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ISSN 1713-7845

JOURNAL

of the

ENTOMOLOGICAL

SOCIETY

ONTARIO

Volume

One Hundred and Thirty-Six

2005

Published October 2006

ISSN 1713-7845

JOURNAL

of the

ENTOMOLOGICAL SOCIETY

ONTARIO

Volume One Hundred and Thirty-Six

2005

Published October 2006

THE ENTOMOLOGICAL SOCIETY OF ONTARIO

President:

J. HUBER

Natural Resources Canada, Canadian Forest Service

c/o Eastern Cereal and Oilseed Research Centre

960 Carling Ave., Ottawa, ON, K1A 0C6

huberjh @agr.gc.ca

President-Elect:

B. HELSON

Natural Resources Canada, Canadian Forest Service

1219 Queen St E., Sault Ste. Marie, ON, P6A 2E5

bhelson @nrcan.ge.ca

Past President:

F. HUNTER

Dept. of Biological Sciences, Brock University

St. Catharines, ON, L2S 3A1

hunterf@brocku.ca

Secretary:

D. HUNT

Agriculture and Agri-Food Canada G.P.C.R.C.

2585 County Road 20, Harrow, ON, NOR 1G0

huntd @agr.gc.ca

Treasurer:

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Natural Resources Canada, Canadian Forest Service

1219 Queen St E., Sault Ste. Marie, ON, P6A 2E5

kbarber @nrcan.gc.ca

Librarian:

J. BRETT

Library, University of Guelph

Guelph, ON, NIG 2W1

Directors:

P. BOUCHARD

Agriculture and Agri-Food Canada

960 Carling Ave., Ottawa ON, K1A 0C6

H. FRASER (2005-2007)

Ontario Ministry of Agriculture and Food

Vineland Resource Centre, P.O. Box 8000

4890 Victoria Avenue North

Vineland, ON, LOR 2E0

M. PICKLES (2005-2007)

3252 Garner Road, Niagara Falls, ON, L2E 6S4

L. TIMMS (2006-2008)

Faculty of Forestry, University of Toronto

Toronto, ON, M5S 3B3

S. VANLAERHOVEN (2006-2008)

Dept. of Biology, University of Windsor

Windsor, ON, N9B 3P4

G. UMPHREY

Dept. of Mathematics and Statistics

University of Guelph, Guelph ON, N1G 2W1

(2004-2006)

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EDITORIAL COMMITTEE

Scientific Editors:

M. RICHARDS*

Dept. of Biological Sciences, Brock University

St. Catharines, ON, L2S 3A1

Y. H. J. PREVOST

Faculty of Forestry and the Forest Environment

Lakehead University, Thunder Bay, ON, P7B 5E1

Technical Editor: A. Rutgers-Kelly

Layout Artist: A. Rutgers-Kelly

Associate Editors:

A. BENNETT

Agriculture and Agri-Food Canada.

960 Carling Ave., Ottawa ON, K1A 06C

N. CARTER

Ontario Ministry of Agriculture and Food

Vineland, ON, LOR 2E0

R. HARMSEN

Biology Department, Queen’s University

Kingston, ON, N7L 3N6

Y. MAUFFETTE

Faculté des sciences, Département des sciences biologiques

Université 4 Québec Montréal, Montréal, QC, H3C 3P8

D. J. PREE

Agriculture and Agri-Food Canada

Southern Crop Protection and Food Research Centre

P.O. Box 6000, Vineland ON, LOR 2E0

J. SKEVINGTON

Agriculture and Agri-Food Canada

Eastern Cereal and Oilseed Research Centre

960 Carling Ave., Ottawa, ON, K1A 0C6

*Mailed MS’s to:

4338 Wildmint Square Ottawa ON, K1V 1N5

Electronic MS’s to miriam.richards@brocku.ca

JESO Volume 136, 2005

JOURNAL

of the

ENTOMOLOGICAL SOCIETY OF ONTARIO

VOLUME 136 | 2005

This volume of the Journal of the Entomological Society of Ontario marks many

transitions, and the beginnings.of some new directions. This volume of JESO was co-edited

by Yves Prévost, outgoing editor, and myself. I sincerely thank Yves for all the work he has

put in as JESO Editor over the last few years and also for helping me to learn how to guide

manuscripts through the review process and eventually to publication. I would have been

pleased to thank Yves a year ago had I| written the Editor’s message for the previous volume

— but my appreciation of his work and effort has grown all the more for having experienced

the job at first hand.

I also wish to thank JESO’s Editorial Board, some of whom are new to the job, and

some of whom are old hands: Andy Bennett, Neil Carter, Dolf Harmsen, Yves Mauffette,

and Jeff Skevington. Their scientific understanding and editorial skills ensure the scientific

quality of our Journal and have helped me enormously as I learn this new job. Our new

Technical Editor, Amy Rutgers-Kelly, has the job of transforming the manuscripts into final,

print-ready format.

This brings me to another set of transitions, and to my major goals as Editor of the

Journal. I believe that electronic publishing is critical to the survival of the Journal. Volume

136 marks our transition to completely electronic submission and manuscript processing.

This is an important shift in the production of the Journal and is the first step in the transition

to electronic publication. For the first time, authors of papers in the current volume will

receive a pdf version of their papers so that they may distribute them electronically. The

next step will be electronic publication of the Journal, in addition to paper publication.

When we have achieved this goal, then JESO will become available to a much wider

entomological audience than is currently possible with paper distribution only. However,

in order for our authors’ work to reach the audience it deserves, we also need to be listed

by electronic journal listings services. In this age of electronic database searches, articles

can be overlooked if their journals are not listed (the JESO Editor herself has failed to find

articles published before her time in JESO!). In order to be included in these listings, JESO

must commit to a regular publication schedule. To do this we intend to first close the gap

between volume year and actual publication year; this should be accomplished by 2007.

Finally, it is my pleasure to introduce the contents of V136, which cover the

entomological gamut from taxonomy to ecology to applied entomology, each study focusing

on a different order - from wasps and bees, to flies, thrips, beetles, moths, aphids, and even

mites. I hope that the scientific quality of these papers and their readability will encourage

all our readers to consider submission for publication in future issues of JESO.

Happy reading! Miriam H. Richards

Editor

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Ontario alvar ground beetles JESO Volume 136, 2005

GROUND BEETLES (COLEOPTERA: CARABIDAE) FROM

ALVAR HABITATS IN ONTARIO

P. BOUCHARD’, T. A. WHEELER’, H. GOULET

Agriculture and Agri-Food Canada,

K.W. Neatby Building, 960 Carling Avenue,

Ottawa, Ontario, Canada K1A 0C6

email: bouchardpb@agr.gc.ca

Abstract J. ent. Soc. Ont. 136: 3—23

An inventory of ground beetles (Coleoptera: Carabidae) was conducted in ten

alvar sites, representing four alvar types, in southern Ontario. We identified

142 species from 8647 specimens. Species richness and numbers of specimens

were generally higher in alvar grasslands. Alvar pavement and alvar shrubland

generally had lower species richness and specimen numbers. Each site had

between four and seven dominant (over 5% of individuals collected at the

site) species, which varied between alvar types and localities. Three of the

dominant species (Agonum nutans, Chlaenius purpuricollis, and Pterostichus

novus) have rarely been collected in non-alvar sites in the region. Most of

the species collected are associated with open habitats or grassy meadows.

The carabid fauna collected was dominated by widespread or eastern North

American species, although some northern and southern species were near

the limits of their range. The known distribution of Cicindela denikei was

extended eastward from northwestern Ontario. Nine introduced European

species were collected, and only two (Carabus nemoralis and Pterostichus

melanarius) were dominant at any site.

Introduction

Alvars are naturally open areas of thin soil overlying flat limestone or dolostone.

The vegetation is generally sparse and dominated by grasses (Poaceae), sedges (Cyperaceae),

and shrubs. Trees are rare because there are few areas with sufficient soil accumulation. Six

types of alvars are recognized based on the percentage of exposed bedrock, herb and shrub

cover, and tree cover (Catling and Brownell 1995). North American alvars are concentrated

in the Great Lakes region, where the limestone was denuded by glaciation and the sites have

been maintained as natural openings by multiple factors including fires, grazing by large

herbivores, lack of soil, and a seasonal pattern of flood-drought-flood in spring, summer,

and fall, respectively. There are 250 to 300 known alvar sites in the Great Lakes region,

' Author to whom all correspondence should be addressed.

> Department of Natural Resource Sciences, Macdonald Campus, McGill University,

Ste-Anne-de-Bellevue, Québec, Canada H9X 3V9

Bouchard et al. JESO Volume 136, 2005

mostly in southern Ontario, but also in New York, Michigan, Ohio, Quebec, and Vermont

(Catling and Brownell 1995; Reschke et al. 1999).

The flora of alvars is well known: 347 species of native plants have been identified

in Great Lakes alvars, of which 28% are considered characteristic of alvars. There are also

several endemic species. The lack of introduced European flora is notable, although many

of these species have invaded alvars recently because of artificial or man-made stresses.

Due to the combination of present conditions (geology, hydrology, etc.) and postglacial

history of alvars, plant species with northern, western, and southern Nearctic affinities

coexist in these sites (Catling and Brownell 1995).

Surveys of arthropods in North American alvars have been sporadic compared to

surveys in Europe. Approximately 1800 species of arthropods, including more than 700

species of Coleoptera, have been recorded in the Great alvar of Oland (Sweden) alone

(Lundberg 1983; Coulianos and Sylvén 1983). As for North America, Catling and Brownell

(1995) documented a number of rarely collected species of Lepidoptera, Coleoptera, and

Hymenoptera in Ontario alvars. There are also 18 species of leafhoppers (Hemiptera:

Auchenorrhyncha) occurring in Great Lakes alvars that are normally associated with prairie

habitats (Bouchard et al. 2001). Bouchard et al. (1998) provided phenology and habitat data

on three species of Carabidae (Coleoptera) that are abundant in Ontario alvars but rarely

collected elsewhere in Ontario.

As part of the International Alvar Conservation Initiative (Reschke et al. 1999),

the objective of this study was to conduct a faunal inventory of the ground beetles

(Coleoptera: Carabidae) in southern Ontario alvars to provide baseline data on the species

and communities in this unique ecosystem.

Methods

Ten sites, representing four alvar types (pavement, shrubland, savanna grassland,

grassland) were sampled in southern Ontario in 1996-1997 (Table 1). Sample sites were

described and mapped in Bouchard et al. (1998; 2001). Examples of each alvar type are

shown in Figure 1.

Sampling methods at each site consisted of one Malaise trap, 16 uncovered pitfall

traps (white plastic beer cups, 9 cm in diameter), 16 pan traps (355 ml yellow plastic bowls,

15 cm in diameter), and two flight intercept traps, distributed randomly throughout the

site. Pan traps and pitfall traps were set with their upper rim flush with the ground surface,

which in alvars with thin soil cover restricted their use to cracks in the bedrock. Propylene

glycol or ethylene glycol was used as the preserving fluid and a drop of liquid detergent or

Kodak Photofio® was added as a wetting agent. All traps were serviced twice a month and

specimens were preserved in 70% ethanol prior to mounting. Hand collecting and sweeping

were used to supplement trap catches at every visit to the sites. All traps operated from

mid-May until mid-September. At Site 9, small mammals disturbed the pitfall traps and pan

traps frequently throughout the summer. Although the data from this site were included in

the species list and calculation of overall numbers of beetles, the site was omitted from the

calculation of dominant species and rarefied estimates of species richness.

In order to determine whether certain species of carabids were characteristic of

Ontario alvar ground beetles JESO Volume 136, 2005

TABLE 1. Location of study sites in Ontario alvars.

Alvar

Site Location region Coordinates Alvar type Sampling

] Misery Bay Manitoulin N 45°47’26” alvar June - Sept 1996

Prov. Nat. Res. Island W 082°45’00” pavement

2 10 km W Manitoulin N 45°49718” alvar. June - Sept 1996

Evansville Island W 082°41°04” — shrubland

3 10 km SW Manitoulin. N 45°52’12” alvar savanna June - Sept 1996

Gore Bay Island W 082°31°48” grassland

4 10kmWGore Manitoulin N 45°53’45” alvar June - Sept 1996

Bay Island W 082°34’41” grassland

5 SkmE Napanee N_ 44°20719” alvar June - Sept 1997

Camden East Plain W 076°47°49” grassland

6 3kmN Miller Bruce N 45°07°46” alvar June - Sept 1997

Lake Peninsula ~ W 081°26’44” ~— pavement

fi Cabot Bruce N 45°14°44” alvar June - Sept 1997

Head Peninsula W 081°18’28” — grassland

8 1.5 km NE Carden N 44°41°02” alvar June - Sept 1997

Dalrymple Plain W 079°05°31” — grassland

9 7.5 km E Carden N 44°38°27” alvar June - Sept 1997

Seabright Plain W 079°03’59” shrubland

10 5 km N Smith Falls N 45°16714” alvar June - Sept 1997

Almonte Plain W 076°10°58” grassland

all alvars sampled, characteristic of particular types of alvars, or whether communities are

more affected by the fauna at the regional scale, we identified the species found in dominant

numbers at each site. Dominant species were defined as any species comprising more

than 5% of carabid specimens collected at that site (Frank and Nentwig 1995). Dominant

species were identified for each alvar site and for all sites pooled.

Buddle et al. (2005) provided strong arguments for including rarefaction curves in

biodiversity studies. We used EstimateS, version 6.0bl (Colwell 2001) to generate individual-

based rarefied estimates of observed species richness for all sites sampled (except site 9,

see comments above). Each curve is the result of 100 randomizations without replacement.

