced

some of the fleas reported on here. Full data

for the mammal specimens can be obtained

at http://arctos.database.museum by track-

ing the University of Alaska Museum of the

"PO Box 60985, Boulder City, NV, United States 89006

* Associated Regional and University Pathologists, Inc., Salt Lake City, UT, United States

; Biology Department, Idaho State University, Pocatello, ID, United States

* Museum of Southwestern Biology, University of New Mexico, Albuquerque, NM, United States

66 J. ENTOMOL. SOC. BRIT. COLUMBIA 102, DECEMBER 2005

Table

Mammalian hosts of the 10 taxa of fleas with present records for the southeastern Alaska

mainland.

Mammal

Sorex cinereus Kerr, masked shrew

Tamiasciurus hudsonicus (Erxleben), red squirrel

Glaucomys sabrinus (Shaw), n. flying squirrel

Peromyscus keeni (Rhoads), Keen’s mouse

Clethrionomys rutilis (Pallas), n. red-backed vole

C. gapperi (Vigors), s. red-backed vole

Microtus pennsylvanicus (Ord), meadow vole

M. longicaudus (Merriam), long-tailed vole

Synaptomys borealis (Richardson), n. bog

lemming

Zapus hudsonius (Zimmerman), meadow jumping

mouse

Martes americana (Turton), marten

Mustela vison Schreber, mink

' New host record for southeastern Alaska.

> New for southeastern Alaska.

> New for Alaska.

* New host record.

> Probably from marten.

North AF number listed under Material

Examined.

In the laboratory the fleas were prepared

for microscopic study by transferring them

from the labeled field vials of 70% ethanol

to a rinse in distilled water, then submerged

in 10% KOH until sufficiently bleached (1

hr to 3 days), rinsed two or three times in

distilled water, dehydrated in graduated

ethanols (to 90%), degreased in oil of win-

tergreen, rinsed in xylene and mounted in

Canada balsam on labeled microscope

slides. Voucher specimens were deposited

Fleas

Hystrichopsylla o. occidentalis Holland

Ceratophyllus ciliatus protinus Jordan

Orchopeas caedens (Jordan)

Opisodasys vesperalis (Jordan)

H. o. occidentalis

Catallagia charlottensis (Baker)

C. c. protinus

Megabothris abantis (Rothschild)

Opisodasys k. keeni (Baker)

Malaraeus telchinus (Rothschild)?

H. o. occidentalis’

C. c. protinus

M. abantis

H. o. occidentalis’

C. charlottensis

M. abantis'

M. telchinus°

C. charlottensis'

M. abanitis'

Megabothris asio megacolpus (J ordan)*

M. abantis

M. abantis'

O. k. keeni’

M. abantis

Hystrichopsvlla dippiei spinata Holland

H. d. spinata’

in the United States National Museum

(USNM) and the Canadian National Collec-

tion of Insects and Arachnids (CNC). Fol-

lowing is a list of collectors and their acro-

nyms used in this paper: C. J. Conroy

(CJC), J. A. Cook (JAC), J. Foreit (JF), R.

Heinen (RH), S. O. MacDonald (SOM), S.

R. Peterson (SRP), A. M. Runck (AMR), C.

T. Seaton (CTS), K. D. Stone (KDS), A. A.

Tsvetkova (AAT) and M. J. Wike (MJW).

Specimens without acronyms are in the

collections of the authors.

J. ENTOMOL. SOC. BRIT. COLUMBIA 102, DECEMBER 2005

SPECIES ACCOUNTS

HYSTRICHOPSYLLIDAE

Hystrichopsylla dippiei spinata Holland,

1949

Material examined: USA: AK: all from

Juneau area; 30.4 km NW, between Amalga

Harbor and Windfall Lake, 19 ex Martes

americana (Turton), 13-x11-89, SRP; ca. 34

km NW, Yankee Basin trail, 12 same host,

30-xii-89, SRP; Eagle River trail, 12 ex

Mustela vison Schreber or Martes ameri-

cana, 30-x11-89, SRP.

These three records continue a series

begun in 1987 with the first record reported

by Haas ef al. (1989). The range of this

large flea was only extended ca. 3.3 km

farther NW of Juneau. Although most hys-

trichopsyllids are associated with insecti-

vores or rodents, this flea infests mustelids.

The new records provide support for marten

as the true host. The Eagle River mink and

marten were packed together in the same

container. Only one valid record (19) for

mink (Haas et al. 1979) exists. Therefore,

the original host for the Eagle River record

may have been the marten. Thus, 24 speci-

mens (844, 169) and 18 collection re-

cords (Haas et al. 1978, 1979, 1980, 1982,

1989, present study) in southeastern Alaska

including islands are now known. Except

for one mink, two humans, and the uncer-

tain host record noted above, all host speci-

mens were marten. There are no Alaskan

records from shrews or rodents although

such collections have been made in British

Columbia and Oregon (Holland 1957; Hop-

kins and Rothschild 1962; Lewis ef al.

1988). Most specimens were from skunks

(Spilogale spp.). Holland (1949) also had

two females from marten and ermine

(Mustela erminea) from Vancouver Island

that he excluded from the series of H. spi-

nata new species.

Hystrichopsylla_ occidentalis occiden-

talis Holland, 1949

Material examined: USA: AK: Berg

Bay, 56°21°49”N, 132°00’29"W, 19 ex

Clethrionomys gapperi (Vigors) [AF

21819], 4-vi-97, CTS. Echo Cove, 59°

31°45”N, 134°21°58”W, 13 ex Clethriono-

mys rutilis (Pallas) [AF21587], 16-vu-97,

CTS. Frosty Bay, S side, 56°03’28”N, 131°

58’01”"W, 19 ex C. gapperi [AF22904], 8-

vil-97, CTS. Klukwan, 17 km W & 30 km

N, Kelsall River drainage, 14, 19 ex Sorex

cinereus Kerr [AF8096], 30-vui-94, JAC,

SOM. Taku River, Canyon Island, 58°

33’N, 133°41’W, 19 ex Peromyscus keeni

(Rhoads) [AF8272], 17-vii-94, JAC; same

data but C. gapperi [AF2856], 16-vii-94,

JAC.

This small relative of H. d. spinata is a

hygrophilous parasite of shrews, voles (type

host: Clethrionomys gapperi) and mice and

occurs in a long narrow range with a notice-

able concentration of collection localities

along the coast from northern California to

southwestern Alaska (Holland 1949: Map

6; Campos and Stark 1979: Fig. 60; Hol-

land 1985: Map 12; Lewis et al. 1988: p.

63). The six new mainland records for five

new localities as well as the four earlier

records indicate that this flea is uncommon

on trapped hosts, in accordance with its

behaviour like a nest flea. Consequently,

the majority of the fleas reside in nests of

the hosts. For example, from two nests of

Microtus oeconomus (Pallas) on the Chilkat

Peninsula, Haas (1982) collected a total of

534 and 49° H. o. occidentalis. This ex-

ceeds the total of 2¢¢ and 59 from the

six trapped host specimens listed above.

The predominance of nest populations was

confirmed for H. occidentalis linsdalei Hol-

land with large samples of nests (179) and

hosts (877) during a 2.5-y survey in north-

ern California by Stark (2002). He reported

a seasonal association with higher popula-

tions in nests of voles than on trapped voles

except during a short time in fall.

Sorex cinereus is a new host record for

southeastern Alaska. Haas ef al. (1980)

listed Sorex monticolus Merriam (as Sorex

vagrans Baird - see MacDonald and Cook

1996) as a host of H. o. occidentalis near

Yakutat. The two species of red-backed

voles, C. gapperi and C. rutilis, are also

new host records for H. o. occidentalis in

southeastern Alaska.

68 J. ENTOMOL. Soc. BRIT. COLUMBIA 102, DECEMBER 2005

CTENOPHTHALMIDAE

Catallagia charlottensis (Baker, 1898)

Material examined: Hosts were Pero-

myscus keeni unless otherwise indicated.

USA: AK: Gwent Cove, 54°57’00”N, 130°

20’00”W, 19 [AF26560], 17-viii-98, SOM.

Haines, Chilkoot Lake, 59°18’39”"N, 135°

34°02”W, 292 [AF4593], 11-vi-93, MJW.

Klukwan, 5 km W of, Klehini River, 59°

24’°39”N, 136°00’09"W, 19 ex Microtus

pennsylvanicus (Ord) [AF8036], 29-vi-94,

JAC. Mosquito Lake, 59°27’08”N, 136°

01°38"W, 299 [AF28822], 5-vi-99, AMR.

Reflection Lake, W side, 56°00’°33”N, 131°

34°32”W, 19 ex Clethrionomys gapperi

[AF29075], 30-vi-99, AMR. Rudyerd Bay,

Point Louise, 55°32’42”N, 130°52°13”W,

12 ex C. gapperi [AF29307], 10-vii-99,

AMR. Rudyerd Bay, 55°33’16’N, 130°

51°33”W, 19 [AF29381], 12-vii-99, AMR.

Smeaton Bay mouth, 55°18’09”N, 130°

50°38”W, 299 [AF29276], 9-vii-99, AMR.

Taku River, Canyon Island, 58°33’N, 133°

41’W, 14 [AF8274], JAC; same data but

192 [AF8276]. Turner Creek, 58°10’40’N,

133°57°30"°W, 192 [AF10126], 20-vii-94,

SOM, JAC, CTS. Walker Cove, Ledge

Point, 55°42’20”"N, 130°53’34"W, 1°

[AF29458], 14-v11-99, AMR.