Measures of standard deviation were obtained from all randomizations for each site.

Carabidae were identified using Lindroth (1961; 1963; 1966; 1968; 1969a; 1969b).

Classification and geographic distribution follow Bousquet and Larochelle (1993). Habitat

preferences were based primarily on data in Lindroth (1961; 1963; 1966; 1968; 1969a;

1969b), although other recent information was incorporated when available (Freitag 1999;

Bouchard et al. | JESO Volume 136, 2005

FIGURE |. Examples of the four alvar types sampled during this study: A) alvar savanna,

Manitoulin Island; B) alvar grassland, LaCloche Island; C) alvar pavement, Manitoulin

Island; D) alvar shrubland, Manitoulin Island.

Larochelle and Lariviere 2003). Species known to occupy four or more of the eight habitat

categories were recorded as generalists.

In order to assess the potential interactions of carabid species in different alvar

sites, we report on the dispersal ability of each species as determined by the condition

of their hind wings. The condition of the hind wings (brachypterous or macropterous)

was recorded from Lindroth (1961; 1963; 1966; 1968; 1969a; 1969b) and Larochelle and

Lariviere (2003). All specimens are deposited in the Lyman Entomological Museum,

McGill University, Ste-Anne-de-Bellevue, Quebec, or the Canadian National Collection of

Insects, Ottawa, Ontario.

Results

Species richness and abundance

We collected 8647 ground beetles, representing 142 species (Table 2). Excluding

site 9, in which most traps were lost during the sampling period, the number of specimens

collected per site ranged from 324 (site 8) to 2188 (site 10), and the number of species

ranged from 21 (site 2) to 67 (site 5). The four sites with the highest species richness were

alvar grasslands (sites 5, 10, 8, 4; Table 2). Sites 5 and 10 had the highest numbers of

specimens (1841 and 2188, respectively) and species (67 and 57, respectively). Species

richness and numbers of specimens collected were also high in the alvar savanna grassland

JESO Volume 136. 2005

Ontario alvar ground beetles

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Ontario alvar ground beetles JESO Volume 136, 2005

(site 3), with 44 species and 1201 specimens. Species richness was lowest in the alvar

shrubland (site 2, 21 species) although more specimens were collected in that site than in

sites 6 (alvar pavement), 7 and 8 (alvar grasslands) (Table 2).

Dominant species

A total of 24 carabid species were collected in dominant numbers. Each site had

between four and seven dominant carabid species (Table 3). Only two of the dominant

species (Poecilus |. lucublandus (Say) and Pterostichus novus Straneo) were present at all

sites (Table 3). Poecilus 1. lucublandus was the most frequently collected species overall

(1342 specimens, Table 2). Agonum cupripenne (Say) was dominant in five sites and present

in all except one. Calathus gregarius (Say) was dominant in four sites. Of the remaining 20

species, five were dominant in two sites and 15 were dominant only in one. Eight different

species ranked first in dominance at the nine sites analyzed (Table 3).

Estimates of species richness

Rarefied estimates of species richness are presented in Figure 2. Overall rarefaction

curves (Fig. 2a) show that sampling was incomplete in several sites and additional carabid

species remained undetected. Site 10 (alvar grassland, Smith Falls Plain) was the only

site for which the accumulation of new species begins to level off (at about 1500-2000

specimens). Six of the sites (1, 3, 4, 5, 7, 10) have overlapping or similar richness based on

the subsamples common to all sites (Fig. 2b; N = 300 specimens). These sites include the

only alvar savanna sampled, one of the alvar pavements, and most alvar grasslands. Site 2

(alvar shrubland, Manitoulin Island) appears to be significantly less diverse than all other

sites while site 8 (alvar grassland, Carden Plain) is the most species rich of all sites (Fig.

2b). The species richness of site 6 (alvar pavement, Bruce Peninsula) is slightly lower than

at site 8 but greater than at all other sites.

Introduced species

Nine introduced European species were collected comprising 6.3% of the total

species richness: Agonum muelleri (Herbst), Amara familiaris (Duftschmid), Am. lunicollis

Schiedte, Carabus nemoralis O. F. Miiller, Clivina fossor (L.), Harpalus affinis (Shrank),

Ophonus puncticeps Stephens, Pterostichus melanarius (Illiger), and Trechus quadristriatus

(Shrank). The number of introduced carabid species collected at each site (Table 2) ranged

between one (sites 2 and 9) and five (sites 3, 5, 6). The two sites with the highest proportion

of introduced species were sites 6 and 3, representing 10.6% and 11.4% of the species

collected at those sites, respectively. Conversely, less than 5% of the carabid species

collected on sites 2, 8 and 9 were introduced. Only three species (Cr. nemoralis, Cl. fossor,

and Pt. melanarius) were represented by more than ten specimens (Table 2). There was no

consistent pattern in the distribution of introduced species between sites.

For all sites combined, 5.1% of the specimens collected belonged to introduced

species. The proportion of introduced species was highest at site 4 (12.3% of all specimens),

whereas introduced species comprised less than 2.5% of all specimens at sites 1, 7, 8, 9,

and 10 (Table 2).

Bouchard et al. JESO Volume 136, 2005

TABLE 3. Dominant species of Carabidae collected in Ontario alvars. Site numbers

correspond to those in Table 1. Dn = rank of dominant species at site (e.g. DI = most

dominant species at site); P - species present but not dominant at site * = introduced

species.

Alvar site

Species 1 Zo Sido oe 6 7 8 10 Total

Agonum cupripenne P D3 DA«DBru\ BP oP Dat ob D4

Agonum nutans Db, wP Pic! BPomih D6

Agonum rufipes P D4 P

Amara pallipes Biiz BP D4

Amara pennsylvanica P D1

Anisodactylus harrisii P D3 P eB Se

Anisodactylus nigerrimus P Pidot®stod: hie

Bembidion mimus P eo: Poe (D2ienPRoDPA see

Calathus gregarius 12): B2qrD2ni 2Pe on DG uiePal # D5

Carabus maeander P Bi aif Pain opP

Carabus nemoralis* P D4 P

Carabus serratus DSs-cBvihPO! SB D4 P P

Chlaenius p. purpuricollis P D4. oP) ok Fs IDS

Dyschirius globulosus P Pv in Be) (ePonoD&bigaBse enbiveekh

Harpalus erythropus P P D6

Harpalus faunus P BP ootBeD D2 D2

Harpalus somnulentus P Beg oii PB Dao Biker

Poecilus 1. lucublandus P P DI) Dd) 2do Ps DS esas D1

Pterostichus commutabilis PresP P r PS “D6. «F

Pterostichus coracinus D3: P P P

Pterostichus femoralis Pao D5

Pterostichus melanarius* Pin Dé (Pier P P

Pterostichus novus Db} DE DS 70 LP PO Diiobe: Riu Ll D3

Synuchus impunctatus D4 Poi? P

Number of dominant species 5 4i) BS! efe) ot bind? bot astie) oe 6

Total number of species 41 21 44 52 68 47 39 56 £57 142

Vagility

Fully developed hind wings are known in at least some specimens of 91% of the

species collected (Table 2). Thirteen species are brachypterous (Table 2).

Habitat associations and geographic affinities

A large number of carabids have previously been associated with open, bare

ground (50 species, Table 2). A similar number of species occur in grassy meadows (48

species). The third and fourth most common habitats are marshes and forests (37 and 25

Ontario alvar ground beetles JESO Volume 136, 2005

8 7 P40

50 at

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Number of individuals

FIGURE 2. Rarefied estimate of species richness for ground beetles (Carabidae) sampled

in nine Ontario alvar sites. Data points were plotted for every 25" specimen and measure of

variance (+ SD) for every 100" specimen. A) rarefaction including all specimens sampled;

B) rarefaction based on first 500 specimens (arrow indicates subsample sizes for comparison

of species richess). Sites are separated by alvar type.

Baathawtics al JESO Volume 136, 2005

species, respectively). Less than ten percent of the species occur in either riparian habitats,

wet meadows, open wet sand, or are generalists in their habitat requirements.

Most of the species collected are widespread in North America, the rest are

restricted to eastern North America. Some species are near the limits of their distribution

in the study sites. Species such as Agonum metallescens (LeConte), Amara lunicollis,

Bradycellus lecontei Csiki, Cicindela I. longilabris Say, and Lebia moesta LeConte are

boreal species near the southern limit of their ranges. In contrast, species like Brachinus

tenuicollis LeConte, Carabus sylvosus, Cyclotrachelus s. sodalis, Cymindis americanus

Dejean, Dicaelus teter, Lophoglossus scrutator (LeConte), and Selenophorus opalinus

(LeConte) are southern or southeastern species near the northern limit of their ranges. Most

of these species were collected in very low numbers. Of the above-mentioned species only

three (Ci. /. longilabris, Cr. sylvosus, and Cy. s. sodalis) were represented by more than five

specimens (Table 2).

All the species collected have previously been recorded in Ontario and few range

extensions were noted. Cicindela denikei Brown was previously known only from a small

area near the borders of Ontario, Manitoba, and Minnesota (Kaulbars and Freitag 1993;

Freitag 1999). Manitoulin Island represents a significant southeastern extension of the

known range and the species appears to be abundant in appropriate alvar sites on the island.

We did not collect Ci. denikei in alvars on the mainland.

All of the introduced species are widespread in North America except Ophonus

puncticeps, which is at the western edge of its North American range in Ontario, and Trechus

quadristriatus, which is known in North America only from Quebec, Ontario, Michigan,

and Wisconsin (Bousquet and Larochelle 1993).

Discussion

Species richness and abundance

The total number of carabid species recorded in each alvar site sampled ranged

from 21 to 67. The lowest species richness occurred in the alvar shrubland of Manitoulin

Island (site 2, Table 2). Approximately 65% of this site is covered with shrubs such as

common juniper (Juniper communis L. Cupressaceae) whereas the rest is composed almost

entirely of large blocks of limestone separated by narrow and deep cracks. This type of

habitat can be compared to similar alvars with poor vegetation diversity in Sweden (Sylvén

1983). In Europe, this alvar type, although not as rich in carabid species as sites that are

more diverse botanically, is thought to support a unique insect fauna and should not be

discarded from a conservation point of view based on low species number (Coulianos and

Sylvén 1983).

Carabid species richness was consistently higher in alvar grasslands, with most

sites supporting more than 50 species each (Table 2). These results are comparable to those

reported for European alvars (Coulianos and Sylvén 1983) where the highest number of

arthropod species was recorded in sites with rich vegetation. Alvar pavement and alvar

savanna sites were also species-rich with between 40 and 50 carabid species each.

Recent investigations of carabid diversity in different types of open habitats in

northeastern North America have reported between 26 and 76 carabid species from a single

Ontario alvar ground beetles JESO Volume 136, 2005

site (Table 4). Additionally, Canadian agroecosystems typically support between 40 and 60

carabid species at a single site (Goulet 2003). The two alvar shrublands sampled during our

study fall below the carabid species richness values recorded in other open habitats. On the

other hand, alvar grasslands occupy the higher end of the scale of carabid species richness

with more than 50 species. The alvar pavements as well as the alvar savanna support

carabid species richness similar to that of typical agroecosystems.

Generally speaking, the more species-rich sites (grasslands, savanna, and

pavements) support larger populations of ground beetles based on our trap catches. More

than 1000 specimens were collected on sites 3, 4, and 5, and more than 2000 specimens at site

10 (Table 2). However, carabid abundance was not closely correlated with species richness

in all sites. For example, the species-rich alvars at Miller Lake (site 6) and Dalrymple (site

8) supported comparatively very low numbers of specimens (less than 400 each).

Dominant species

Species such as Poecilus |. lucublandus and Pterostichus novus were dominant

in several sites and present in all alvars sampled. Whereas Po. /. lucublandus is a species

commonly encountered in grassy meadows throughout its range (Lindroth 1966; Tyler

and Ellis 1979; Levesque and Levesque 1987, 1994; Boivin and Hance 1994; Byers et al.

2000), the presence of Pt. novus in Ontario alvars in such numbers was not expected. The

latter species has been collected in high numbers in different types of forests outside of

Ontario (e.g. Snider and Snider 1986; Epstein and Kulman 1990). The numerous captures

of this species in southern Ontario alvars (> 700 specimens) indicate that this species is

closely associated with this type of habitat (Bouchard et al. 1998). Three species (Agonum

cupripenne, Dyschirius globulosus, and Harpalus somnulentus) were recorded in dominant

numbers in some alvars and were present in all sites except for the alvar shrubland on

Manitoulin Island (Table 3). As mentioned above, the alvar shrublands support the lowest

carabid beetle species richness of all alvars sampled. The harsh microclimatic conditions

in alvar shrublands seem to be an important factor in excluding certain species found

commonly in all other sites sampled.

Five dominant species were collected only in alvar grasslands: Agonum nutans, Ag.

rufipes, Amara pallipes, Am. pennsylvanica, and Harpalus erythropus. Of these, Ag. nutans

was the most numerous carabid in the alvar grassland of Manitoulin Island and was the

only species present in all other alvar grasslands. Although widespread in North America,

most of the specimens of Ag. nutans recorded in Canada prior to this study were from

the shore of Lake Erie (Lindroth 1966). Based on the uncommon catches of this species

outside of alvars in Ontario, it now appears that Ag. nutans is very closely associated with

alvar grasslands in the province (Bouchard et al. 1998). The presence of Carabus serratus

in dominant numbers only in alvar pavements is also noteworthy. This species, although

collected in small numbers in other alvar types, seems to prefer sites with moss or lichen-

covered, flat limestone with sparse grasses and shrubs growing in cracks.

Some species of dominant ground beetles were either only recorded in or only

dominant in the two alvars of eastern Ontario (Amara pennsylvanica and Harpalus faunus).

Other species such as Prerostichus coracinus and Pt. melanarius were collected in dominant

numbers only in alvars of Manitoulin Island. These observations indicate that the ground

beetle community of Ontario alvars can also be influenced by regional assemblages.

JESO Volume 136, 2005

Bouchard et al.

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Ontario alvar ground beetles JESO Volume 136, 2005

Estimates of species richness

The majority of alvar sites sampled in this study have overlapping or similar

rarefaction curves when estimates are standardized to sampling effort (Fig. 2b). The two

major exceptions to this trend are sites 2 and 8. Site 2 (the only alvar shrubland included

in the analysis) is significantly less diverse than all other sites. This result is consistent

with studies on Swedish alvars (Coulianos and Sylvén 1983) and is thought to reflect the

lower microhabitat diversity available to ground beetles in alvar shrublands. Site 8 (alvar

grassland, Carden Plain) is the most species-rich alvar sampled. The high number of

singletons (N=22) and doubletons (N=11) in this site, combined with the low number of

specimens (N=324) result in a rarefaction curve that show no signs of leveling off (Fig.

2a).

Introduced species

Of the approximately 470 species of ground beetles that occur in eastern Canada,

41 (8.7%) are introduced European species that have become established predominantly in

disturbed ecosystems. Although the number of introduced carabid species in disturbed sites

may represent a small proportion of the overall species richness (when compared to native

species), these species can often dominate trap catches (Goulet 2003).

The number of introduced ground beetle species in the sampled Ontario alvars

ranged between one, in the alvar shrubland sites, and five, in sites on Manitoulin Island, the

Napanee Plain and the Bruce Peninsula, respectively (Table 2). The number of introduced

species recorded in other recent studies on the carabid fauna of various open habitats in

eastern North America ranged between 4 and 13 (Table 4). The alvar shrublands in Ontario,

as for similar alvars in Sweden (Coulianos and Sylvén 1983), can be considered relatively

undisturbed by human activity. This hypothesis is supported the current study by the very

low number of introduced species that have invaded these harsh habitats. The alvar sites

with four or five introduced species are usually sites with rich vegetation that have been

used in the past as pastures for farm animals (e.g. alvar savanna and alvar grassland on

Manitoulin Island).

Ground beetle communities, such as those recorded by Levesque and Levesque

(1994) in raspberry plantations, can be composed of more than 80% introduced species

in some sites. The overall percentage of introduced specimens in the Ontario alvars was

low in most sites with values below 5% (Table 2). The site with the greater percentage of

introduced specimens was the alvar grassland of Manitoulin Island (12.3%), a site that has

been used in the past for grazing. Even with a value of more than 12%, the alvar grassland

of Manitoulin Island supports what can be considered a relatively undisturbed ground beetle

community when compared to those reported in other studies (Table 4).

Vagility

The majority of ground beetles collected in Ontario alvars have the ability to fly

at some stage during their life cycle. Of the thirteen brachypterous species recorded during

our study, only three occur in dominant numbers in at least one site (Carabus nemoralis,

Pterostichus coracinus, and Pt. novus). Carabus nemoralis is an introduced species that

has a large population in the alvar grassland of the Napanee Plain. Prerostichus coracinus

Bouchard et al. JESO Volume 136, 2005

occurs in large numbers on the alvar pavement and alvar shrubland on Manitoulin Island.

Because alvar pavement sites on Manitoulin Island are now preserved, and because this

species has close associations with forested areas neighboring alvar sites, its survival on

the island seems secure. Pferostichus novus, however, is closely associated with alvars in

southern Ontario (Bouchard et al. 1998) and the reduced dispersal ability of this species

could pose a threat to local populations in certain areas.

Habitat associations and geographic affinities

Given the nature of alvars, it is reasonable to predict that the carabid fauna would

be dominated by species associated with open dry habitats. Because of the occurrence

of spring flooding and the frequent persistence of temporary pools in many of the sites,

hygrophilous species would be expected to comprise another important component of the

fauna. Forest species and those associated with riparian habitats would be expected in —

lower numbers, usually as a result of movement from adjacent suitable habitats that border

or surround many of the alvar sites.

These predictions were largely confirmed by our results (Table 2). More than

thirty percent of all species are known to occur in open bare ground or grassy meadows

throughout their North American range (e.g. Amara spp.). Nineteen of the twenty-four

dominant species in Table 3 (79.2%) typically occur in dry open habitats or grassy meadows.

The presence of seasonal flooding has a major influence on the ground beetle communities

of most alvars, with 26% of all species recorded being associated with marsh habitats (e.g.

some Agonum spp.). Populations of Ag. nutans, a species rarely collected in Ontario which

seems closely associated with alvar grasslands, are thought to increase with the presence

of small bodies of water in those habitats (Bouchard et al. 1998). Bembidion mimus and

Carabus meander are typically associated with wet habitats throughout their ranges and

are found in dominant numbers in one Ontario alvar site each (Table 3). Forest ground

beetles make up a lesser component of the alvar fauna (18% of all species). Most of the

forest species were collected in small numbers except for Prerostichus coracinus which was

found in dominant numbers at two sites on Manitoulin Island. Both sites are surrounded

by forests. Ground beetles known to occur in riparian habitats, open wet sand, and wet

meadows make up only a small percentage of the Ontario alvar communities.

The Carabidae, dominated by widespread and eastern Nearctic species, do not

show the same geographic pattern as the plants. The flora of Ontario alvars consists of a

combination of southern, northern, and western species, along with some endemic species

(Catling and Brownell 1995). The presence of boreal and western plant species probably

resulted from range expansion of this flora in periglacial communities along the front of

the continental ice sheet. Following glacial retreat, relict populations remained in suitable

open habitats such as alvars. The southern flora probably colonized alvars later, during

the expansion of prairie communities in the Hypsithermal (Catling and Brownell 1995).

The presence of western carabid species such as Chlaenius p. purpuricollis in Great Lakes

alvars probably results from the existence of more continuous prairie habitat during the

Hypsithermal. This pattern is also seen in the distribution of several species of leafhoppers

(Homoptera: Cicadellidae) (Bouchard et al. 2001).

Most ground beetles are generalized predators, and their patterns of distribution

and habitat association are generally associated with climatic and physical features of the

Ontario alvar ground beetles JESO Volume 136, 2005

habitat rather than the distribution of prey species or plant communities (Campbell et al.

1979). As aresult, close correspondence between geographic or habitat affinities of carabids

and plants was not expected in this study. Nevertheless, a small number of species showed

notable patterns of distribution.

Agonum nutans, Chlaenius p. purpuricollis, and Pterostichus novus were all

dominant in this study and have rarely been collected in Ontario except in alvars. Because

of this, Bouchard et al. (1998) considered them alvar-associated species in the region.

However, all three have been collected in other habitats outside of Ontario.

Agonum nutans was present in all the alvar grasslands, but was dominant in only

one. It was not collected in other alvar types. Based on the few published records of this

species, Bouchard et al. (1998) considered Ag. nutans associated with open grassy areas in

the Great Lakes region.

Pterostichus novus was collected at all alvar sites, and was one of the most

dominant species. Although it is apparently associated with alvars in Ontario (Bouchard

et al. 1998), many specimens have been collected in a range of habitats including upland

and mesic deciduous forests and mesic old fields in Michigan and Minnesota (Snider and

Snider 1986; Epstein and Kulman 1990). Because of variation in habitat use, phenology,

and morphological characters throughout its range, Bouchard et al. (1998) suggested that

Pt. novus may represent a complex of species.

Chlaenius purpuricollis purpuricollis was collected in six of the sites and was

dominant in two. The main range of Ch. p. purpuricollis extends over the prairie ecotone

and they are found in well drained, open grasslands. In Ontario it has been recorded only

from alvars.

Cicindela denikei has a restricted range in northwestern Ontario, southeastern

Manitoba, and northeastern Minnesota and is associated with dry open substrates, usually

near forest stands (Kaulbars and Freitag 1993). The Manitoulin Island population is

apparently disjunct from the western population and given its apparent habitat preferences,

Ci. denikei may be restricted to alvars in Ontario.

The major obstacle to characterizing the carabid community of the Great Lakes

alvars is the lack of similar studies on native, open habitats other than alvars in the region. If

the dominant species identified in this study are also dominant elsewhere in the region, it may

be in habitats such as savannas, tallgrass prairie outliers, or sand beach and dune ecosystems.

Comprehensive inventories of Carabidae using standardized sampling programs should be

undertaken in more of those habitats in order to establish the distribution, abundance, and

habitat preferences of “alvar” carabids in the Great Lakes region.

Acknowledgments

We thank the Ontario Ministry of Natural Resources, conservation groups, and

private landowners for permission to collect in the alvar sites. Judith Jones and John Morton

provided information on alvar flora. Naomi de Ville and Steven Foldi assisted with field

work. Yves Bousquet (Agriculture and Agri-Food Canada) confirmed species identifications

of Carabidae and Richard Freitag (Lakehead University) confirmed the identity of Cicindela

denikei. Yves Bousquet and Andrew Bennett (Agriculture and Agri-Food Canada) provided

Bouchard et al. JESO Volume 136, 2005

useful comments on this manuscript. Frédéric Beaulieu and Maxim Larrivée assisted with

rarefaction analyses. Funding was provided by The Nature Conservancy, The Federation

of Ontario Naturalists, Fonds québécois de la recherche sur la nature et les technologies

(Quebec), and the Natural Sciences and Engineering Research Council of Canada.

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Pea leafminer trapping JESO Volume 136, 2005

INFLUENCE OF COLOUR AND TRAP HEIGHT ON CAPTURES

OF ADULT PEA LEAFMINER, LIRIOMYZA HUIDOBRENSIS

(BLANCHARD) (DIPTERA: AGROMYZIDAE), IN CELERY

A. D. MARTIN, R. S. VERNON!, R. H. HALLETT?

Department of Environmental Biology, University of Guelph,

Guelph, Ontario, Canada, NIG 2W1

email: rhallett@uoguelph.ca

Abstract dvent: Soe Ont: 436925435

Sticky trap colour preference and spatial distribution of adult pea leafminer

in celery were evaluated in 2001 and 2002 for use in developing an integrated

approach to managing this pest. Colour preference was determined by

exposing traps of various colours (red, blue, violet, green, white, and yellow)

and materials (cardboard and acetate) to leafminer populations in celery for

24-48 hours. To evaluate the vertical distribution of flying adults, yellow

sticky cards were positioned at standard heights (10, 30, 50, 70, and 90

cm) within celery crops of varying height for 24-48 hours. All cards were

returned to the lab where sex and total number of adult pea leafminer were

determined. Both sexes of adult pea leafminer were preferentially attracted

to yellow opaque or translucent sticky cards, with highest captures occurring

about 20 cm below the crop the top of the celery crop canopy.

Introduction

The pea leafminer, Liriomyza huidobrensis (Blanchard), was initially identified

in the Holland Marsh region of Ontario in 1999 after causing significant economic loss in

leafy vegetable crops (McDonald et al. 2000). This polyphagous pest is established in the

sub-tropical and temperate regions of North and South America, Europe, and Asia (Spencer

1973; Weintraub and Horowitz 1995). Since its discovery in Ontario, the pea leafminer

has remained geographically isolated within the Holland Marsh region, where it appears to

survive the winter within greenhouses (Martin et al. 2005). Local crops experiencing damage

include lettuce (Lactuca sativa Linnaeus), spinach (Spinacia oleracea Linnaeus), celery

(Apium graveolens Linnaeus), Asian crucifers (Brassica spp.), greenhouse ornamentals,

greenhouse cucumbers (Cucumis sativus (Linnaeus)), and onions (A//ium cepa Linnaeus).

Insect monitoring is an important management practice required to track pest

presence within a field effectively and time control measures accurately. Sampling methods

used for monitoring leafminers include adult counts on sticky traps, pupal collections, counts

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

Canada, VOM 1A0

? Author to whom all correspondence should be addressed.

Martin et al. | JESO Volume 136, 2005

of mines, and live larval counts within leaves (Levins et al. 1975; Poe et al. 1978; Johnson

et al. 1980). Not all of these techniques are reliable or efficient, as large errors in estimation

may occur, counts of adult and pupal stages are not representative of larval populations,

and a detrimental time delay in implementing control measures may occur by attempts to

forecast future populations (Zehnder and Trumble 1984; Heinz and Chaney 1995). In the

Holland Marsh, pea leafminer monitoring in celery, Apium graveolens Linnaeus, occurs

indirectly from systems in place for established pests or by visual damage assessments.

Although sticky card captures may not provide enough information to accurately time pest

control programs, this is the simplest and most efficient monitoring method used in the area

and it provides growers with information about levels of adult infestation in their crops.

The potential for severe economic losses make early detection and continued monitoring

of this pest particularly important. The purpose of this research was to determine the most

effective colour and placement of sticky traps within the celery canopy to maximize adult

pea leafminer captures.

Materials and Methods

All experiments were conducted in plots of celery cv. Florida 683 grown on muck

soil (60% organic matter) at the University of Guelph Muck Crops Research Station,

Kettleby, ON.