The distribution patterns of this hy-

grophilous nest flea and of H. 0. occiden-

talis are similar (Holland 1963: Fig 2, 1985:

Map 14; Lewis et al. 1988: p. 82; Haas ef

al. 1989: Fig 2; Lewis and Haas 2001). In

southeastern Alaska both fleas have been

recorded from P. keeni (as P. maniculatus)

and M. oeconomus (nests) at mainland lo-

calities (Haas 1982; Haas et al. 1982; Hol-

land 1985). Our new records of these two

fleas and a record of C. charlottensis ex C.

rutilis in Haines (Holland 1985) linked the

fleas as parasites of C. rutilis, C. gapperi,

and M. pennsylvanicus on the mainland.

Thus far however, only C. charlottensis is

known from M. /ongicaudus in our survey

area (Haas et a/. 1982). Another similarity

in our records for H. 0. occidentalis and C.

charlottensis from trapped hosts was the

infrequency of more than a single specimen

collected per host. The larger number of H.

0. occidentalis specimens in nests of M.

oeconomus than on all trapped hosts applies

to C. charlottensis for its occurrence in the

same two nests found along the shore of the

Chilkat Peninsula. Both nests were infested

with breeding populations from which were

collected a total of 13¢¢ (four reared) and

1199 (three reared); an additional male

was collected from a third nest. Again,

more specimens (25) were collected from

nests (3) than those (15) from trapped hosts

12).

CERATOPHYLLIDAE

Ceratophyllus ciliatus protinus Jordan,

1929

Material examined: Hosts were Tamias-

ciurus hudsonicus (Erxleben) unless other-

wise indicated. USA: AK: Berg Bay, 56°

21°49"N, 132°00’29"W, 400, 299

[AF21810], 2-vin-97, CTS. Chickamin

River, Wolf Cabin, 55°46’N, 130°53’W,

14 [AF4930], 25-vii-93, SOM. Dyea Na-

tional Historical Park, 59°30’24”’N, 135°

20°52”W, 14, 299 [AF12532], 2-vii-95,

CTS. Gwent Cove, 54°57’°00”N, 130°

20°00”W, 204, 12 [AF26585], 19-viii-98,

SOM. Klukwan, 5 km W, Klehini River,

59°24’39"N, 136°00°09”W, 13 [AF8117],

l-vui-94, JAC. Peterson Creek, Juneau

Quad., 58°29°N, 134°477W, 19 ex

Clethrionomys rutilis [AF8243], 11-vu-94,

JAC. Rudyerd Bay, 55°33’16”’N, 130°

51°33°W, 229 [AF29409], 13-vii-99,

AMR. Taku River, Canyon Island, 58°

33’N, 133°41’W, 1% ex Peromyscus keeni

[AF8272], 17-vu-94, JAC; same data but

23 [AF8273]. Walker Cove, Hut Point,

55°42’48”N, 130°54’04”W, 12 ex P. keeni

[AF29442], 13-v1i-99, AMR.

This member of the Vancouverian group

(Holland 1963) has a typical Northwest

Pacific coast distribution similar to two

other members of the group, H. 0. occiden-

talis and C. charlottensis (Haddow et al.

1983: Map 17; Holland 1985: Map 71;

Lewis et al. 1988: p. 179). These authors

described the changes of preferred hosts

along the coast from south to north with

Townsend’s chipmunk (Neotamias town-

sendii (Bachman)) in Oregon, Douglas’s

squirrel (Zamiasciurus douglasii

(Bachman)) in southwestern British Colum-

J. ENTOMOL. SOc. BRIT. COLUMBIA 102, DECEMBER 2005

bia, and the red squirrel (7. hudsonicus) in

southeastern Alaska. Originally thought to

be the only truly specific flea of the red

squirrel in this area of Alaska, this view has

been modified as the result of collection of

the much wider-ranging true red squirrel

flea, Orchopeas caedens Jordan, along the

Taiya River in 1995 (see below).

Megabothris abantis (Rothschild, 1905)

Material examined: USA: AK: Bartlett

Cove, 10 km NW of Gustavus Airport, 58°

27N, 135°53’W, 14 ex Clethrionomys

rutilis [AF2379], 16-vii-92, collector un-

known. Berg Bay, 56°21°49°N, 132°

00°29”"W, 19 ex C. gapperi [AF21819], 4-

vill-97, CTS. Chickamin River, Wolf

Cabin, 55°46’N, 130°53’W, 1¢ ex Synap-

tomys borealis (Richardson) [AF4973], 26-

vul-93, SOM et al. Chilkat Peninsula, Mud

Bay, 59°09°45”N, 135°21’28”’W, 10, 19

ex Clethrionomys rutilis (2) [AF22019,

22020], 6-v11-97, CTS et al. Frosty Bay, S

side, 56°03’28”N, 131°58’01”W, 19 ex C.

gapperi [AF22904], 8-vii-97, CTS. Kluk-

wan, 5 km W of, Klehini River, 59°

24’39"N, 136°00°09"W, 2¢¢, 12 ex

Zapus hudsonius (Zimmerman) (2)

[AF8068, 8070], 30-vi-94, JAC. Nakat

Inlet, 54°57’N, 130°45’W, 19 ex C. gap-

peri [AF4265], 8-vu1-93, JAC, SOM. Rud-

yerd Bay, 55°33°16”’N, 130°51733”W,

22 2ex C. gapperi [AF29315], 10-vii-99,

AMR; same data but 192 [AF29375], 12-

vii-99; same data but 192 [AF29408], 13-

vul-99; same data but 55°41°58”N, 130°

31°12”W, 19 ex P. keeni [AF22589], 8-vi-

99, RH, AMR. Rudyerd Bay, Point Louise,

55°32°42”N, 130°52°13°W, 429 ex C.

gapperi (2) [AF29304, 29307], 10-v1i-99,

AMR. Salmon River, mouth of Texas

Creek, 56°01°37”N, 130°04’14”"W, 19 ex

P. keeni [AF12736], 2-viii-95, CTS. Smea-

ton Bay mouth, 55°18’09”"N, 130°

50°38°W, 146, 12 ex C.. gapperi

[AF29290], 10-vii-99, AMR; same date but

13 ex P. keeni [AF29292]. Stikine River,

Figure 8 Lake, 56°42’N, 132°15’W, 19 ex

P. keeni [AF2628], 14-vii-92, SOM; same

data but [AF2650], 15-vii-92. Taku River,

Canyon Island, 58°33’N, 133°41’W, 14 ex

C. gapperi [AF8254], 16-vii1-94, JAC; same

data but 19 [AF8270], 17-vii-94; same data

but 1d, 299 [AF8271]; same data but

229 ex M. pennsvivanicus [AF8268], 16-

vul-94. Turner Creek, 58°10°40”N, 133°

57°30°W, 2 2° ex P. keeni [AF10126], 20-

v1i-94, SOM, JAC, CTS; same data but 19

ex C. gapperi [AF10119]; same data but 19

ex Microtus longicaudus |AF10120]. Unuk

River mouth, 56°05’N, 131°06’W, 12 ex

C. gapperi [AF4359], 20-vi1-93, SOM et al.

Walker Cove, Hut Point, 55°42’48”N, 130°

54’04"W, 235, 12 ex P. keeni (2)

[AF29428, 29442], 13-vii-99, AMR; same

data but 12 ex C. gapperi [AF29434].

Walker Cove, Ledge Point, 55°42’20”N,

130°53°34"W, 3229 ex C._ gapperi

[AF29416], 13-vii-99, AMR. Willard Inlet,

inlet 2 km NW of mouth of, 54°49’N, 130°

39°W, 19 ex P. keeni [AF4299], 9-vii-93,

JAC, SOM. Yakutat, 59°30°47°N, 139°

40’°46”W, 19 ex C. rutilis [AF7769], 26-

vi1-94, CJC, AAT.

Megabothris abantis is a common vole

flea in southern regions of Alaska. Holland

(1958) originally grouped it with C. char-

lottensis and C. c. protinus because all three

have a similar Pacific Coast distribution.

Subsequently, he (Holland 1963: Fig. 2)

classified M. abantis as a member of the

Cordilleran Group B because it is not re-

stricted to the coastal strip but ranges

widely eastward into the Rocky Mountains

(Haddow eft al. 1983: Map 76, Holland

1985: Map 76).

Other than the closely grouped collec-

tion sites on the southeastern Alaska

mainland, Holland (1958), Haas eft al.

(1980), Haas (1982), and Haas ef al. (1982)

recorded only six other localities, all north-

west of the Taku River: Chilkat Peninsula,

near Juneau, Klondike Highway at Moore

Creek, Mosquito Lake, Taiya River, and

Yakutat. Twenty-nine of our 34 new

mainland records of M. abantis fill the large

void mapped by Haas et al. (1989: Fig. 5)

southeast of the Taku River with 16 new

localities.