Trap Colour. Card stock (white poster board, Hilroy, Toronto, ON) was painted

with two to three coats of exterior or interior acrylic latex paint (The General Paint Store,

Cambridge, ON), and cut into 28 x 10 cm cards. Cards were folded in half (14 x 10 cm)

with the painted surface exposed. Just prior to placement in the field, traps were coated

with medium grade Sticky Stuff® (Olson Products, Medina, OH). Commercial, translucent

yellow sticky traps (Cooper Mill Ltd., Madoc, ON) were also included as a standard (14 x

10 cm). All traps were fastened to wooden stakes using bullclips and were oriented facing

north/south between the two centre rows of a four row celery bed.

In 2001, six colours (paint formulations provided in parentheses) were evaluated:

white (71-011: Al/2, T1/2), violet (81-054: E44, L2, V1Y40), blue (71-052: ACO79N),

green (71-054: A2Y, T1, Kx6), yellow (71-054: A2Y10, T1, Kx8), and red (15-101). Traps

were placed at the top of the canopy (approximately 62 cm). All treatments were replicated

six times on three consecutive days (28-30 August) in a completely randomized design.

After each 24 hour exposure period, cards were collected and the sex and number of all pea

leafminers on the total trap surface were determined using a dissecting microscope (25x).

In 2002, the heights of 30 randomly selected celery plants were measured and

the average crop height was determined prior to each experimental period. All traps were

placed at half of the average crop height for that experimental period, and were arranged

in a completely randomized design with five replications per exposure period. In order to

examine the effect of light transmission through traps on pea leafminer captures, paints were

applied to both card stock and plastic (overhead transparency film, Basics Office Products,

Kitchener, ON) cards to create opaque and translucent traps, respectively. Due to low

captures on red, blue, and violet traps in 2001, only white, green, and yellow were included.

Traps were established in the field on 6 and 20 August, 5 September, and 2 October 2002

Pea leafminer trapping JESO Volume 136, 2005

for 48 hours after which they were returned to the lab where sex and total number of adult

pea leafminer were determined.

Spectral reflectance curves of all trap colours and types were determined by

spectrophotometer (DataFlash 100 spectrophotometer, Datacolour, Lawrenceville, NJ)

and are presented in Figure 1. For translucent and commercial yellow traps, reflectance

values were determined for traps placed against both white and black backgrounds; spectral

reflectance curves were created using the mean percent reflectance values from both

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100

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80 White

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Wavelength (nm)

FIGURE 1. Spectral reflectance curves of A) opaque trap colors and B) translucent trap

colors, determined by DataFlash 100 spectrophotometer.

Martin et al. JESO Volume 136, 2005

backgrounds (Figure 1B).

Trap height. In 2001, yellow commercial sticky traps measuring 14 x 10 cm were

fastened individually to wooden stakes using bullclips at heights of 10, 30, 50, 70, or 90

cm from the soil to the bottom of the trap and positioned between the two centre rows of a

four row celery bed. Sticky cards were arranged in a completely randomized design with

nine replications per day for two days. On 3 and 6 September 2001, traps were exposed for

24 hours, after which the sex and total number of pea leafminer adults on each trap were

determined. Average crop height was approximately 65 cm throughout this experiment.

In 2002, yellow commercial individual sticky cards were fastened to wooden stakes

at 10, 30, 50, 70, and 90 cm above the soil, using a completely randomized design with eight

replications. Traps were positioned between the two centre rows of a four row celery bed

on | and 14 August, and 17 September 2002, when mean crop heights were 30, 50 and 70

cm, respectively. Traps were exposed for 48 hours after which sex and total number of pea

leafminer adults on each trap were determined.

Statistical analyses. Colour data from both years and height data from 2001 were

analyzed by analysis of variance (ANOVA) using PROC GLM (SAS Institute, 1999) after

transformation by log (x + 0.5). In 2002, numerical heights were renamed according to

their relative placement within the canopy (i.e., 60 cm below, 40 cm below, 20 cm below,

at, 20 cm above, 40 cm above, and 60 cm above the crop canopy); data for all exposure

periods were pooled and analysed as described above. Since trap placement was based on

crop height at the time of trap exposure not all relative trap positions could be tested at each

exposure period (i.e. 60 cm below the canopy was not applicable when crop height was 50

cm). Two basic assumptions of ANOVA, i.e. 1) independent treatment and model effects

and 2) random, independent, and normally distributed errors, were verified prior to analysis.

Height and colour were ranked in order of attractiveness by males, females, and total using

Tukey’s Honestly Significant Difference test. In all cases, a = 0.05 and actual, rather than

transformed, data are presented.

Results

Trap colour. In 2001, the commercial yellow sticky card captured the most female

(F = 85.26; df = 6, 109; P<0.0001 ) and male (F = 87.06; df = 6, 109; P<0.0001) pea

leafminers, followed by painted yellow and green sticky cards (Table 1). More males

were captured on white than violet sticky cards, but there were no significant differences

in captures of females on white, blue, red, and violet sticky cards. Significant date (F =

5.55; df= 2, 109; P= 0.0051) and replicate (F = 4.80; df= 5, 109; P = 0.0005) effects were

observed.

In 2002, significantly more males (F = 10.53; df = 6, 107; P<0.0001) and females

(F = 18.60; df = 6, 125; P<0.0001) were captured on yellow sticky cards than on all other

trap types (Table 2). For females, commercial and translucent yellow sticky cards were

significantly more attractive than opaque yellow cards, but for males there was no difference

between the three types of yellow cards. Translucent green and white traps did not capture

more pea leafminer than their opaque counterparts. There were significant date*treatment

interactions for males (F = 2.69; df= 18, 107; P= 0.0009) and total pea leafminer captured

Pea leafminer trapping JESO Volume 136, 2005

TABLE 1. Effect of trap color on the number of male, female, and total adult pea leafminer,

Liriomyza huidobrensis, captured on sticky traps in three 24-hour trapping sessions between

28 and 30 August 2001. Data for all trapping sessions were combined.

Mean (< SE) Number of Pea Leafminer Adults Captured!

Trap Colour Male Female Total

Commerical Yellow 138.734 14.85 a 48.73+448 a 187.47+16.19 a

Yellow 32.83+ 7.60 b 15.39 + 3.58 b 48.22+11.02 b

Green 19.56+ 2.41 b 8.33 + 0.90 b 27.39% 315 b

White Ti | Te ae bhi Aa 3.22 + 0.67 ¢ S222 [are

Blue 5.00+ 1.25 cd 2.56 + 0.37 ¢ LIZ ase

Red 2.78+ 0.45 cd 23272 +036 € 5.33+ 0.62 ec

Violet 194+ 0.38 d 1.94+0.76 ¢ 3.89+ 1.06 ed

‘Means in the same column followed by the same letter are not significantly different,

ANOVA and Tukey’s HSD comparisons of means, a= 0.05.

TABLE 2. Effect of trap color and translucence on the number of male, female, and total

adult pea leafminer, Liriomyza huidobrensis, captured on sticky traps in 2002. Data for all

dates were combined.

Mean (+ SE) Number of Pea Leafminer Adults Captured’

Trap Colour Male Female Total

Commercial Yellow 14.95+5.20 a 404021446 a 55.35419.21 a

Translucent Yellow 10.90+3.52 ab 34.00+11.68 a 44.90 14.84 a

Opaque Yellow 9.10+2.76 ab £7.00 2:03:71 och 26.20+ 8.24 b

Translucent Green 6.95+2.94 be 12.20+ 4.57 be 19.15+ 7.38 be

Opaque Green 4.05 + 1.23 bed 6.85+ 1.91 be 10.90+ 2.89 be

Translucent White 2.142,152..,.¢ 10.00+ 3.76 be 12.74+ 5.10 ed

Opaque White 1.85+0.61 cd 456-4. wLSGi © 6.40+ 1.79 d

'Means in the same column followed by the same letter are not significantly different,

ANOVA and Tukey’s HSD comparisons of means, a= 0.05.

Martin et al, | JESO Volume 136, 2005

(/ = 2.02; df= 18, 107; P= 0.0142), These interactions were apparently due to low insect

captures on 6 and 20 August, which led to a lack of significant model effects on 6 August

for males, As patterns of capture on the remaining dates (5 September and | October) were

almost identical to those for all dates combined, all data were pooled (Table 2),

Trap height, In 2001, captures of both males (3 September: /’ = 65,90; df = 4,

32; P=<0,0001, 6 September: / = 54,01; df= 4, 32; P<0,0001) and females (3 September

I= 121,62; df= 4, 32; P<0,0001, 6 September; /' = 135.49; df= 4, 32; P<0,0001) were

significantly higher on traps placed at either 30 cm or 50 cm height than at other heights,

with more males than females being captured at 30 em on 3 September (Table 3), Male

and female captures at 10 cm trap height were low to intermediate and captures decreased

with increasing trap heights above 50 cm, A significant date*treatment interaction for both

males (= 10,56; df = 4, 72; P<0,0001) and females (/° = 7,00; df = 4, 72; P<0,0001)

captured prevented the pooling of data from both experimental periods, On 3 September,

precipitation likely reduced male and female captures at 50 cm relative to the more sheltered

placement at 30 cm,

In 2002, male ("= 37.60; df= 6, 98; P<0.0001) captures were significantly higher

on traps placed between 20 cm below and 20 cm above the crop canopy, than on traps at

lower or higher positions (Table 4), Female (/° = 32,80; df= 6, 104; P<0,0001) captures

were significantly higher on traps placed below the crop canopy than on traps placed at or

above the crop canopy. Total (4° = 17,08; df= 6, 98; P<0,0001) captures were highest on

traps placed at canopy height or below, There were significant date*treatment interactions

for males (4° = 8.59: df = 6, 98; P<0,0001) and total (4 = 3,26; df = 6, 98; P = 0.0058)

captured, These interactions were apparently due to low insect captures on | August,

leading to lack of a significant treatment effect for males (/' = 0.65; df= 4; 98; P = 0.6342).

As patterns of capture on the remaining dates (14 August and 17 September) were almost

identical to those for all dates combined, pooled data are presented here.

Discussion and Conclusions

The effect of colour on specific behaviours of insects is not well known, but it

is generally accepted that attractive colours elicit more alighting by insects (Bernays and

Chapman 1994), Adult pea leafminer, both male and female, are attracted to yellow and

green sticky traps but not to white, blue, red and violet traps, Previous studies of other

leafminers, as well as other dipterans, have shown an attraction for yellow and green,

with yellow being the most common colour when sticky cards are used for monitoring

(Chandler 1981; Affeldt et al. 1983; Harris and Miller 1983; Zoebisch and Schuster 1990;

Jones and Schreiber 1994; Degen and Stidler 1996), It is uncertain why so many insects

respond strongly to yellow; however, these wavelengths are in the range of 560 to 580

nm and are not far from the peak sensitivity of an insect’s green sensitive pigment (540

nm). The reflectance intensity of peak yellow wavelengths between 560 and 580 are also

generally much higher than the peak wavelengths reflected by green pigments, and it has

been hypothesized that yellow simply represents a ‘supernormal’, or more highly attractive

version of green to certain insects (Bernays and Chapman 1994), In contrast, Delia antiqua,

the onion fly, is more attracted to white painted surfaces than yellow cardboard in the field

JESO Volume 136, 2005

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Martin et al. | JESO Volume 136, 2005

TABLE 4. Effect of trap placement relative to the celery canopy on the number of male,

female, and total adult pea leafminer, Liriomyza huidobrensis, captured on yellow sticky

card traps in 2002. Data for three dates combined (1 and 14 August, and 17 September

2002; mean crop height was 30, 50 and 70 cm, respectively).

Mean (+ SE) Number of Pea Leafminer Adults Captured’

Position

relative to crop

height n Male Female Total

60 cm above 8 0.50+0.27 b 0.12+0.12 d 0.63 + 0.26 ¢

40 cm above 16 0.81 +0.43 b 0.19+0.14 d 1.00 + 0.56 ¢

20 cm above 24 4.79+1.29 a 2.83 + 0.93 ¢ 7:62 £2.17

At crop height 24 “13.79 3.69-a 6.59+ 1.75 b 20.29+5.25 a

20 cm below 24 ~=11.76+4.00 a 16.20 + 3.86 a 27.96+5.51 a

40 cm below 16 0.87+0.31 b 13.47+2.97 a 13.33 22.95:-8

60 cm below 8 SS ag ey 14.50+3.79 a 14.50+3.79 a

'Means in the same column followed by the same letter are not significantly different,

ANOVA and Tukey’s HSD comparisons of means, a = 0.05.

(Vernon and Bartel 1985) and insects attacking the flowers of plants, such as Frankliniella

occidentalis, are frequently attracted to blue sticky traps (Gillespie and Vernon 1990).

In 2002, captures of adult pea leafminers were numerically higher on traps that

allowed for the transmission of light as opposed to opaque traps of the same colour.

Translucent traps appear brighter than their opaque counterparts, due in part to the combined

reflectance of yellow wavelengths from, and the transmission of yellow wavelengths

through, the trap.

In the 2001 trap colour experiment, approximately twice as many males as females

were captured, while more than three times as many females as males were trapped in

2002. This result is likely due to the placement of traps within the canopy (top in 2001 and

middle in 2002) rather than a reflection of the sex ratio present in the field. In 2001, the sex

ratios of the trap height (1 : 0.43 males to females) and colour (1 : 0.80) experiments were

more similar than in 2002, when the sex ratio in the colour experiment (1 : 4.15) was more

strongly biased towards females than height experiment (1 : 1.18) for three similar dates.

Even though sex ratios of adults emerging from colony-reared pupae indicate a 1:1 sex ratio

(Parrella 1987), Jones and Parrella (1986) captured 83.5% males and 16.5% females in a

greenhouse when traps were placed 0.3 m above the canopy. In potatoes, about twice as

many females as males were caught on sticky traps 10 cm above the ground, but relatively

equal sex ratios were found at heights up to 70 cm, which was 20 cm above the crop canopy

(Weintraub and Horowitz 1996).

In the trap height experiments, captures of female pea leafminers were highest

when traps were positioned within the celery canopy. Male pea leafminers were most

Pea leafminer trapping JESO Volume 136, 2005

frequently captured on traps that were located within 20 cm above or below the top of the

canopy despite the mean height of the crop changing over 40 cm throughout the duration

of the experiment. In contrast, captures of male Liriomyza spp. were highest in the middle

and lower portions of a tomato canopy in a study by Zehnder and Trumble (1984), which

may be related to canopy architecture. Increased captures of pea leafminer males in the

upper portion of the celery canopy may be explained by high flight activity as they actively

search for food and mates; while females spend more time on leaves for oviposition. This

interpretation is supported by the finding that significantly more pea leafminer larvae were

found in cucumber leaves within the lower canopy than at higher positions on the plant

(Abou-Fakhr Hammad and Nemer 2000). Our findings suggest that trap height studies

should be designed, and recommendations expressed, in relation to the height of the crop

canopy rather than height above the ground to more accurately reflect insect behaviour.

Combined captures of male and female pea leafminer adults were highest in the

middle portion of the celery canopy. These results correspond with other studies on the

spatial distribution of Liriomyza within plants when sex is not considered. In potato, more

pea leafminer were captured at or just below crop height than closer to the ground (Weintraub

and Horowitz 1996). More L. trifolii and L. sativae adults were captured by placing cards

at low to middle canopy heights in tomatoes and peppers (Zehnder and Trumble 1984;

Chandler 1985; Zoebisch and Schuster 1990), possibly indicating host-dependent spatial

distributions. There are several reasons for high captures of adult Liriomyza in the middle

of the crop canopy. Adult longevity is prolonged at cooler temperatures (Parrella 1987),

and due to an absence of direct sunlight, temperatures are cooler within the crop canopy

than above it. Maximum daily air temperatures at the time of the experiment in 2001

were 27°C with temperatures frequently rising above 30°C in 2002. Female fecundity of

L. trifolii is greatly reduced as temperatures approach 35°C, with maximum fecundity at

30°C (Leibee 1984). Female pea leafminers may remain within the crop canopy in order to

maximize their fitness. Larvae developing lower within the canopy may also be protected

from temperature extremes and parasitoids by the dense foliage.

In the Holland Marsh region of Ontario, pea leafminer populations remain low

through July and August but rapidly reach economically damaging levels from early

September to October (Martin et al. 2005). Adult pea leafminer have a high attraction to

sticky cards that reflect through the yellow portion of the spectrum, as opposed to blue.

Translucent yellow sticky cards placed 20 cm below the top of the crop canopy are most

efficient at capturing both male and female adult pea leafminer in celery. Although sticky

trap captures cannot as yet be used to adequately time chemical sprays to target larvae, they

can be used as an indicator of pea leafminer presence and movement of adults throughout

a field (Zehnder and Trumble 1984; Heinz and Chaney 1995). In addition, a rapid increase

in adults on sticky traps can be used to herald the need for more extensive larval monitoring

within the crop.

Acknowledgements

The authors would like to thank Diane Stanley-Horn, Ryan Gorman, Christine

Bahlai, and Sheila Goodfellow for technical assistance; the staff of the Muck Crops Research

Martin et al. . JESO Volume 136, 2005

Station for field plot preparation; Stuart Eaton of Cloverdale Paint and Paper, Surrey, BC,

for spectral analysis of trap colours; and Drs. Mark Sears and Mary Ruth McDonald for

their valuable comments. This research was supported by the Food Systems 2002 Pest

Management Research Program, Ontario Ministry of Agriculture and Food; the Ontario

Fruit and Vegetable Growers’ Association; the Canada-Ontario Research and Development

Fund; and the University of Guelph-OMAF Plants Program.

References

Abou-Fakhr Hammad, E. M. and N. M. Nemer. 2000. Population densities, spatial pattern

and development of the pea leafminer (Diptera: Agromyzidae) on cucumber, swiss

chard, and bean. Journal of Agricultural Science 134: 61-68.

Affeldt, H. A., R. W. Thimiyan, F. F. Smith, and R. E. Webb. 1983. Response of the

greenhouse whitefly (Homoptera: Aleyrodidae) and the vegetable leafminer

(Diptera:Agromyzidae) to photospectra. Journal of Economic Entomology 76:

1405-1409.

Bernays, E. A. and R. F. Chapman. 1994. Host Plant Selection by Phytophagous Insects.

Chapman & Hall, Inc. New York. 312pp.

Chandler, L. D. 1981. Evaluation of different shapes and colour intensities of yellow traps for

use in population monitoring of Dipterous leafminers. Southwestern Entomologist

6: 23-27.

Chandler, L. D. 1985. Flight activity of Liriomyza trifolii (Diptera: Agromyzidae) in

relationship to placement of yellow traps in bell pepper. Journal of Economic

Entomology 78: 825-828.

Degen, T. and E. Stadler. 1996. Foliar form, colour and surface characteristics influence

oviposition behaviour of the carrot fly. Entomologia Experimentalis et Applicata

83: 99-112.

Gillespie, D. R. and R. S. Vernon. 1990. Trap catch of western flower thrips (Thysanoptera:

Thripidae) as affected by colour and height of sticky traps in mature greenhouse

cucumber crops. Journal of Economic Entomology 83: 971-975.

Harris, M. O. and J. R. Miller. 1983. Colour stimuli and oviposition behaviour of the onion

fly, Delia antiqua (Meigen) (Diptera: Anthomyiidae). Annals of the Entomological

Society of America 76: 766-771.

Heinz, K. M. and W. E. Chaney. 1995. Sampling for Liriomyza huidobrensis (Diptera:

Agromyzidae) larvae and damage in celery. Environmental Entomology 24: 204-

Ot.

Johnson, M. W., E. R. Oatman, and J. A. Wyman. 1980. Effects of insecticides on populations

of the vegetable leafminer and associated parasites on summer pole tomatoes.

Journal of Economic Entomology 73: 61-66.

Jones, V. P. and M. P. Parella. 1986. The movement and dispersal of Liriomyza trifolii

(Diptera: Agromyzidae) in a chrysanthemum greenhouse. Annals of Applied

Biology 109: 33-39.

Jones, C. J. and E. T. Schreiber. 1994. Colour and height affects oviposition site preferences

of Toxorhynchites splendens and Toxorhynchites rutilus rutilus (Diptera: Culicidae)

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in the laboratory. Environmental Entomology 23: 130-135.

Leibee, G. L. 1984. Influence of temperature of development and fecundity of Liriomyza

trifolii (Burgess) (Diptera: Agromyzidae) on celery. Environmental Entomology

13: 497-501.

Levins, R. A., S. L. Poe, R. C. Lettell, and J. P. Jones. 1975. Effectiveness of a leafminer

control program for Florida tomato production. Journal of Economic Entomology

68: 772-774.

Martin, A. D., R. H. Hallett, M. K. Sears, and M. R. McDonald. 2005. Overwintering ability

of Liriomyza huidobrensis (Blanchard) (Diptera: Agromyzidae), in southern

Ontario, Canada. Environmental Entomology 34: 743-747.

McDonald, M. R., M. K. Sears, T. Clarke, J. Chaput, and S. A. Marshall. 2000. Pea leafminer,

a new pest of leafy vegetables in Ontario, Canada. Hortscience 35: 392.

Parrella M. 1987. Biology of Liriomyza. Annual Review of Entomology 32: 201-224.

Poe, S. L., P. H. Everett, D. J. Schuster, and C. A. Musgrave. 1978. Insecticidal effects on

Liriomyza sativae larvae and their parasites on tomato. Journal of the Georgia

Entomological Society 13: 322-327.

SAS Institute. 1999. SAS for Windows, ver. 8.1. SAS Institute Inc. Cary, NC.

Spencer K. A. 1973. Agromyzidae (Diptera) of Economic Importance. Series Entomologica,

The Hague 9:1-144. ©

Vernon, R. S. and D. L. Bartel. 1985. Effect of hue, saturation and intensity on colour

selection by the onion fly, Delia antiqua (Meigen) (Diptera: Anthomyiidae) in the

field. Environmental Entomology 14: 210-216.

Weintraub, P. G. and A. R. Horowitz. 1995. The newest leafminer pest in Israel, Liriomyza

huidobrensis. Phytoparasitica 23:177-184.

Weintraub, P. G. and A. R. Horowitz. 1996. Spatial and diel activity of the pea leafminer

(Diptera: Agromyzidae) in potatoes, Solanum tuberosum. Environmental

Entomology 25:722-726.

Zehnder, G. W. and J. T. Trumble. 1984. Spatial and diel activity of Liriomyza species

(Diptera: Agromyzidae) in fresh market tomatoes. Environmental Entomology

13:1411-1416.

Zoebisch, T. G. and D. J. Schuster. 1990. Influence of height of yellow sticky cards on

captures of adult leafminer (Liriomyza trifolii) (Diptera: Agromyzidae) in staked

tomatoes. Florida Entomologist 73: 505-507.

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New records of Ontario aculeate Hymenoptera JESO Volume 136, 2005

NEW RECORDS OF NATIVE AND INTRODUCED ACULEATE

HYMENOPTERA FROM ONTARIO, WITH KEYS TO EASTERN

CANADIAN SPECIES OF CERCERIS (CRABRONIDAE)

AND EASTERN NEARCTIC SPECIES OF CHELOSTOMA

(MEGACHILIDAE)

MATTHIAS BUCK', STEVEN M. PAIERO, STEPHEN A. MARSHALL

Department of Environmental Biology, University of Guelph,

Guelph, Ontario, Canada, N1G 2W1

email: mbuck@uoguelph.ca

Abstract J. ent. Soc. Ont. 136: 37—52

The Palaearctic Ancistrocerus gazella (Vespidae) and Spilomena troglodytes

(Crabronidae) are recorded for-the first time from the Nearctic region based

on material from Ontario (both species) and the northeastern United States

(A. gazella). Seven species are recorded for the first time from Canada

(C), one from eastern Canada (eC) and two from the eastern Nearctic (eN).

Sierolomorphidae: Sierolomorpha nigrescens (eN); Sphecidae: Jsodontia

elegans (eN), I. philadelphica (C); Crabronidae: Tachysphex punctifrons

(eC), Ectemnius paucimaculatus (C), Cerceris bicornuta (C); Colletidae:

Hylaeus hyalinatus (C); Megachilidae: Chelostoma campanularum (C), Ch.

rapunculi (C), Hoplitis anthocopoides (C). The recently recorded Stictia

carolina (Crabronidae) is confirmed as established in southern Ontario. A key

to the twenty eastern Canadian species of the genus Cerceris (Crabronidae)

is provided, separating for the first time males of several species in the echo,

clypeata, and nigrescens species groups. The three eastern Nearctic species

of Chelostoma (Megachilidae) are also keyed.

Introduction

Despite substantial recent faunistic work many Ontario Aculeata families still

remain relatively poorly documented. Until recently less than 59% of Ontario spheciform

wasps had been recorded (Buck 2004). Unpublished data show similar ratios for other

families of aculeate wasps, e.g., Vespidae 64% (Buck et al., in prep.), Pompilidae 60%,

Mutillidae 33% (Buck unpubl.). The present paper updates the recently published

checklist of Ontario spheciform wasps (Buck 2004) by adding six species of Sphecidae and

Crabronidae, which brings the provincial total up to 284 species. Five new records in four

other families (Sierolomorphidae, Vespidae, Colletidae, Megachilidae) are also presented.

Species inventories are important because the Ontario Hymenoptera fauna is in

' Author to whom all correspondence should be addressed.

Buck et al. ; | JESO Volume 136, 2005

a constant state of flux. Faunal change is being effected by various causes such as local

extirpations, introductions of exotic species (e.g. Smith 1991; Paiero and Buck 2004;

Romankova 2004; Buck 2004, 2005), and natural range extensions. The present paper

provides examples for the latter two categories. As documented recently for three groups

of solitary wasps (Crabronidae, Pompilidae, Vespidae: Eumeninae; see Buck 2005), the

increase in the number of exotic aculeate species is due mostly to introductions of cavity-

nesting species (including mud daubers), whereas the ground-nesting fauna has remained

largely unaffected. Six introduced species that are newly recorded in this paper, as well

as several previously recorded introduced bees (Paiero and Buck 2004; Romankova 2004;

Smith 1991) are likewise cavity-nesters or construct free-standing mortar nests. Some of

the newly recorded species have been present in Ontario (or North America) for a long

period of time without being noticed (either due to misidentification or because old material

was identified only recently). Range extensions of native species are also often overlooked

or detected with delay due to a lack of consistent sampling and a shortage of taxonomic

expertise. The suspected case of a northward range extension into Ontario of one of the

largest and most conspicuous spheciform wasps in North America (the ‘horse guard’, Stictia

carolina, Crabronidae) was reported earlier (Buck 2004). New data provided in this paper

now suggest that the species has in fact become established in Ontario.

Materials and Methods

All specimens, unless noted otherwise, are deposited in the University of Guelph

Insect Collection, Department of Environmental Biology, Guelph, Ontario.

Acronyms of depositories: AMNH — American Museum of Natural History, New

York, New York; CNCI — Canadian National Collection of Insects, Ottawa, Ontario; GAM

— private collection of Parker Gambino, Brewster, New York; GUS — private collection of

Josef Gusenleitner, Linz, Austria; ROME — Royal Ontario Museum, Toronto, Ontario.

Abbreviations: MOD — mid ocellar diameter.