The recorded hosts of M. abantis on the

mainland are Sorex monticolus (as S. va-

grans), P. keeni (as P. maniculatus), C.

rutilis, M. oeconomus (nests), M. longi-

70 J. ENTOMOL. SOC. BRIT. COLUMBIA 102, DECEMBER 2005

caudus and Z. hudsonius (Haas 1982; Haas

et al. 1982). Thus, three of the seven hosts

of our 34 new mainland records of M

abantis are new for the region: C. gapperi

(18), M. pennsylvanicus (1), and S. borealis

(1). Clethrionomys gapperi was the most

commonly collected mammal infested with

this flea and produced 5¢¢ and 2099

fleas, more than half of the 45 fleas (1204,

339°) collected. With the emphasis of the

mainland survey on areas south of Juneau,

only two C. rutilis with fleas (1¢, 19) were

trapped. The abundance on C. gapperi

alone, however, was concordant with the

classification of host parasitism by Haddow

et al. (1983) with the top ranking of mem-

bers of the genus Clethrionomys along with

Microtus as the only major hosts of this

flea. Our data for Microtus spp., however,

were insufficient for analysis with only

399 fleas ex one M. pennsylvanicus and

one M. longicaudus. Consequently, the sec-

ond best source of fleas from trapped hosts

was P. keeni with 11 fleas (3306, 899)

from nine mice. Although M. abantis is not

a nest flea, as indicated above with H. o.

occidentalis and C. charlottensis, more

specimens were found in a series of M.

oeconomus nests collected on the Chilkat

Peninsula shoreline (Haas 1982) than on a

larger number of trapped hosts. Four nests

were infested with M. abantis; of these, two

had breeding populations from which were

collected a total of 45 fleas (18¢¢ (9

reared), 2729 (13 reared)). With the addi-

tion of our specimens, 90 specimens have

been collected. The sex ratio of 1¢: 229

appears typical for M. abantis. Marshall

(1981) calculated 30% males in a sample of

456 specimens from trapped hosts in New

Mexico (Haas et al. 1973).

Megabothris asio megacolpus (Jordan,

1929)

Material examined: All ex meadow

voles, Microtus pennsylvanicus. USA: AK:

Klukwan, 11 km E & 12 km S, 59°

20°44"N, 135°46711”"W, 192 [AF8067], 30-

vi-94, JAC. Klukwan, 10 km E & 9 km S,

59°21°58”N, 135°47°58"W, 19 [AF8081],

30-vi-94, JAC; same data but [AF8164],

204, 19, 1-vii-94; same data but

[AF8165], 19; same data but [AF8185],

13, 2-vii-94.

Previously unknown in Alaska, M. a.

megacolpus was first collected (three 74,

four 9) in late June and early July 1994

by one of us (JAC) at two localities south-

east of Klukwan in one of the small exten-

sions of the range of the host, Microtus

pennsylvanicus, from Canada into south-

eastern Alaska. The range of the host in this

area, subspecies M. p. alcorni Baker, ex-

tends south from southwestern Yukon

across northwestern British Columbia and

into the Chilkat River valley of Alaska as

far south as Haines (Miller and Kellogg

1955).

The distribution of this vole flea coin-

cides almost completely with the range of

M. pennsylvanicus over much of northern

North America from Yukon Territory to

Quebec south into the Rocky Mountains

and the western Great Lakes (Haddow et al.

1983: Map 82; Holland 1985: Map 77). The

Stikine and Taku River valleys on the

southeastern Alaskan mainland also support

populations of M. pennsylvanicus

(MacDonald and Cook 1996) and may

eventually yield additional specimens of M.

a. megacolpus.

Malaraeus telchinus (Rothschild, 1905)

Material examined: Hosts Peromyscus

keeni except as indicated. USA: AK: Gwent

Cove, 54°57°00”N, 130°20’00”W, 346

[AF26560], 17-vii-98, SOM. Rudyerd Bay,

55°33°16"N, 130°51733"W, 19

[AF29317], 10-vii-99, AMR; 14, 19

[AF29361], 11-vii-99, AMR; 299

[AF29379]; 299 [AF29381]; 19

[AF29382], 12-vii-99, AMR; same locality

but 19 ex C. gapperi [AF29375], 12-vii-99,

AMR; same data but [AF29378]. Rudyerd

Bay, Point Louise, 55°32’42”N, 130°

53°34°W, 14, 222 [AF29372], 12-vii-99,

AMR. Walker Cove, Ledge Point, 55°

42’20”N, 130°53’34”"W, 19 [AF29417],

13-vii-99, AMR; 14, 399 [AF29458], 14-

vii-99, AMR; same locality but 19 ex C.

gapperi [AF29416], 13-vii-99, AMR.

The western vole and mouse flea,

telchinus, eluded detection in Alaska until

334 were collected from one P. keeni at

J. ENTOMOL. SOc. BRIT. COLUMBIA 102, DECEMBER 2005

Gwent Cove (across Pearse Canal from

Pearse Island, British Columbia) in 1998.

The next two localities were farther north at

Rudyerd Bay where 24:4 and 999 were

collected ex six P. keeni, and 299 ex 2 C.

gapperi. The collector (AMR) then moved

north to Ledge Point on the south shore of

the mouth of Walker Cove and established

the most northern locality for M. te/lchinus

in North America with 1% and 49 9 ex two

P. keeni and 19 ex C. gapperi. This new

locality is ca. 236 km northwest of Kitimat,

the most northern mainland locality in Brit-

ish Columbia (Holland 1985). Holland

(1949, 1985) listed other M. telchinus off-

shore records close to southeastern Alaska

on the Queen Charlotte Islands and Pitt

Island. Most of his many records were clus-

tered farther southeastward in British Co-

lumbia at inland montane and coastal lo-

calities (Holland 1985: Map 84). South of

Canada these populations diverge into a

coastal branch that reaches southern Cali-

fornia and a montane branch that almost

bypasses the Great Basin to reach the mesic

habitats on mountains and high plateaus in

Arizona and New Mexico (Haddow ef al.

1983: Map 74).

Malaraeus telchinus is recorded from a

wide range of hosts. Haddow et al. (1983)

listed four Peromyscus species as the major

hosts of M. telchinus but omitted P. sitken-

sis Merriam (= P. keeni, see Hogan et al.

1993); no Clethrionomys species were

listed. Holland (1949, 1985) only listed four

records from C. gapperi including one for

Kitimat. The great majority of hosts on the

mainland were P. maniculatus with P. keeni

in the Queen Charlotte Islands. Clethriono-

mys californicus (Merriam) is a major host

in Oregon; Lewis et al. (1988) reported

more specimens of M. telchinus from this

vole than from each of two Peromyscus

species, three Microtus species, and Lem-

miscus curtatus (Cope). This wide host

range of M. telchinus confirms adaptability

for changing major hosts when entering a

region with a different fauna of potential

hosts (e.g. moving from mainland British

Columbia and P. maniculatus to mainland

southeastern Alaska and P. keeni). We ex-

pect that M. te/chinus occurs north of its

present known northern range limit of

Walker Cove because P. keeni occurs on

the mainland north to Haines and Skagway

(MacDonald and Cook 1996) and this flea

occurs in “rather mesic habitats” elsewhere

(Haddow et al. 1983: p. 108).

Opisodasys vesperalis (Jordan, 1929)

Material examined: All ex northern fly-

ing squirrels, Glaucomys sabrinus (Shaw).

USA: AK: Chilkat River, 6.3 km WNW of

Haines, 59°15’42”N, 135°33’35”"W, 24,

12 [AF12539], 4-vii-95, CTS. Rudyerd

Bay, 55°33°16”N, 130°57°33”W, 242,

3292 [AF29318], 10-vii-99, AMR. St.

James Bay, W _ side Lynn Canal, 58°

34’30"N, 135°09’30"W, 10, 299

[AF10306], 9-1-95, JF.

Glaucomys sabrinus, the host of O. ves-

peralis, is found along the mainland of

southeastern Alaska, and on islands in the

Alexander Archipelago south of Frederick

Sound (MacDonald and Cook 1996). Re-

cords of its fleas are few with only one for

the mainland and one for Revillagigedo

Island (Haas et al. 1982, 1989: Fig. 7). Our

three new records are the first for trapped

hosts (the mainland collection near Skag-

way was from a nest). Lacking a pleural

arch, O. vesperalis is a crawling nest flea

(rather than a jumping flea) and remains in

the nest when the host is absent (Traub

1972). Collections from other mammals are

strictly accidental and rare; the flea is

“essentially specific to Glaucomys_ sabri-

nus” (Haddow et al. 1983: p. 130). Opiso-

dasys vesperalis is among the minority of

fleas considered to be “ultraspecific’’, 1.e.,

“limited to infestation of a single species of

host” (Traub 1985: p. 332). It 1s a west

coastal flea ranging from southeastern

Alaska to northern California and eastward

into montane squirrel habitat as far as Idaho

and Montana (Haddow et al. 1983: Map

107; Holland 1985: Map 89; Lewis ef al.

1988: p. 201; Haas et al. 1989: Fig 7).

Opisodasys keeni keeni (Baker, 1896)

Material examined: USA: AK:

Chickamin River, Wolf Cabin, 55°46’N,

130°53’W, 34.4, 192 ex Peromyscus keeni

[AF4953], 26-vi1-93, SOM et al.; same data

i2 J. ENTOMOL. Soc. BRIT. COLUMBIA 102, DECEMBER 2005

but 12 ex Synaptomys borealis [AF4973].

Crescent Lake, 58°11’N, 133°19’W, 52,

729 ex P. keeni (4) [AF8309, 8310, 8311,

8316], 22-vii-94, SOM. Echo Cove, 58°

31°45”N, 134°54’28"W, 20, 529 ex P.

keeni [AF21759], 21-vi-97, CTS. Gwent

Cove, 54°57’00”N, 130°20°00”"W, 302,

492° ex P. keeni [AF26560], 17-viii-98,

SOM. Reflection Lake, SW _ side, 55°

59°59"N, 131°33’59”"W, 53.3, 429 ex P.

keeni [AF29116], 1-vii-99, AMR. Rudyerd

Bay, 55°18’09”N, 130°50’°38”W, 14, 49°

ex P. keeni (2) [AF29317, 29320], 10-vii-

99, AMR; same data but 299 (2)

[AF29360, 29361], 11-vii-99; same data but

43g, 1199 (5) [AF29379, 29380, 29381,

29382, 29383], 12-vi1-99; same data but

34.4, 19 (2) [AF29407, 29408], 13-vii-99.