Sierolomorphidae

Sierolomorpha nigrescens Evans, 1961

CANADA, Ontario: Thunder Bay Distr., 4, 2, Sleeping Giant Provincial Park,

Marie Louise Lake Campground, 48°21°47”N, 88°47°53”W, 9-14 July 2002, forest trail,

white pan traps, M. Buck.

Recorded for the first time from the eastern Nearctic. Previously, the species was

known from Saskatchewan west to the Yukon and south to California, Arizona, and Colorado

(Evans 1961). Evans (l.c.) suspected that S. nigrescens might be a western subspecies of S.

canadensis (Provancher). He mentions a range overlap between the two species, which is

incompatible with the hypothesis of subspecies status. The biology of Sierolomorphidae is

unknown. Most species appear to be associated with wooded areas and might be parasitoids

of wood-boring insects.

New records of Ontario aculeate Hymenoptera JESO Volume 136, 2005

Vespidae

Ancistrocerus gazella (Panzer, 1798)

CANADA, Ontario: York Reg., 2, Etobicoke, 16 August 1995, garden, B. Larson.

Peel Reg., 2, Cooksville, 17 June 1993, field vegetation, R. Krupke. Wellington Co., °,

Guelph, 6 October 1992, field, D. Bennett; 2, Guelph, Speed River, 7 October 1997, sweep

net, R. Vincent; Guelph, University Campus, 4, 31 August 2001, S. M. Paiero, 2, 3-5

September, 2, 6 September 2002, M. Buck, 3, 2, 16 August, 3, 25 August, 9, 26 August,

2, 30 August, 22, 1 September, 292, 3 September 2004, M. Buck. Halton Reg., 2, Milton,

Derry Rd. & 4" Line, 43°31°31”N, 79°50’25”W, 5 August 2002, S. M. Paiero. Welland

Co., 22, Thorold, 21 August 1983, M. D. Forward. UNITED STATES, Massachusetts:

2, Cape Cod, 13 August 1978, W. A. Attwater. New York: 2, New York, Central Park, 1

October 1961, P. H. Arnaud (AMNH)); 9, Kings Co., Brighton Beach area, 8 August 1962,

S. H. Hessel and R. B. Tarsy (AMNH); 1 specimen, Newburgh, Fostertown, 26 June 1967,

P. P. Babiy (GUS*). New Jersey: 2, Bergen Co., Closter, 26 June 1962, J. G. Rozen et al.

(AMNH). Delaware: | specimen, Wilmington, 11 June 1974, P. P. Babiy (GUS*). (*data

kindly provided by J. Gusenleitner; material not examined by the authors).

The oldest Nearctic specimen of this Palaearctic species examined was collected

in New York in 1961. Due to its similarity to another introduced Palaearctic species, A.

parietum (L.), which has been known from eastern North America for a long time, A.

gazella was overlooked for almost half a century. For identification Gusenleitner’s (1995)

key to central and southern European Ancistrocerus was used. Ancistrocerus gazella differs

from A. parietum by the following characters: transverse carina of tergum 1 with small

median incision only (with deep, V-shaped incision in parietum), and metanotum with

complete yellow band (in some males divided medially or absent; black or with small,

evanescent yellow spots in parietum). The Palaearctic range of A. gazella includes most of

Europe (except northern Scandinavia) east to the Caucasus, North Africa (Morocco), and

Madeira (Bliithgen 1961). Like the closely related A. parietum, it nests in a great variety

of natural and man-made cavities including hollow stems, borings in wood, hollows in

brick-and-mortar walls, or metal rails (Bliithgen I.c.). In central Europe the species has two

generations (Bliithgen l|.c.). The flight period in Ontario is similar, and probably includes

two generations as well.

Sphecidae

Isodontia elegans (F. Smith, 1856)

CANADA, Ontario: Essex Co., 2, La Salle, Brunet Park, 29 July 2005, S. M.

Paiero. UNITED STATES, New York (data kindly provided by P. Gambino; material

not examined by the authors): Bronx Co., Harris Park Annex, 29, 19 July 1995, &, 2, 16

July 1996, P. Gambino (GAM); Bronx Co., 4, East 211" Street at Woodlawn Cemetery, 12

August 1997, P. Gambino (GAM); Bronx Co., Van Cortlandt Park at Gunhill Road, 9, 6

June 2000, 4, 27 June 2001, 24, 2, 3 July 2005, P. Gambino (GAM); Westchester Co., Q,

Croton Point Park, 19 August 1999, P. Gambino (GAM).

Buck et al. 7 ) JESO Volume 136, 2005

First published records of J. elegans from the eastern Nearctic region. The natural

range of this species extends from British Columbia south to California, Texas and northern

Mexico (Bohart and Menke 1963). The easternmost previously published records are from

western Nebraska and east-central Texas. Besides the material listed above, two eastern

Nearctic records of /. elegans have been posted on an amateur entomological website on the

internet. One record (supplemented by an image of a correctly identified /. elegans) is from

West Chicago Prairie, DuPage Co., Illinois on 2 July 2005 (Marlin 2005). Another contributor

to the website mentions collecting the species “in Cincinnati [Ohio] in the 1990s” (Eaton

2005). /sodontia elegans is recognised easily by the brownish colour of the metasoma, and

there is therefore no reason to doubt the identity of the mentioned material. The occurrence

of this species in the eastern Nearctic is probably due to accidental introduction. Because

of their nesting habits (in borings in wood, stems, etc.), species of this genus are prone to

be introduced accidentally to other geographic areas. This has also happened to the closely

related eastern Nearctic species /. mexicana (Saussure), which was accidentally introduced

to southern Europe and Hawaii (Bohart and Menke 1963; Bitsch et al. 1997). Examples

of western-eastern Nearctic introductions are rare in aculeate wasps. Besides the western

Trypoxylon bidentatum Fox, which might have become established in Ontario (see Buck

2004), /. elegans appears to be the only example for this introduction pattern.

Isodontia philadelphica (Lepeletier, 1845)

CANADA, Ontario: Kent Co., 4, Rondeau Provincial Park, South Point Trail

East, 42°15°35”N, 81°50’°53”W, sandy savannah, visiting Melilotus albus Medikus flowers,

28 July 2005, M. Buck.

First record for Canada. Harrington (1902) and Walker (1913) erroneously

recorded the species from Ontario (Buck 2004). The previously known range extends from

Connecticut, New York, and Illinois south to Florida and west to California (Bohart and

Menke 1963). Jsodontia philadelphica is easily distinguished from other species in the

genus by the mainly dark pubescence of the body.

Crabronidae

Spilomena troglodytes (Vander Linden, 1829)

CANADA, Ontario: Wellington Co., 2, Guelph, University of Guelph Campus,

16 August 2004, on Solidago flowers, M. Buck. Other material examined: FINLAND: <2,

®, Tavastia australis, Janakkala, 5 July 2002, swept from Salix fragilis L., V. Vikberg.

Spilomena troglodytes is a trans-Palaearctic species (Pulawski 2005) that is

recorded here for the first time from North America. The species was identified using

Palaearctic keys by Vikberg (2000), Dollfuss (1986), and Lomholdt (1975) and compared

to authoritatively identified material from Finland that was kindly provided by V. Vikberg.

In Bohart and Smith’s (1995) key to Nearctic Spilomena species, S. troglodytes runs to

couplets 15 (females of S. pusilla (Say) and S. hainesi N. Smith) and 20 (males of S. barberi

Krombein and S. pusilla). The female of S. troglodytes is distinguished from S. pusilla

by the apically compressed tergum 6 which bears a median carina (or double carina) in

its apical third (tergum rounded and ecarinate in S. pusilla and S. barberi, undescribed

New records of Ontario aculeate Hymenoptera JESO Volume 136, 2005

for the Californian S. hainesi). From S. hainesi it can be separated by the short basal

flagellomeres (longer than broad in S. hainesi according to Bohart and Smith (1995)). The

male differs from S. pusilla by the less extensive yellow facial markings (not surrounding

antennal bases dorsally); both sexes differ from S. barberi by the scarcely pubescent apical

portions of terga 3—6 (pubescence fairly dense and in a clearly defined band in S. barberi).

Spilomena troglodytes nests in borings in wood, preferably those made by anobiid beetles,

and in thatched roofs; it provisions its brood with nymphal Thysanoptera (Lomholdt 1975).

The species was probably introduced accidentally to North America with timber or other

substrates containing nests.

Tachysphex punctifrons Fox, 1891

CANADA, Ontario: Leeds and Grenville Co., 3, Lake Opinicon, Perth Road

Village, Queens University Biological Station, 44°33’57”N, 76°19°31”W, 1-6 August

2005, L. Best.

Recorded for the first time from eastern Canada. Western Canadian records are

from Manitoba to Alberta. In the eastern United States the species occurs along the Atlantic

Seaboard from Massachusetts to Florida and west to North Dakota, Idaho, Utah, and New

Mexico (Pulawski 1988). The species is rare in the Great Lakes region (F. E. Kurczewski,

in litt.), where it has been recorded from Michigan, Illinois, Wisconsin, and Minnesota

(Pulawski 1988).

Ectemnius paucimaculatus (Packard, 1866)

CANADA, Ontario: Kent Co., 3, Rondeau Provincial Park, Marsh Trail North,

11 July 2005, visiting flowers of Daucus carota L., M. Buck.

Recorded for the first time from Canada. This species was wrongly recorded from

the Ottawa area by Harrington (1902) (see Buck 2004). It is very similar to E. stirpicola

(Packard) with which it has been confused. However, the shape of the clypeus (illustrated

by Bohart and Kimsey 1979) is a very reliable diagnostic character despite the quite

subtle difference between the two species. The colouration of tergum 5, another character

mentioned by Bohart and Kimsey (l.c., couplet 21: with a pair of yellow spots in stirpicola,

without spots in paucimaculatus) has proven unreliable. The senior author has examined

several E. stirpicola from Ontario (in CNCI) that lack yellow spots on tergum 5, and in

some melanistic specimens the yellow markings of the metasoma are reduced to a single

pair of spots on tergum 2. Ectemnius paucimaculatus has been recorded previously from

Illinois and New York south to Florida (Krombein 1979).

Stictia carolina (Fabricius, 1793)

CANADA, Ontario: Kent Co., 3, Rondeau Provincial Park, South Point Trail

East, 42°15°35”N, 81°50’53”W, dunes, visiting flowers of Melilotus albus, 28 July 2005,

M. Buck.

This large and conspicuous species was recorded recently for the first time from

Canada based on a single male from Point Pelee, Ontario (Buck 2004). At the time it was

unclear whether the recorded specimen was just a straggler or whether the species had

recently expanded its range into southern Ontario. No further collecting was done at Point

Pelee since the first discovery but the new finding of Stictia carolina approximately 65 km

Buck et al. JESO Volume 136, 2005

ENE of Point Pelee indicates that this species has apparently become established along the

western part of Lake Erie in southern Ontario.

Cerceris bicornuta Guérin, 1845 (Fig. 1)

CANADA, Ontario: Lambton Co., 2, Walpole Island, Chief’s Road, sand pits,

42°39°39''N, 82°29’°41”W, 8 August 2005, dug out from ground burrow, S. M. Paiero. Essex

Co., °, Windsor, Broadway Park, 28 July 2005, M. D. Bergeron.

First record from Canada. In the United States the species has a transcontinental

distribution from Massachusetts, southern New York, lower Michigan and Illinois south

to Florida and west to California and Oregon (Scullen 1965). Because of the unusual

colouration of the female and male morphology (see Fig. 1 and key below) this species

is easily recognisable. With twenty species, Cerceris is the largest genus of spheciform

wasps in eastern Canada but most species are difficult to identify with the current literature.

Scullen’s (1965) revision of the genus provides good illustrations of certain diagnostic

features but his key is often misleading and difficult to use for a non-expert. Furthermore,

males in several species groups (arelate-dentifrons, atramontensis-clypeata-halone-

prominens, echo-finitima) have never been separated. With recent renewed interest in the

genus (Marshall et al. 2005) we take the opportunity to provide a novel key to the eastern

Canadian species of Cerceris that remedies these problems.

FIGURE 1. Female Cerceris bicornuta from Windsor, Ontario, July 2005 (photo by S. A.

Marshall).

New records of Ontario aculeate Hymenoptera JESO Volume 136, 2005

Key to the eastern Canadian species of Cerceris Latreille

Notes. Four species from the northeastern United States might be found in

Canada in the future but are not included in the key: C. alaope Banks (Massachusetts,

southern New York), C. compar Cresson (distributed widely throughout New England

states, Pennsylvania, Ohio, Michigan, Minnesota), C. juwcunda Cresson (New York), and

C. mandibularis Patton (southern New York, southern Pennsylvania). The species marked

by asterisk (*) are not known from any other Canadian province besides Ontario. It should

be noted that colouration shows geographic variation in most species. The present key is

designed for eastern Canada and adjacent regions, and some colour characters will not

necessarily work for southern or western specimens of the species included here.

Females

(Antenna with ten flagellomeres, metasoma with six apparent segments.)

l. Clypeal process with broadly lamellate apex. Scutellum with a pair of yellow spots

~ Clypeal process not lamellate oe variably developed, in some specimens very

small and virtually absent. Scutellum usually black, rarely with complete yellow

band or a pair of yellow spots or largely ferruginouG...............:.cccsssceesseeeeseeeeetteeeeeees 3

2. Width of lamellate portion of clypeal process less than length of scape; lamella

inserted at level of lower eye margin. Metanotum black........... C. rufopicta F. Smith

— Width of lamellate portion of clypeal process ca. 1.5x length of scape; lamella

inserted far above level of lower eye margin. Metanotum with complete yellow

a as Re 2 ows pn awe cj~nayeddaccndnManeintanansaebinss C. compacta Cresson*

x 3 Tegula conspicuously humped (as in Fig. 7) with coarsely punctate summit and/or

mesopleuron with distinct, tooth-like ventrolateral tubercle near middle (Fig. 9).