Rudyerd Bay, Point Louise, 55°32°42”N,

130°52’13”W, 23 Sex P. keeni [AF29372],

12-vii-99, AMR. Smeaton Bay mouth, 55°

18’09”N, 130°50’38”"W, 10, 329 ex P.

keeni (3) [AF29276, 29278, 29279], 9-vii-

99, AMR; same data but 4¢¢, 19

[AF29292], 10-vu-99. Smeaton Bay, E

Skull Creek, 55°17’°27”N, 130°49°25”W,

1d, 19 ex P. keeni [AF29283], 9-vii-99,

AMR. Stikine River, Figure 8 Lake, 56°

42’N, 132°15’°W, 14, 19 ex P. keeni

[AF2627], 14-vii-92, SOM; same data but

229 [AF2628]; same data but 19

[AF2650], 15-vii-92, Taku River, Canyon

Island, 58°33’N, 133°41’W, 1 ex P. keeni

[AF8272], 17-vi1-94, JAC et al.; same data

but 266, 229 [AF8273]. Unuk River

mouth, 56°05’N, 131°06’W, 23°97, 229 ex

P. keeni (3) [AF4355, 4360, 4428], 20-21-

vul-93, SOM et al. Walker Cove, Hut Point,

55°42°48”N, 130°54’04”"W, 3404, 229 ex

P. keeni (3) [AF29428, 29438, 29442], 13-

vil-99, AMR. Walker Cove, Ledge Point,

55°42’20”N, 130°53’34”W, 1¢ ex P. keeni

[AF29458], 14-vii-99, AMR.

This common and abundant Peromyscus

flea is another member of the Vancouverian

Group (Holland 1963) with a west coastal

distribution including the Queen Charlotte

Islands, extending eastward into montane

habitats of its major host P. maniculatus in

British Columbia and Alberta (Holland

1949:Map 28, 1985: Map 88). Southeast of

Canada O. k. keeni occurs in Montana,

Utah, Colorado and New Mexico (Ecke and

Johnson 1952; Stark 1959; Eads and Cam-

pos 1983; Haddow et al. 1983: Map 108;

Fagerlund ef a/. 2001; Ford ef al. 2004). Its

coastal range extends from northern Cali-

fornia as far north as Skagway, Alaska

(Haas et al. 1982; Haddow ef al. 1983: Map

108; Lewis et a/. 1988: p. 203). In British

Columbia, O. k. keeni and M. telchinus

have the same host, P. maniculatus (P.

keeni in the Queen Charlotte Islands), and

distribution (Holland 1985: Maps 84 & 88).

Host sharing by these fleas also occurs in

southeastern Alaska: with M. telchinus, P.

keeni was host for nine collections, C. gap-

peri for three; with O. k. keeni, P. keeni was

host for 37 collections (434.4, 5599) and

S. borealis, a new host record, was host for

one (Q).

Orchopeas caedens (Jordan, 1925)

Material examined: USA: AK: 500 m S

of Taiya River bridge, 59°30’11’°N, 135°

20°44”"W, 19 ex Tamiasciurus hudsonicus

[AF12525], 1-vu-95, CTS, KDS.

The collection of 192 of the common,

“ultraspecific” (Traub 1985) red squirrel

flea, O. caedens, in southeastern Alaska

was unexpected. The niche was already

filled by C. ciliatus protinus, an arboreal

squirrel flea well-adapted to the west

coastal maritime climate on the coast and

islands of British Columbia and north

through the length of southeastern Alaska.

Until now, there has been no record of O.

caedens within the range of the red squirrel

in this region. Elsewhere, O. caedens 1s

found throughout most of the transcontinen-

tal range of the red squirrel and occurs with

other red squirrel fleas, such as Ceratophyl-

lus vison Baker and Tarsopsylla octodecim-

dentata coloradensis (Baker) in nests in

Alaska west of the Yukon Territory (Haas

and Wilson 1982). The limiting factor for

O. caedens in southeastern Alaska 1s proba-

bly the high humidity and precipitation of

the coastal climate. The Taiya River valley

lies in the area of lowest mean annual pre-

cipitation in southeastern Alaska, 1.e., less

than 101.6 cm (Watson 1959).

J. ENTOMOL. Soc. BRIT. COLUMBIA 102, DECEMBER 2005

DISCUSSION

Nineteen mammal fleas have been docu-

mented for southeastern Alaska (Haas ef ai.

1989). Three of these are known only from

islands: the bat flea, Myodopsylla gentilis

Jordan and Rothschild, and the bear flea,

Chaetopsylla_ tuberculaticeps (Bezzi), on

Admiralty and Chichagof Islands (Haas ef

al. 1979, 1980, 1989) and the dog flea,

Ctenocephalides canis (Curtis), on Revilla-

gigedo Island (Holland 1985: p. 38). Our

three additions (M. asio megacolpus, M.

telchinus and O. caedens) bring the

mainland total to 19 taxa. Because of the

high humidity and precipitation in the area,

these new records (especially that of O.

caedens) were not expected.

We have tried to collect the northern

Peromyscus. maniculatus flea, Aetheca

thamba (Jordan), in the Klondike Highway

pass at the Alaska/British Columbia border

without success. Holland (1949: Map 40,

1958: Fig. 5, 1985: Map 73) mapped and

discussed the split distribution pattern of

transcontinental A. thamba (then Monopsyl-

lus thambus). Most of the many records

range from southern Yukon Territory (e.g.,

1.6 km S of Carcross) eastward into north-

em British Columbia (e.g., Atlin) to north-

ern and southern Alberta, northwestern Sas-

katchewan, and southwestern Northwest

Territories. A small disjunct population

exists in Quebec and Labrador. The prox-

imity of many collections of a cold climate

flea from a common and abundant host are

conditions favourable for the collection of

A. thamba in the northern Alaska/British

Columbia border passes.

The bushy-tailed woodrat, Neotoma

cinerea (Ord), is one of several wide-

ranging mammals of British Columbia that

has established itself in corridors of some

major rivers such as the Taku, Stikine and

Unuk that transect the Alaska/British Co-

lumbia border mountains (MacDonald and

Cook 1996). The common woodrat flea,

Orchopeas agilis (Rothschild) (formerly O.

sexdentatus agilis), is the most widespread

member of the sexdentatus group. It ranges

from high mountains with the cool sum-

mers required by N. cinerea in New Mexico

and Colorado northwestward to British Co-

lumbia (e.g., Atlin) and questionably

Yukon Territory (Finley 1958; Haas et al.

1973; Holland 1985: Map 94; Lewis 2000;

Haas et al. 2004). Haddow ef al. (1983:

Map 115) mapped two localities on the

Alaska/British Columbia border but none in

Alaska. We believe O. agilis probably oc-

curs on the southeastern Alaskan mainland

where N. cinerea occurs. Well sheltered

nests should be good sources of fleas in that

region.

Catallagia ioffi Scalon (formerly C.

jellisoni Holland; see Lewis and Haas

2001) is an uncommonly collected, Holarc-

tic mammal flea with only six known lo-

calities in Canada scattered from near Daw-

son in Yukon Territory southeastward

through British Columbia (e.g., Atlin) to

Banff National Park, Alberta (Holland

1954; Hopkins and Rothschild 1962; Haas

and Johnson 1981; Holland 1985 :Map 17).

The likelihood of collecting specimens on

the southeastern Alaskan mainland is sug-

gested by this elongated distribution pattern

in western Canada as well as the diversity

of known hosts, e.g., P. maniculatus, N.

cinerea, C. rutilis, C. gapperi, M. pennsyl-

vanicus and Lemmus — trimucronatus

(Richardson). The latter species has not

been recorded for southeastern Alaska

(MacDonald and Cook 1996).

ACKNOWLEDGEMENTS

We thank C. J. Conroy, J. Foreit, R.

Heinen, S. R. Peterson, C. T. Seaton, K. D.

Stone, A. A. Tsvetkova and M. J. Wike for

collecting, labeling and preserving the flea

specimens regardless of difficult field con-

ditions. We thank the US Fish and Wildlife

Service, USDA Forest Service, and Alaska

Department of Fish and Game for financial

and logistic support for the fieldwork. Mi-

chael K. MacDonald kindly reviewed the

manuscript.

74 J. ENTOMOL. Soc. BRIT. COLUMBIA 102, DECEMBER 2005

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Haddow, J., R. Traub and M. Rothschild. 1983. Distribution of ceratophyllid fleas and notes on their hosts,

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Ceratophyllidae: key to the genera and host relationships. Privately published by M. Rothschild and R.

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implications of karyotypic, electrophoretic, and mitochondrial-DNA variation in Peromyscus from the

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70:1-306.

Holland, G.P. 1954. A new species of Catallagia Rothschild from the Rocky Mountains of Alberta

(Siphonaptera: Hystrichopsyllidae: Neopsyllinae). The Canadian Entomologist 86: 381-384.

Holland, G.P. 1957. Notes on the genus Hystrichopsylla Rothschild in the New World, with descriptions of

one new species and two new subspecies (Siphonaptera: Hystrichopsyllidae). The Canadian Entomolo-

gist 89: 309-324.

Holland, G.P. 1958. Distribution patterns of northern fleas (Siphonaptera), pp. 645-658. /n Proceedings,

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sity Press, Cambridge.

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(Siphonaptera: Ceratophyllidae: Ceratophyllinae). Journal of Vector Ecology 25: 164-189.

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description of a new species (Siphonaptera: Ctenophthalmidae: Neopsyllinae: Phalacropsyllini). Journal

of Vector Ecology 26: 51-69.

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MacDonald, S.O. and J.A. Cook. 1996. The land mammal fauna of Southeast Alaska. Canadian Field-

Naturalist 110: 571-598.

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Stark, H.E. 2002. Population dynamics of adult fleas (Siphonaptera) on hosts and in nests of the California

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Traub, R. 1972. Notes on zoogeography, convergent evolution and taxonomy of fleas (Siphonaptera), based

on collections from Gunong Benom and elsewhere in south-east Asia. II. Convergent evolution. Bulle-

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United States No. 60-49. U.S. Department of Commerce, Washington, DC.

J. ENTOMOL. SOc. BRIT. COLUMBIA 102, DECEMBER 2005

SCIENTIFIC NOTE

Infestation of Bent Grass by a New Seed Pest, Chirothrips

manicatus (Thysanoptera: Thripidae), in Oregon

SUJAYA RAO! and STEPHEN C. ALDERMAN’

Bent grasses (Agrostis spp.) (Poaceae:

Pooideae) are used extensively on golf

courses. Oregon is the largest bent grass

seed producing state in the U.S.A., with

over 3,900 ha under cultivation of Agrostis

stolonifera L. (= Agrostis palustris Hud-

son) (creeping bent grass), Agrostis castel-

lana Boissier & Reuter (dryland or High-

land bent grass) and Agrostis capillaris L.

(= Agrostis tenuis Sibthorp) (colonial bent

grass) (USDA-ODA 2002).

In June 2004 Chirothrips manicatus

Haliday (Thysanoptera: Thripidae) were

detected within individual florets of

Agrostis spp. in Oregon. Voucher speci-

mens were deposited in the Oregon State

Entomology Museum, Corvallis, OR.

Each floret with a thrips produced no seed.

Chirothrips manicatus 1s widespread in

North America. In Oregon it has been col-

lected from flowers of various plants (Post

1947), but this is the first report of it devel-

oping in florets of Agrostis. It has been

reported on A. fenuis in New Zealand

(Mound and Walker 1982) and on Agrostis

sp. in Europe (zur Strassen 2003), but there

is no information on its impact on seed

production on these hosts. It 1s reported as

a pest of orchard grass in New Zealand

(Doull 1956).

To determine the extent to which C.

manicatus was present in commercial

Agrostis seed production fields in Oregon,

we surveyed 13 bent grass seed production

fields in July 2004 (Table 1). The fields

were located in the Silverton Hills area in

Table 1.

Incidence of Chirothrips manicatus in Agrostis seed production fields in the Willamette Valley

in western Oregon. Means + SE are based on collections of 17 to 50 panicles from each of four

transects in a diamond pattern in each field.

Field Agrostishost Cultivar Mean % Meanno. Meanno. Mean % seed

panicles thrips per seeds per loss due to

with thrips panicle panicle thrips

] A. castellana Highland 87.9+4.1 DAE tee a 452 + 105 5.1+1.4

2 A. castellana Highland 49.8+7.3 6.8 + 0.9 426 + 53 0.9+0.2

3 A. castellana Highland 60.0 + 5.7 10.315 526 + 55 L1+04

4 A. castellana Highland 26.2 +5.4 54 2.5 235 220 1.34 0.7

5 A. castellana Highland 8052 7.1 17.9+2.3 3792 11 4.1+1.8

6 A. castellana Highland 32.9+4.3 23232 471 +10 10+0.4

f| A. castellana Highland 648+ 10.1 17.6220 467+9 2.5205

8 A. castellana Highland 38.0278 6.0+ 0.7 Zi te2Z 0.8 + 0.3

9 A. stolonifera | Crenshaw 2.0+ 1.4 4842.5 528 + 44 0.02 + 0.01

10 A. stolonifera Princeville 9523.1 1440.1 452.222 0.03 + 0.01

1] A. stolonifera Pennlinks 0 0 a2 ead 0

12 A. stolonifera _ Penncross 0 0 292212 0

13 A. capillaris Alistar 0.5.24 0.5 0.03’ 425+ 13 0.01 + 0.01

Five individuals in a single panicle

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

* USDA-ARS, National Forage Seed Production Research Center, Corvallis, OR 97331

> Author to whom correspondence should be addressed

78 J. ENTOMOL. SOc. BRIT. COLUMBIA 102, DECEMBER 2005

the Willamette Valley in western Oregon.

In each field, 17 to 50 panicles were col-

lected at random along each of four tran-

sects in a diamond pattern. Panicles were

examined under a stereo microscope and

the number of thrips recorded (Table 1).

Seeds from each panicle were threshed by

hand to avoid seed loss. Caryopses were

separated from the lemma and palea using

a scarifier (Model PSS1000, Mater Interna-

tional, Inc., Corvallis, OR) and debris was

removed with an air column (Alderman ef

al. 2003). Total seed weight from each

transect was determined and the weight of

a subset of 200 seeds from each transect

was used to estimate the total number of

healthy seeds in panicles collected from

each transect. Percentage seed loss was

estimated as: [number of infested seeds /

(number of infested seeds + estimated

number of healthy seeds)| x 100, where the

number of infested seeds equals the num-

ber of thrips, based on our observation of

one thrips per floret and destruction of a

single seed by each thrips.

Overall, 32.5% of 2,310 panicles from

the 13 fields that were examined were 1n-

fested with thrips and the abundance of the

thrips appeared to be linked to the host

(Table 1). The greatest infestation was ob-

served in Highland bent grass (A. castel-

lana) which is the most common cultivar

grown for seed in Oregon. Individual bent

grass florets contained a single C. manica-

tus pupa or adult (apterous male or winged

female) with its head towards the base of

the floret (Figure 1). The thrips were en-

closed firmly between the lemma and the

palea, and were not easily dislodged. In

florets where a thrips was present, organic

debris was visible but there was no trace of

the caryopsis (seed). These data negate

previous speculation that small seeded

plants such as Agrostis spp. are unlikely

hosts for C. manicatus (Doull 1956).

It is not known how long C. manicatus

has been present on Agrostis in Oregon. As

there is no external indication of C. mani-

catus, it is possible that its presence could

have been undetected. Female C. manica-

tus overwinter within florets in the field

(Doull 1956) and therefore it is also possi-

ble that C. manicatus has emerged as a pest

due to the phase-out of field burning in the

late 1980’s.

We thank B. Matson; G. Gingrich; Ore-

gon bent grass seed producers; the USDA-

ARS Systematic Entomology Laboratory,

Beltsville, MD, for identification of C.

manicatus; L. Mound, CSIRO, Australia,

for discussions; and T. Cook, R. Halse,

W.P. Stephen and L.A. Mound for review-

ing the manuscript.

Figure 1. Chirothrips manicatus within a floret of Agrostis castellana.

REFERENCES

Alderman, S.C., D.M. Bilsland, J.A. Griesbach, G.M. Milbrath, N.W. Schaad and E. Postnikova. 2003. Use of a

seed scarifier for detection and enumeration of galls of Anguina and Rathayibacter species in orchard grass

seed. Plant Disease 87: 320-323.

Doull, K.M. 1956. Thrips infesting cocksfoot in New Zealand. II. The biology and economic importance of the

cocksfoot thrips Chirothrips manicatus Haliday. New Zealand Journal of Science and Technology 38: 56-65.

Mound, L.A. and A.K. Walker. 1982. Terebrantia (Insecta: Thysanoptera). Fauna of New Zealand 1: 1-113.

Post, R.L. 1947. Thysanoptera in Oregon. PhD. dissertation, Oregon State University, Corvallis, U.S.A.

USDA-ODA. 2002. U. S. Department of Agriculture - Oregon Department of Agriculture. 2001-2002 Oregon

Agriculture & Fisheries Statistics, Oregon Agricultural Statistic Service, Portland, Oregon, U.S.A.

zur Strassen, R. 2003. Die terebranten Thysanopteren Europas und des Mittelmeer-Gebietes. Die Tierwelt

Deutschlands. 74. Teil. [The terebrantian Thysanoptera of Europe and the Mediterranean region. The animal

world of Germany. Part 74.]. Goecke & Evers, Keltern, Germany.

J. ENTOMOL. SOC. BRIT. COLUMBIA 102, DECEMBER 2005

SCIENTIFIC NOTE

Comparative Activity of the Codling Moth Granulovirus

Against Grapholita molesta and Cydia pomonella

(Lepidoptera: Tortricidae)

LAWRENCE A. LACEY'”, STEVEN P. ARTHURS!

and HEATHER HEADRICK'

ABSTRACT— The granulovirus of codling moth, Cydia pomonella L., CpGV, is now com-

mercialized for codling moth control in pome fruit in the USA and Canada. It is highly spe-

cific for codling moth and related species. Comparative assays of CpGV against neonate

larvae of another introduced tortricid pest, the oriental fruit moth, Grapholita molesta Busck,

revealed a 557 and 589 fold lower susceptibility of neonate larvae compared with the LCs

and LCos; values derived for C. pomonella.