Pygidial plate narrowed towards base; basal width at most slightly greater than half

maximum width. Scutellum with a pair of yellow spots or complete yellow band......

= Tegula moderately convex and usually smooth, rarely with a few scattered coarse

punctures on summit. Mesopleuron rounded ventrolaterally, in some specimens with

minute angle. Pygidial plate variable. Scutellum usually black...............::ceee rs

4. Clypeal process weakly trilobate, middle lobe broad, with slightly convex apex.

Tegula evenly convex, not humped, with smooth summit. Metanotum black. Yellow

fasciae on terga 2 and 4 complete, broadly interrupted ON 3...........:ececeeeeeneeeeeeeteeees

+4 Clypeal process bilobate or bidentate, its apical margin slightly to conspicuously

emarginate between corners. Tegula distinctly humped, with coarsely punctate

summit. Metanotum with complete yellow band. Yellow fasciae on terga 24

OMENS roo = Ie. ohh add eR cake ctenctiee cab beteensensensbenhcnbensaenctecsensensessceestatedereneenses 5

De Clypeal process narrow (ca. 1.5x MOD), with sharp median incision. Tergum |

largely ferruginous. Ventrolateral tubercle of mesopleuron poorly developed, apical

angle in anterior view much greater than 90°. [Subantennal sclerite and clypeus

os io akasanxgupcno NORM MA CLIRK ION C. crucis Viereck & Cockerell*

Buck et al. JESO Volume 136, 2005

8 |} 9 10

FIGURES 2-10. Diagnostic features of Cerceris adults. Male head, lateral view: 2 — C.

halone, 3 — C. clypeata, 4— C. occipitomaculata. Male flagellomeres VIII—X]I, dorsal view:

5 — C. atramontensis (arrow pointing to posterior swelling of flagellomere), 6 — C. bicornuta.

Male tegula, posterior view: 7 — C. echo, 8 — C. finitima. Ventrolateral tubercle of female

mesopleuron, anteroventral view: 9 — C. echo. Male hind basitarsus, posterodorsal view: 10

— C. bicornuta.

~ Clypeal process broad (> 2 MOD), very shallowly emarginate. Tergum 1 black,

usually marked with yellow. Tubercle of mesopleuron prominent, apical angle in

anterior view < 90° (Fig: 9). ......icc.cciéshosssanedasoudes einnanoosteepecedaewes semen enttnenaet atm 6

6. Subantennal sclerite and clypeus almost completely yellow. Clypeus essentially

flat above process; apex of process extending ventrally to level of clypeal

tit +1] eee rsecann cM roe ey Mec) C. finitima Cresson*

— Subantennal sclerite and clypeus black, the latter rarely with small median yellow

spot. Clypeus with median convexity above process; apex of process ending short of

level.of ventral clypeal margin:. \:..c.2tad.e lamibineey...2uonigt C. echo Mickel*

re Clypeal process virtually absent:.. 2c ..2jecs Gennaio alee 8

_ Clypeal process prominent, of variable shape....cscscis<is0.c0-sss22<-scencentonsoaesebsced ee emaneas 9

New records of Ontario aculeate Hymenoptera JESO Volume 136, 2005

10.

11.

13.

14.

#5.

Disc of clypeus evenly convex, with a pair of tiny tubercles just above apical margin.

Antennal flagellomeres (VI—)VII-X with linear tyli. Propodeum black. Metasomal

terga black except for broad yellow fascia on tergum 2 and in some specimens small

lateral spots on Siar 3. Wing strongly infuscated. Large species, body length ca.

eae DRY LOE EE SOBA eT I ied OREO RE LENE ae C. fumipennis Say*

Clypeus with an indistinct, curved, ridge-like swelling near middle, area below

swelling flattened. Antennal flagellomeres without tyli. Propodeum with a pair of

yellow spots. Metasoma with subequal yellow fasciae on terga 2-5. Wing weakly

mnuseatods "Smaller; length ca. 10 mim. .i)...55560460 ii iicdccicsedssdiecssesedescesees C. deserta Say

Clypeal process developed as low, conical, median tubercle..............0.ccccccceeseeeeees 10

Clypeal process not conical, its apex emarginate or truncate (in some specimens only

ener aaanCa Sul Pr FEN Rs AU SU RO AEN aed 1]

Clypeal process somewhat flattened dorsoventrally and slightly deflected downward

at apex (lateral view). Inner margin of mandible with low and ill-defined teeth, not

notched. Scutellum black, metanotum yellow. Metasomal terga 2—5S with subequal

SNROMRC TA bhatt ACL Ja ORO C. nitidoides Ferguson

Clypeal process neither dorsoventrally flattened nor deflected, apex rectangular in

lateral view. Second mandibular tooth very enlarged, inner margin of mandible

deeply notched just distal of tooth. Scutellum yellow-banded, metanotum black.

Tergum 2 black, tergum 3 with broad yellow fascia, terga 4 and 5 with narrow yellow

_ |: Be J 0 5.8 ee ar Ee CR Oe PL Ee ee C. insolita Cresson*

Yellow fascia of tergum 2 distinctly wider than on following terga................... 12

Wellow fasciae of metasomal terga 2—5 subequal..................--sccsocescssconsssotenesorenees 16

Head, pronotum, and propodeum with ferruginous markings; scutellum and metasoma

largely ferruginous. Terga 3—5 without yellow fasciae. Pygidial plate about half as

anne ae Dene datas BE NAMIE dese. C. bicornuta Guérin*

Body without ferruginous markings. Terga 3—5 with yellow apical fasciae. Pygidial

Demmacnan Whe as at mmiddle..5.{005 200.020 BO INS I ween 13

Clypeal process (measured from base of clypeus to apex of process along midline)

at least as long as scape. Clypeus with yellow spots laterally (exceptionally

Rasta periiaresereiienirireriataivereel ibd AA dtodd C. clypeata Dahlbom

Clypeal process shorter than scape. Clypeus often without yellow spots............... 14

Clypeal margin with a pair of very prominent and stout paramedian teeth

bordering deep median emargination (depth of depression equals diameter of

scape). Edge of clypeal process strongly curved (often almost semicircularly) in

anteroventral view. Clypeus with yellow markings laterally and medially below

SOMRM NMR SSSETS oe PRU a dcoiiees aides taste soued eoecnnaaeuvenaveevecewesdaseeee C. halone Banks

Paramedian teeth of clypeal margin less robust and less prominent, area between

them moderately emarginate (depth of emargination at most half diameter of scape).

Edge of clypeal process usually straight to slightly curved in anteroventral view.

Clypeus black laterally, rarely with median yellow spot below process..............-.+. 15

Apical corners of clypeal process as far apart as centres of antennal sockets. Process

projecting clearly less than diameter of scape beyond level of flattened lower part of

clypeus (lateral view). Widespread..............csccseeeeseeeseeeees C. atramontensis Banks

Buck et al. | JESO Volume 136, 2005

- Apical corners of clypeal process as far apart as lateral margins of antennal sockets.

Process projecting by at least diameter of scape beyond level of flattened lower part

of clypeus (Ottawa area, one record only)............c::cccceseceeeeeeees C. prominens Banks *

16. | Clypeal margin without median tooth. Rarely collected speci€s............:sseseseeee: 17

- Clypeal margin with low, often rectangular, median tooth...............cc:ccecsseeeesseeeeeees 18

17. | Clypeal margin with one pair of teeth that are twice as far apart as antennal sockets;

margin between teeth straight. Clypeal process parallel-sided and broad, its apical

comers further apart than lateral margins of antennal sockets. Scutum dull between

PUI... cni.cxcnnvsvaxnnshacsanceey haat vestanet an tedeeh hye aa C. occipitomaculata Packard*

~ Clypeal margin with two pairs of teeth, inner pair larger and about as far apart as

antennal sockets; margin between inner teeth emarginate. Sides of clypeal process

distinctly convergent towards apex; apical corners closer to each other than centres

of antennal sockets. Scutum shiny between punctures................... C. astarte Banks*

18. | Clypeal process with deep triangular emargination, its apical edge almost straight in

anteroventral view, rounded over medially. [Median tooth of clypeal margin broad,

rectangular. Markings of body bright yellow. ]...................00 C. dentifrons Cresson

- Clypeal process less deeply and more evenly emarginate; its apical edge acute

medially and usually strongly curved in anteroventral VIeW.............:::c:cccsseeeeeeeeeeees 19

19. | Median tooth of clypeal margin broad, rectangular. Body markings pale yellow to

IVORY $s; is «065 :s:.2- ote dseaieh een mation Saeed bend, cee C. nigrescens F. Smith

— Median tooth of clypeal margin narrow, triangular. Body markings bright yellow....

asrenteddkssenenes died Gch obbieheipes hheed- dees dadad aeglyaeg a culate eae deena ey eaeeal C. arelate Banks

Males

(Antenna with eleven flagellomeres, metasoma with seven apparent segments.)

te Sternum 2 with median subbasal’ swelling.....28 0 jiscscity.i- id. Lea ee Ie 2

& Stemumm 2 flat; .:...c00sés ct Setecele Ge. Beh eda eat) eae Boe 6

2 Clypeus extensively. black, especially laterally: -.: swusi:5..25-taiiish--ab Soko. Nea 3

— Clypeus-yellow except apical margin. :....02.20.5h050ciLnnnthcsstbccedamnaeoens ceeded er -

x Tegula with coarsely punctate summit. Metanotum yellow (yellow spot evanescent

in some specimens). Tergum | usually marked with ferruginous; tergum 3 with

complete yellow apical faseia.s)scic1.3.- ade taseterpeaenss: C. crucis Viereck & Cockerell*

_ Tegula with impunctate summit. Metanotum black. Tergum | black, without

ferruginous markings; tergum 3 with pair of broadly separated yellow lateral spots...

oT EO ee ee ee eRe Te ee een on C. kennicottii Cresson*

4. Tegula moderately and evenly convex, with indistinct punctures. Scutellum black.

Propodeal enclosure smooth with weakly impressed median groove............:::::00e0e

Sain hiiusedaauanbnggn ss¥ee devwoceaunys <anjdubepaduns dat sdudausapeien dbl apenas eerste C. nitidoides Ferguson

— Tegula distinctly humped (Fig.7, 8) and coarsely punctured. Scutellum with pair of

yellow lateral spots (spots evanescent in some specimens). Propodeal enclosure with

distinct transverse TidGeS.3.5.:.50) sensswdenante-tintt sad heels Seller S

5; Metanotum with coarse, contiguous punctures, with no interspaces except along

posterior margin. Tegula moderately convex (Fig. 7), convexity subequal to greatest

New records of Ontario aculeate Hymenoptera JESO Volume 136, 2005

ll.

iP:

13.

diameter of flagellomere III. Apical fascia of tergum 2 with slightly convex or straight

anterior margin. Erect setae of sterna 3—6 shorter (length < 1 MOD)........0000000000....

err te doe: Clit ites ecec ccs vO tee i ceddel Liawia! C. echo Mickel*

Metanotum with extensive shiny interspaces between small punctures. Tegula

extremely convex (Fig. 8), convexity subequal to 1.5x greatest diameter of

flagellomere III. Apical fascia of tergum 2 emarginate anteriorly. Erect setae of

sterna 3—6 long (length > 1 MOD))..............:cccceccescesseseeeeseeeeees C. finitima Cresson*

Setal brushes of clypeal margin very broad, separated by distinctly less than their

own width. Scutellum. with yellow band. Tergum 2 black, lacking apical fascia;

terga | and 3 with broad apical fasciae; fasciae narrow on terga 4 and 5..........0..00..000-

SMa rer roo on Glawin. Geen rat AL. dee C. insolita Cresson*

Setal brushes of clypeal margin separated by at least their own width, not extending

onto median lobe. Scutellum black or with pair of yellow spots, exceptionally with

yellow band. Tergum 2 with well developed apical fascia, other terga variable.......7

Apical fascia of tergum 2 broader than those of following terga........0....00.cccceceeeeee 8

een tit BAAD 4 SUDCGUAN. «20. 5. ccaccscocesinscsntesaocezeccsessesdsesed IR cacenee ee

Flagellomere XI without outstanding setulae on posterior surface..............0000...000- 9

Flagellomere XI with a few outstanding setulae on posterior surface (e.g., Fig. 5).....

ep NObR Te LN boven cede PD Obed). 21 lon. ae eee Vino, wey. ..0..12

Scape and clypeus black, the latter in some cases with small ivory spot(s). Pale

OA CEES nee ee ne eee C. fumipennis Say*

Anterior surface of scape yellow. Clypeus yellow except apical margin. Pale

CE NTA 7 Panne SERPS Gc re TEER EY ee eo ee ee eee ee 10

Tergum 7 with pair of basolateral setal tufts. Sterna 3-5 with conspicuous, dense

erect hair. Hind basitarsus somewhat swollen apically and slightly curved outward

(Fig. 10). Flagellomere XI conspicuously curved (Fig. 6)....... C. bicornuta Guérin*

Tergum 7 with scattered setae laterally. Sterna 3—5 with moderately dense, inclined

hair. Hind basitarsus simple. Flagellomere XI nearly straight......000.0000 ee 1]

Flagellomeres (VIII-)[X—XI with bare posterior patches (devoid of microtrichia).