Since its introduction into North America,

the oriental fruit moth, Grapholita molesta

Busck, has become a widely established pest of

peach, nectarine, apricot, and apple (Rothschild

and Vickers 1991). There is little information

regarding naturally occurring disease of the

oriental fruit moth, with the exception of mi-

crosporidia in adults (Simchuk and Komarova

1983) and Bacillus thuringiensis in larvae

(Grassi and Deseé 1984). Although field trials

with various formulations of B. thuringiensis

have been reported for oriental fruit moth, re-

sults indicate that it is relatively ineffective

(Rothschild and Vickers 1991).

Following its initial discovery in infected

codling moth Cydia pomonella L. larvae in

Mexico in 1964, numerous laboratory and field

studies have confirmed the virulence of the

codling moth granulovirus (CpGV), against its

homologous host (Falcon et al. 1968, Laing

and Jaques 1980, Arthurs and Lacey 2004,

Cossentine and Jensen 2004). In early host

specificity studies, CpGV was also noted to

have larvicidal activity against the pea moth,

Cydia nigricana (Fabricius) (Payne, 1981) and

oriental fruit moth (Falcon ef al. 1968), but

quantitative assays of the virus have not been

reported for the latter species. We conducted

bioassays of the Cyd-X formulation of CpGV

(Certis USA, Columbia, MD) against oriental

fruit moth and codling moth neonates from

colonies maintained at the Yakima Agricultural

Research Laboratory using the materials and

methods described by Lacey et al. (2002). The

codling moth diet described by Brinton et al.

(1969) (BioServ, Frenchtown, NJ, USA) was

used for both species.

Following initial bioassays to determine

mortality ranges, five concentrations of Cyd-X

that produced mortality in neonate larvae rang-

ing from 10 to 96.7% in oriental fruit moth and

36.7 to 96.7% in codling moth were bioassayed

against 30 neonates per concentration. Bioas-

says were conducted on artificial diet in 2-ml

plastic conical autosampler vials (Daigger,

Lincolnshire, IL, USA). A 2-mm diameter hole

in the cap of each vial covered with stainless

steel screen (0.16 mm mesh size) eliminated

condensation. Ten ul of aqueous virus suspen-

sions or 10 ul of water for controls was applied

to the surface of 1 ml of artificial medium

(approximately 100 mm”) in the autosampler

vials. The label specified virus concentration of

Cyd-X is 3 x 10’° granules per liter. After the

surface of the medium had dried, one neonate

larva was added to each vial. The vials were

incubated for 7 d at 25 + 1.7 °C and then as-

sessed for larval mortality. The study was re-

peated for each species on four separate dates.

Each date was treated as an individual replicate

of each concentration (i.e. data were not pooled

before probit analysis).

The results of the assays clearly indicated

that oriental fruit moth neonates are susceptible

to CpGV, but at a significantly lower level than

that observed in codling moth neonates (Table

1). The oriental fruit moth were 557 and 589

times less susceptible to CpGV compared with

codling moth, based on probit (normal sig-

moid) analysis of the LCs) and LCos, respec-

' Yakima Agricultural Research Laboratory, USDA-ARS, 5230 Konnowac Pass Rd., Wapato, WA 98908

* Author to whom correspondence should be addressed

80 J. ENTOMOL. SOC. BRIT. COLUMBIA 102, DECEMBER 2005

tively (StatsDirect Ltd, v. 2.4). Based on our

methods, the calculated LCs) and LCo; of

CpGV for oriental fruit moth are 35 and 540

granules per mm’, respectively, but only 0.06

and 0.9 granules per mm” for codling moth.

Although CpGV must be ingested in order to

infect a larva, Ballard et al. (2000) demon-

strated that the point of entry of codling moth

larvae into fruit may not necessarily be where

virus is acquired; larvae could become infected

by walking or browsing on CpGV-sprayed leaf

surfaces in as little as 3.5 min. Ostensibly virus

picked up on legs or mouth parts in the absence

of browsing leaf surfaces could contaminate

the initial point of entry. In our bioassays, neo-

nates of both species may wander over the

surface of the medium before boring into it. In

the case of codling moth larvae this would

provide ample opportunity to acquire virus

even at the lower concentrations. Huber (1986)

estimated that the LDs, for neonate larvae

could be as low as 1.2 granules per larva.

First generation oriental fruit moth often

feed on shoots and young foliage. Although not

as active against oriental fruit moth as against

codling moth in laboratory bioassays, field

activity of Cyd-X against oriental fruit moth

neonates at label rates used for codling moth

control (0.07-0.44 L/ha) could potentially re-

duce oriental fruit moth populations if signifi-

cant feeding of early instars of the first genera-

tion occurred on treated foliage. Because natu-

ral feeding behavior will influence their sus-

ceptibility to CpGV, further field studies are

warranted.

We are grateful to Rob Fritts Jr. (Certis) for

Cyd-X samples and the Washington Tree Fruit

Research Commission for financial support.

We thank Don Hostetter and Joel Siegel for

helpful reviews of the manuscript.

Table 1.

LCso and LCo; values for CpGV bioassayed against Grapholita molesta and Cydia pomonella neo-

nates. The number of granules per 10 ul is based on dilutions of Cyd-X with a label specified virus

concentration of 3 x 10° granules/L. All probit comparisons were significantly different, P < 0.001

based on dosage log(;0+;) and adjusted for control mortality (<< 7.1%).

Species n LCs) (95% CI) LCys (95% CI) Slope

C. pomonella 714 6.30 (4.74 — 7.91) 91.62 (63.00 — 155.68) 1.41

G. molesta 708 351 x10 (272-4422 x10) 5.40 x 10° (3.66 —9.04 x 10°) 1.39

REFERENCES

Arthurs, S.P. and L.A. Lacey. 2004. Field evaluation of commercial formulations of the codling moth granulovi-

rus (CpGV): persistence of activity and success of seasonal applications against natural infestations in the

Pacific Northwest. Biological Control 31: 388-397.

Ballard, J., D.J. Ellis, and C.C. Payne. 2000. Uptake of granulovirus from the surface of apples and leaves by first

instar larvae of the codling moth Cydia pomonella L. (Lepidoptera: Olethreutidae). Biocontrol Science and

Technology 10: 617-625.

Brinton, F.E., M.D. Proverbs, and B.E. Carty. 1969. Artificial diet for mass production of the codling moth, Car-

pocapsae pomonella (Lepidoptera: Olethreutidae). The Canadian Entomologist 101: 577-584.

Cossentine, J.E. and L.B.M. Jensen. 2004. Persistence of a commercial codling moth granulovirus product on

apple fruit and foliage. Journal of the Entomological Society of British Columbia 101: 87-92.

Falcon, L.A., W.R. Kane, and R.S. Bethell. 1968. Preliminary evaluation of a granulosis virus for control of cod-

ling moth. Journal of Economic Entomology 61: 1208-1213.

Grassi, S. and K.V. Deseé. 1984. Distribution of Bacillus thuringiensis Berl. and prospects of using it in plant

protection. Atti Giornate Fitopatologiche 2: 425-433 (in Italian). Review of Applied Entomology 73: 361.

Huber, J. 1986. Use of baculoviruses in pest management programs, pp. 181-202. Jn R.R. Granados and B.A.

Federici (eds), The Biology of Baculoviruses, Vol. II: Practical Application for Insect Control. CRC Press,

Boca Raton, FL.

Lacey, L.A., P.V. Vail, and D.F. Hoffmann. 2002. Comparative activity of baculoviruses against the codling moth,

Cydia pomonella, and three other tortricid pests of tree fruit. Journal of Invertebrate Pathology 80: 64-68.

Laing, D.R. and R.P. Jaques. 1980. Codling moth: Techniques for rearing larvae and bioassaying granulosis virus.

Journal of Economic Entomology 73: 851-853.

Payne, C.C. 1981. The susceptibility of the pea moth, Cydia nigricana, to infection by the granulosis virus of the

codling moth, Cydia pomonella. Journal of Invertebrate Pathology 38: 71-77.

Rothschild, G.H.L. and R.A. Vickers. 1991. Biology, ecology and control of the oriental fruit moth, pp. 389-412.

In L.P.S. van der Geest and H.H. Evenhuis (Eds.), Tortricid Pests, Their Biology, Natural Enemies and

Control. Elsevier Science Publishers, Amsterdam, The Netherlands.

Simchuk, P.A. and G.F. Komarova. 1983. Microsporidiosis in the oriental peach moth. Zashchita Rasteni 29 (in

Russian); Reviews of Applied Entomology 72: 471 (in English).

J. ENTOMOL. Soc. BRIT. COLUMBIA 102, DECEMBER 2005

SCIENTIFIC NOTE

A novel host association for Monarthrum scutellare

(Coleoptera: Curculionidae: Scolytinae) in British Columbia

LELAND M. HUMBLE’

Monarthrum scutellare (LeConte) is an

ambrosia beetle that ranges from British

Columbia to northern Baja California in

Mexico (Bright and Stark 1973, Wood

1982, Wood and Bright 1992). It is re-

corded to breed in various species of Fa-

gaceae including Chrysolepis, Lithocarpus

densiflora (Hooker & Arnold) Rehder,

Quercus spp., Quercus agrifolia Neé, Q.

garryana Douglas and Q. kellogi Newberry

(Farris 1965, Bright and Stark 1973, Bright

1976, Wood and Bright 1992) with single

records from Abies and sequoia that Bright

and Stark (1973) considered accidental or

erroneous.

On 2 May 2005, a 38-cm length of

“green” split alder firewood and associated

Scolytinae (Coleoptera: Curculionidae)

collected from a recently delivered com-

mercial load of firewood were submitted to

the Canadian Forest Service for identifica-

tion after beetles were observed emerging

from the wood. The half stem section was

split off-centre, included all annual growth

rings, and was 18.2 cm in diameter and 27

years of age. No bark was present on the

piece of firewood, however, a single V-

shaped parental gallery 22 mm in length

was incised in the sapwood and five adult

Alniphagus aspericollis (LeConte)

(Curculionidae: Scolytinae) were associated

with the sample. The gallery shape agrees

with those described by Bright and Stark

(1973) as typical for A. aspericollis. The

presence of a parental gallery of A. asperi-

collis and the structure of the wood

(absence of rays, ring porous wood) con-

firmed that the host attacked was Alnus

rubra Bongard (Betulaceae).

Boring dust was being actively extruded

from ambrosia beetle galleries along the

split face of the wood; however, no en-

trance holes were observed on the outer

face of the bole. The wood was held at

room temperature for adult emergence and

52 female and 55 male M_ scutellare

emerged between 2 May and 17 May 2005.

The sample was then split longitudinally

and the distribution of galleries along the

split face enumerated by growth ring and

growth ring widths measured to the nearest

0.5 mm. All of the 22 M. scutellare galler-

ies visible on the split face were in the wid-

est growth rings (mean + SD = 5.6 + 0.99

mm) from the first nine years of growth. No

galleries were apparent in the outermost

40.5 mm of the xylem comprising the last

18 years of growth.

A band saw was used to cut 1-2 cm

thick cross-sections containing ambrosia

beetle galleries and the galleries dissected.

Bifurcations were evident in four of the five

partial galleries dissected, with three having

a single bifurcation and one bifurcating

twice. The galleries dissected (longest arm)

ranged from 12.5 to 52.8 mm in length and

were heavily stained black, likely by the

ambrosia fungus introduced by the female

beetles (Farris 1965). Although larvae of

Monarthrum species, including M. scutel-

lare (Wood and Stark 1973), M. mali

(Fitch) and M. fasciatum (Say) (Solomon

1995) characteristically develop in

“cradles” excavated above and below the

sidewalls of the parental galleries, no brood

cradles were evident in the dissected galler-

ies or on the radial faces of the split wood.

While no evidence of brood production was

found during gallery dissections, the heavy

staining observed along the length of the

dissected galleries indicates that the ob-

served attack was not recent and the large

' Natural Resources Canada, Canadian Forest Service, 506 West Burnside Road, Victoria, British Columbia

82 J. ENTOMOL. Soc. BRIT. COLUMBIA 102, DECEMBER 2005

numbers of adults recovered suggests that

M. scutellare can attack and breed in red

alder. At least one oak-feeding species of

Monarthrum, M. laterale (Eichhoff), has

been recorded from alder and M. biden-

tatum Wood, M. hoegi (Blandford) and M.

umbrinum (Blandford) breed in A/nus spp.

(Wood 1982, Wood and Bright 1993).

Thus, while M. scutellare is usually associ-

ated with species of Fagaceae, it 1s possible

that hosts in other families may also be util-

ized. Alternatively, the emergent adults

could represent mature adults attempting to

establish brood in the firewood piece. Be-

cause galleries associated with this collec-

tion were incomplete, the ability of M.

scutellare to develop in A. rubra cannot be

determined with certainty. The presence of

brood production in choice and no-choice

breeding trials of M. scutellare in cut stem

sections of native Fagaceae (Quercus gar-

ryana Douglas ex Hooker) and A. rubra or

the discovery of brood in naturally attacked

red alder will be necessary to confirm

breeding in non-traditional hosts. Although

evidence of breeding of M. scutellare in red

alder is currently circumstantial, such novel

host associations have been demonstrated to

occur in other ambrosia beetles. Nuijholt

(1981) reported attack in red alder by two

species of ambrosia beetles, Gnathotrichus

retusus LeConte and Trypodendron linea-

tum (Olivier), which normally utilize conif-

erous species as hosts (Bright 1976, Wood

and Bright 1992). Kunholz eft al. (2000)

subsequently confirmed red alder as a

breeding host for G. retusus, while

Lindgren (1986) documented brood produc-

tion by 7. lineatum in bigleaf maple, Acer

macrophyllum Pursh.

Voucher specimens of A. aspericollis

and M. scutellare have been deposited in

the reference collection at Canadian Forest

Service, Pacific Forestry Centre, Victoria,

British Columbia. L. Safranyik, T. Shore

and A. Carroll, Natural Resources Canada,

Canadian Forest Service reviewed an earlier

version of this manuscript. Their helpful

comments and those of two anonymous

reviewers are gratefully acknowledged.

REFERENCES

Bright, D.L. 1976. The bark beetles of Canada and Alaska Coleoptera: Scolytidae. Canada Department of

Agriculture Publication 1576. Ottawa, ON.

Bright, D.L. and R.W. Stark. 1973. The bark and ambrosia beetles of California Coleoptera: Scolytidae

and Platypodidae. Bulletin of the California Insect Survey 16: 1-169.

Farris, S.H. 1965. Repositories of symbiotic fungus in ambrosia beetle Monarthrum scutellare Lec.

(Coleoptera: Scolytidae). Proceedings of the Entomological Society of British Columbia 62: 30-33.

Kuhnholz, S., J.H. Borden, and R.L. McIntosh. 2000. The ambrosia beetle, Gnathotrichus retusus

(Coleoptera: Scolytidae) breeding in red alder, A/nus rubra (Betulaceae). Journal of the Entomological

Society of British Columbia. 97: 103-104.

Lindgren, B.S. 1986. Trypodendron lineatum (Coleoptera: Scolytidae) breeding in big leaf maple, Acer

macrophyllum. Journal of the Entomological Society of British Columbia 83: 44.

Nijholt, W.W. 1981. Ambrosia beetles in alder. Canadian Forestry Service Research Notes 1: 12. Cana-

dian Forest Service, Ottawa, ON.

Solomon, J.D. Guide to insect borers of North American broadleaf trees and shrubs. U.S. Department of

Agriculture, Forest Service, Agric. Handbk. 706. Washington D.C.

Wood, S.L. 1982. The bark and ambrosia beetles of North and Central America (Coleoptera: Scolytidae),

a taxonomic monograph. Great Basin Naturalist Memoirs 6: 1-1163.

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

nomic Index. Great Basin Naturalist Memoirs 13: 1-1553.

J. ENTOMOL. SOC. BRIT. COLUMBIA 102, DECEMBER 2005

SCIENTIFIC NOTE

Notes on the status of the Eurasian moths Noctua pronuba

and Noctua comes (Lepidoptera: Noctuidae)

on Vancouver Island, British Columbia

CLAUDIA R. COPLEY?” and ROBERT A. CANNINGS’

Two Eurasian cutworm moths

(Lepidoptera: Noctuidae), Noctua pronuba

(Linnaeus) and Noctua comes (Hibner),

both accidentally introduced to North

America, are now sympatric in southwest-

ern British Columbia. The former was first

recorded in Nova Scotia, the latter in Brit-

ish Columbia. This paper reports the occur-

rence of both species for the first time on

Vancouver Island. They are the only spe-

cies of the genus Noctua known in North

America (Lafontaine 1998).

Noctua pronuba (the large yellow un-

derwing) was first reported in North Amer-

ica in Nova Scotia in 1979 (Neil 1981). It is

now known from every Canadian province

and Nunavut (Troubridge and Lafontaine

2005) and, in the USA, from Maine (Wright

1987) to Louisiana (Brou 1997) and Cali-

fornia (Powell 2002). It was first recorded

in BC in 2002 (CNC [Canadian National

Collection of Insects and Arachnids, Ot-

tawa] data) and is now abundant on eastern

Vancouver Island as far north as Sayward

(RBCM [Royal British Columbia Museum,

Victoria] data) and areas of the lower Fraser

River Valley (K. Needham, pers. comm.).

We have yet to hear of any records from the

BC Interior.

Noctua comes (the lesser yellow under-

wing) was first recorded in Canada in Bur-

naby, BC in August 1982 (Neil 1984) al-

though a specimen in the Spencer Entomo-

logical Museum, UBC (University of Brit-

ish Columbia, Vancouver) was collected in

Vancouver in July 1982. This species was

first recorded on Vancouver Island in Vic-

toria in 1990 (PFC [Pacific Forestry Centre,

Victoria] data) and is now abundant in sub-

urban habitats there. Elsewhere in BC N.

comes has been found in the Okanagan Val-

ley and Lillooet and south to Oregon

(Lafontaine 1998, J. Troubridge, pers.

comm. ).

Noctua pronuba has a wingspan of 50-

60 mm and a diagnostic orange-yellow

hindwing with a broad black border. Images

of adults and larvae are in Wright (1987),

Lafontaine (1998), and Neil and Specht

(1987). Noctua comes is similar but nor-

mally has a black mark near the centre of

the orange of the hindwing; genitalia differ-

ences distinguish the two species unequivo-

cally (Lafontaine 1998).

Although N. pronuba is known to be

migratory and a very strong flier (Passoa

and Hollingsworth 1996), its spread may

have been facilitated by human activity. It

has a wide range of host plants, many of

which are part of the horticultural trade,

food-crop industry, or are widespread

weeds. Host plant genera include: Holcus

(J. Tatum, pers. comm.), Poa and other

grasses (Wright and Neil, 1983), Azriplex,

Chrysanthemum, Dianthus, Fragaria, Free-

sia, Gladiolus, Myosotis, Polygonum, Pri-

mula, Ribes, Stellaria, Taraxacum and

Viola. Larvae also eat various common

food crops (Passoa and Hollingsworth

1996, B. Duncan, pers. comm). Noctua

comes has been recorded on Conium, Cor-

nus, Potentilla, Calendula, Cardamine,

Cirsium, Digitalis, Fragaria, Myosotis,

Plantago, Primula but, most often, on Ru-

mex crispus (J. Tatum, pers. comm.) as well

as tobacco and grapes (Sannino and

Espinosa 1999) and Crataegus (Ward

2003).

Life-history details of N. pronuba are

reported in Singh and Kevan (1965),

' Corresponding author, email: ccopley@royalbcmuseum.be.ca

A Royal British Columbia Museum, 675 Belleville Street, Victoria, BC, V8W 9W2

84 J. ENTOMOL. SOC. BRIT. COLUMBIA 102, DECEMBER 2005

Wright and Neil (1983), Morris (1987), and

Passoa and Hollingsworth (1996). In British

Columbia, N. pronuba will likely exhibit

the univoltine life history typical of Euro-

pean and eastern North American popula-

tions.

Each female lays up to 2000 eggs on

leaf undersides (Morris 1987) or non-host

substrates (B. Duncan, pers. comm.). Lar-

vae feed on foliage, crowns and roots of

hosts; immature larvae usually overwinter

(Morris 1987), but in coastal BC mature

larvae also do so (B. Duncan, pers. comm.).

Mature larvae pupate in the soil in the

spring (Passoa and Hollingsworth 1996).

Tachinid flies parasitize N. pronuba in east-

ern North America (J. Troubridge, pers.

comm.), but have not been recorded in BC

on N. comes or N. pronuba (J. Tatum, pers.

comm.). However, Trichogramma wasps

parasitize egg masses in the province (B.

Duncan, pers. comm. ).

As it is a strong, migratory flier (Passoa

and Hollingsworth 1996), can endure very

cold winters (Wright and Neil 1983), and

feeds on a wide array of plants associated

with humans, N. pronuba will probably

colonize all of BC. Adults may lay eggs on

non-plant substrates (B. Duncan, pers.

comm) or hide during the day in objects

around human habitation (although they fly

readily when disturbed), making them good

candidates for transport by vehicles.

Because of its growing abundance in

coastal BC, we believe that N. pronuba may

become an economic pest, although in the

long-term, populations will likely be mod-

erated by increasing parasitism. Noctua

comes will probably have a similar future.

We thank Jane Seed, Karen Needham

and Chris Borkent for data from PFC, UBC

and CNC collections, respectively and the

following colleagues for their permission to

use their personal communications. Jim

Troubridge (Agriculture and Agri-Food

Canada, Ottawa, ON) and Karen Needham

(UBC, Vancouver, BC) provided informa-

tion on distribution. Jeremy Tatum

(University of Victoria, Victoria, BC) and

Bob Duncan (PFC, Victoria, BC) com-

mented on food plants and life history.

REFERENCES

Brou, V. 1997. A Gulf Coast record of the European cutworm, Noctua pronuba (L.). News of the Southern

Lepidopterists’ Society 19: 3.

Lafontaine, J.D. 1998. Noctuoidea, Noctuidae (part). Jn Dominick, R.B. et al. The Moths of America North

of Mexico, Fascicle 27.3. The Wedge Entomological Research Foundation, Washington, DC.

Morris, R.F. 1987. Note on occurrence of the large yellow underwing moth, Noctua pronuba (Linnaeus)

(Lepidoptera: Noctuidae), in Newfoundland. The Canadian Entomologist 119: 403-404.

Neil, K.A. 1981. The occurrence of Noctua pronuba, (Noctuidae) in Nova Scotia: a new North America

record. Journal of the Lepidopterist’s Society 35: 248.

Neil, K.A. 1984. Noctua comes, a noctuid new to North America (Lepidoptera: Noctuidae: Noctuinae). The

Canadian Entomologist 116: 479-480.

Neil, K.A. and H.B. Specht. 1987. Sixth-instar larvae of Noctua pronuba (L.)(Lepidoptera: Noctuidae). The

Canadian Entomologist 119: 209-214.

Passoa, S. and C.S. Hollingsworth. 1996. Distribution, identification and rate of spread of Noctua pronuba

(Lepidoptera: Noctuidae) in the northeastern United States. Entomological News 107: 151-160.

Powell J.A. 2002. Noctua pronuba reaches the Pacific Coast. News of the Lepidopterists’ Society 44: 120.

Sannino, L. and B. Espinosa. 1999. On the morphology of Noctua comes (Lepidoptera: Noctuidae). I] Ta-

bacco 7: 35-43.

Singh, M.P. and D.K. Kevan. 1965. Notes on three common British species of agrotid moth. Entomologists

Record and Journal of Variation 68: 233-235.

Troubridge, J.T. and J.D. Lafontaine. 2005. The Moths of Canada. Part 1, Noctuoidea of Western Canada

and Part 2, Noctuoidea of Eastern Canada. Canadian Biodiversity Information Facility, Government of

Canada, Ottawa, ON. http://www.cbif.gc.ca/spp_pages/misc_moths/phps/mothindex_e.php.

Ward, J. 2003. Moths

of Northamptonshire.

Northamptonshire Moth Group. _http://

www.northamptonshirewildlife.co.uk/nmoths/nmoths.htm

Wright, B. 1987. The European yellow underwing, Noctua pronuba (L.)(Lepidoptera: Noctuidae) in the

Atlantic provinces (Canada) and the State of Maine (USA). The Canadian Entomologist 119: 993-997.

Wright, B. and K.A. Neil. 1983. Noctua pronuba, a European cutworm, established in Nova Scotia

(Lepidoptera: Noctuidae). The Canadian Entomologist 115: 1047-1048.

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—___ IAIN

Entomological Society of British Columbia

Volume 102 Issued December 2005 ISSN #0071-0733 |

| Directors of the Entomological Society of British Columbia, 2005-2006.................:00cc0- 2

Naomi C. DeLury, Gary J.R. Judd and Mark G.T. Gardiner. Antennal detection of sex

pheromone by female Pandemis limitata (Robinson) (Lepidoptera: Tortricidae) and its

impact on their calling behaviour ....4.....:0:0:5.5e.cesccdenececee ones 3-12

| Michael K. Bomford and Robert S. Vernon. Moisture tempers impairment of adult

Otiorhynchus sulcatus (Coleoptera: Curculionidae) climbing ability by fluoropolymer,

talc dust, and lithium grease .2...i...05.2iececencnes censuses scene Gece ee 13-20

Lawrence C. Wright, Wyatt W. Cone and David G. James. Sources of Spring and Fall

Hop Aphid, Phorodon humuli (Schrank), (Homoptera: Aphididae) Migrants in South

Central Washington «...0:.5..:0.:Heassschoengestaanoiusnesends ies uemtetetnenCee eu sae tartan 21-26

Cynthia L. Broberg and John H. Borden. Host preference by Saperda calcarata Say

(Coleoptera: Cerambycidae) ..2....3.050 ssc ect sacaek eee ee ees oe 27-34

Peter J. Landolt, Alberto Pantoja and Daryl Green. Yellowjacket Wasps (Hymenoptera:

Vespidae) Trapped in Alaska with Heptyl Butyrate, Acetic Acid and Isobutanol 35-42

Rex D. Kenner. Redescription of Haliplus dorsomaculatus (Coleoptera: Haliplidae) with

a New Synonymy and Comments on Habitat and Distribution .................. eee 43-56

Robert A. Cannings and John P. Simaika. Lestes disjunctus and L. forcipatus (Odonata:

Lestidae): An evaluation of status and distribution in British Columbia ............. 57-64

Glenn E. Haas, James R. Kucera, Amy M. Runck, Stephen O. MacDonald and Joseph A.

Cook. Mammal Fleas (Siphonaptera: Ceratophyllidae) New for Alaska and the South-

eastern Mainland Collected During Seven Years of a Field Survey of Small Mammals

lasdentsondansameeedvesesceedaroncnan ae speaentencedelnade Shes ageeeaeeln i: em ks eeee tae teee eee 65-76

SCIENTIFIC NOTES

Sujaya Rao and Stephen C. Alderman. Infestation of Bent Grass by a New Seed Pest,

Chirothrips manicatus (Thysanoptera: Thripidae), in Oregon .............:eeeeeeeeeeeeeee 77-78

Lawrence A. Lacey, Steven P. Arthurs and Heather Headrick. Comparative Activity of

the Codling Moth Granulovirus Against Grapholita molesta and Cydia pomonella

(Lepidoptera: Tortricidae) .....sc0.:.0cciasecondscccescocctes cote epg: aes <ceiieele ss aeeane arene ena 79-80

Leland M. Humble. A novel host association for Monarthrum scutellare (Coleoptera:

Curculionidae: Scolytinae) in British Columbia ....2......... ee 81-82

| Claudia R. Copley and Robert A. Cannings. Notes on the status of the Eurasian moths

Noctua pronuba and Noctua comes (Lepidoptera: Noctuidae) on Vancouver Island,

British Columbia ..3...00:..56.s:h..s0c2s.sectsaeees eee ee eee tee 83-84