Yellow area of median clypeal lobe more rounded ventrally. Metanotum black........

MR Sh het cons cats. cetera dnsic ysis deg Seu Hae da. cation bene be eernsnes C. rufopicta F. Smith

Apical flagellomeres without bare posterior patches, evenly covered with microtrichia.

Yellow area of median clypeal lobe more or less triangular and pointed ventrally.

Paeamotum wih yellow band....0..)..202.4.... 00002. C. compacta Cresson*

Clypeus conspicuously flattened (Fig. 2). [Flagellomere IX with low posterior

swelling, visible as slight convexity in profile; as in Fig. 5.]............ C. halone Banks

Shypetawitinthe usual:slight convexity: (Fig. 3)...2.5..0. 0c ieee lalasees 13

Flagellomere [X without posterior swelling (straight in profile but with the usual bare

patch). Lower surface of flagellum orange..................:::eceeeeees C. clypeata Dahlbom

Flagellomere IX with low posterior swelling visible in profile (Fig. 5). At least median

pacsoniel farpeilum black ventrally. .:!.2..0.............006. 00 i AE a

SPADA Tire SOLO 3.0. Lectin cc C. atramontensis Banks and C. prominens Banks*

Note: Males of C. atramontensis and C. prominens cannot be separated based on

morphological characters. While the former is one of the most common species of

the genus in Ontario (distribution: southern Ontario north to Killarney Provincial

Buck eral. | JESO Volume 136, 2005

Park) the latter is known only from a few specimens collected around 1900 in the

Ottawa area (Buck 2004).

14. Median clypeal lobe with lateral teeth only, median tooth absent. Posterior fringe

of erect setulae present on whole length of flagellomeres XI and X, fairly dense at

base of flagellomere XI. Propodeal enclosure smooth except for weakly impressed

mediag STOVE. i.clath eae 5. Re ee dae oe C. astarte Banks*

~ Median clypeal lobe with median tooth (indistinct in atypical specimens). Posterior

fringe of erect setulae interrupted or sparse near base of flagellomere XI, setulae on

flagellomere X restricted to apical half or less. Propodeal enclosure with more or less

distinct longitudinal ridges:sci.2001.685 nie, Maat ee Bei 15

15. | Clypeus flattened, with welt-like transverse swelling above margin of median lobe.

Width of median clypeal lobe > 1/3 clypeal width, with distinct emarginations

between tecthixi)...chest 26. Mets besoin ...elal. aie C. deserta Say

— Clypeus convex, without transverse swelling above margin of median clypeal lobe.

Width of median clypeal lobe < 1/3 clypeal width; emarginations between teeth

IAGISTITICE, 2. ances Sed decavenkd be Sobsbadin tod dubedee nd en eae NRC LAIR dies SOULE, A ae 16

16. Clypeus more strongly convex (Fig. 4). [Body markings yellow] (Ontario, one

record :only)i.sis2. ctliniae 2 I a. C. occipitomaculata Packard*

- Clypeus weakly convex (as in Fig. 3). Mostly commonly collected species.......... 17

17. Body markings ivory to pale yellow. [Apical flagellomeres as in C. arelate]| (see

RROD OU aN dea ons cs ddesthn ta beh oscil, satiate C. nigrescens F. Smith

~ Body markings bright yellow or slightly palef..............c ese eeeceeesesseeesseeeeesteeeeseeeeeeees 18

18. | Flagellomeres (V—)VI—XI each with bare patch (devoid of microtrichia) posteriorly

(patches becoming smaller on more basal flagellomeres)......... C. dentifrons Cresson

- Only flagellomeres (IX—)X—XI with bare patch ventrally (small on flagellomere IX if

PPOBEME CAA... dacveinsdtusiibih ds Michel kip OMIA Re aoe C. arelate Banks

Note: This character requires careful examination under critical lighting.

Colletidae

Hylaeus (Spatulariella) hyalinatus F. Smith, 1843

CANADA, Ontario: Halton Reg., Oakville, 16 Mile Creek nr. Hwy 407, 4, 2,21

August 2004, 4, 25 June 2005, M. Buck. Essex Co., 4, W of Harrow, 28 June 1993, edge

of farmer’s field, pheromone trap, J. Doherty.

Newly recorded from Canada. This Palaearctic species was first recorded from

the Nearctic region by Ascher (2001) based on material collected in New York in 1997 and

later. All New York state records are from the Ithaca area (Tompkins Co.) and the New York

City area (Bronx Co., New York Co., Westchester Co.) (Ascher et al. 2006). The earliest

specimen from Ontario was collected in 1993 (see above), and now represents the oldest

known record from North America. Hylaeus hyalinatus is distinguished easily from other

northeastern Hylaeus by the well-developed omaulus and the protruding spatulate process

of male sternum 8. The male terminalia and facial markings were illustrated by Ascher

(2001). This species nests in a great variety of cavities in and above the ground, including

abandoned solitary wasp or bee nests, hollow twigs, and borings in wood, etc. (Ascher

ke)}.

New records of Ontario aculeate Hymenoptera JESO Volume 136, 2005

Megachilidae

Chelostoma (Chelostoma) campanularum (Kirby, 1802)

CANADA, Ontario: York Reg., <, Etobicoke, 29 June 1997, backyard, C. S.

Onodera; 23, 102, Toronto, Humber River nr. old mill, 11 July 1999, T. Romankova

(ROME). Wellington Co., 32, Guelph, 22 July 2004, on Campanula, S. M. Paiero; °,

Guelph, Wellington St. & Fife Rd., 4 September 2004, at roots of uprooted tree, M. Buck.

Halton Reg., 3, Oakville, 21 July 1976, W. A. Attwater. Welland Co., 2, Welland, 27 June

1977, R. G. Bennett.

Newly recorded from Canada. This is another Palaearctic species that has

apparently been introduced accidentally to North America. Previously, the species

was known only from New York, where it was first collected in 1973 (Eickwort 1980).

Chelostoma campanularum nests in borings in wood or hollow twigs, and was probably

introduced with shipments of wood (e.g., wooden pallets) containing nests. The species is

oligolectic on Campanula (Eickwort l.c.).

Chelostoma (Gyrodromella) rapunculi (Lepeletier, 1841)

CANADA, Ontario: Halton Reg., 3, 2, Oakville, 16 Mile Creek nr. Hwy 407,

25 June 2005, visiting flowers of Echium vulgare L., M. Buck.

As the previous species, Ch. rapunculi is native to the Palaearctic region, and is

recorded for the first time from Canada. It was first discovered in the Nearctic region by

Eickwort (1980) based on specimens collected in New York as early as 1962. The biology

is similar to Ch. campanularum, with females being oligolectic on Campanula, though

our specimens were visiting flowers of viper’s bugloss (Echium vulgare). The differences

between the two introduced and the single native eastern Nearctic species of Chelostoma

are summarized in the key below.

Key to the eastern Nearctic species of Chelostoma Latreille

| Female (ten flagellomeres; metasomal sterna with scopa)............:cc:ccessssesseeeeseseeseeees 2

— Maencioven fagellomeres; scopa absent) iv..0..6/iicil ail cide isn attesecteets +

2. Terga 1-4 with apical fasciae of white appressed pubescence. Body length 8-11

set 52 ES ER ERS in os eo ae ee Ch. rapunculi (Lepeletier)

Terga 14 without fasciae of appressed pubescence. Body length 5-8 mm.............. 3

3. Length of mandible approximately 2/3 eye height (Eickwort 1980: Fig. 3).

Flagellomeres VIII and IX at least as long as wide. Setae of mid basitarsus

aera rpm certs each ction au taelte Ch. philadelphi Robertson

+ Length of mandible approximately half eye height (Eickwort 1980: Fig. 2).

Flagellomeres VIII and IX wider than long. Setae of mid basitarsus conspicuously

pemnmmneenrre ty Bre cecil 2 le Ch. campanularum (Kirby)

4. Apical tergum trilobate, median lobe below paired lateral lobes; lobes truncate

apically (Eickwort 1980: Fig. 6). Clypeus truncate apically. Sternum 2 with

prominent, nearly semicircular protuberance (posterior view). Body length 8-11

NN a Sev pak aden de hice senicheadetsread exdtebeneaanebaes Ch. rapunculi (Lepeletier)

Buck et al. | JESO Volume 136, 2005

- Apical tergum with paired lobes only, lacking median lobe. Sternum 2 with low,

transverse, welt-like swelling. Body length 5—8 mm.................ccccccceeseseecceeseeeeceeenes 5

3: Apical tergum quadridentate, lateral pair of teeth about half the size of paramedian

pair (Eickwort 1980: Fig. 4). Flagellomere II longer than wide and longer than

fageliomere It. 2501.0... La I Ch. philadelphi Robertson

- Apical tergum bidentate, only paramedian pair of teeth present, elongate (Eickwort

1980: Fig. 5). Flagellomere II wider than long, at most as long as flagellomere I.........

sn ndviddd ba c0d sat sive ogelboct MUSIC bs LE RIOTRDER CAAe, Lee Ch. campanularum (Kirby)

Hoplitis (Hoplitis) anthocopoides (Schenk, 1853)

CANADA, Ontario: Peel Reg., 2, Forks of the Credit, gravel pit NW of Provincial

Park, 43°49°24”N, 80°0°57°W, 5 August 2002, white pan traps, M. Buck. Wellington

Co., 4, Rockwood, Valley Rd., 43°46’56”N, 80°8’28”W, 21 July 2004, on rock and mortar

walls of ruin of house, M. Buck; 64, 42, Guelph, Niska Rd., Guelph Bird Sanctuary, 11

June 2005, abandoned gravel pit, M. Buck; 3¢, 292, Guelph, Wellington & Fife Rds., 12

June 2005, abandoned lot, M. Buck; 34, &, same locality, reared from mortar nests on

rocks collected on 1 June 2005 (emergence dates in lab: 4, 14 June, 3, 15 June, 2, 2, 30

June 2005), M. Buck. Halton Reg., Milton, Woodland Trails camp, 6 Line Nassagaweya,

43°32’51”N, 79°59°35”W, &, 27 June 2005, 3, 29, 8 July 2005, S. M. Paiero.

This species is also native to Europe and is newly recorded from Canada. It was

known previously from New York, where it was first collected in 1969 (Eickwort 1970).

According to S. Droege (in litt.) the species now also occurs in West Virginia (Hampshire

Co., 2004). The biology of H. anthocopoides was studied thoroughly by Eickwort (1973).

Unlike native Hoplitis (which belong to different subgenera) this species is a true mason bee,

i.e. it builds “mortar and pebble” nests. The nests are constructed on exposed areas of rocks

(large or small), rubble, stone walls, etc. The females are oligolectic on. viper’s bugloss, an

introduced European weed that is widespread in disturbed areas with poor soil. This species

can be separated from other species of Hoplitis using Mitchell (1962) in conjunction with

the supplementary couplets provided by Eickwort (1970).

Acknowledgements

We thank J. M. Carpenter (AMNH), C. Darling (ROME), and J. T. Huber (CNCI)

for the opportunity to study material under their care, as well as P. Gambino (Brewster,

NY) and J. Gusenleitner (Linz, Austria) for providing specimen data from their private

collections. V. Vikberg (Turenki, Finland) is thanked for his help in identifying Spilomena

troglodytes and for donating comparative material of this species to the Guelph collection.

L. Best (University of Guelph) is thanked for making his personal collection available to

us and for depositing Tachysphex punctifrons and other interesting Hymenoptera material

in the University of Guelph collection. S. Droege (USGS Patuxent Wildlife Research

Center, Beltsville, MD) kindly informed us of an additional state record for Hoplitis

anthocopoides.

New records of Ontario aculeate Hymenoptera JESO Volume 136, 2005

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Insecticide resistance in oriental fruit moth JESO Volume 136, 2005

STATUS OF RESISTANCE TO INSECTICIDES IN POPULATIONS OF

THE ORIENTAL FRUIT MOTH GRAPHOLITA MOLESTA (BUSCK)

(LEPIDOPTERA: TORTRICIDAE) IN SOUTHERN ONTARIO

D. J. PREE, K. J. WHITTY, M. K. POGODA!, L. A. BITTNER

Agriculture and Agri-Food Canada, S.C.P.F.R.C.,

P.O. Box 6000, 4902 Victoria Avenue North,

Vineland Station, Ontario, Canada, LOR 2E0

email: pogodam@agr.gc.ca

Abstract J. ent. Soc. Ont. 136: 53-70

Populations of Oriental fruit moth Grapholita molesta (Busck) (Lepidoptera:

Tortricidae) were assessed for levels of resistance to organophosphorus

(OP) and pyrethroid insecticides approximately 10 years after initial assays

identified the resistance, and 6-8 years after a resistance management

strategy was introduced for use in peach production systems. Resistance to

OP insecticides was detected at all three locations tested (Niagara, Norfolk,

and Essex). Resistance frequencies had increased at one site (Jordan Station

Experimental Farm) that had been monitored closely in 1999; however,

frequencies at that site did not increase over the three years reported here.

Results also indicated that pyrethroid resistance had declined in the Niagara

area, occurred at low levels in the Norfolk area, and was not found in the

Essex area. Mechanisms and cross resistances between OP and carbamate

insecticides appeared similar to those described in earlier studies. Resistance

was associated with elevated general esterase activity and the presence of an

acetylcholinesterase which was less sensitive to inhibition than in susceptible

populations. Resistance to azinphosmethyl and phosmet was expressed at low

levels but high levels of resistance was expressed to the methyl carbamates,

carbaryl, or carbofuran. Chlorpyrifos was equally toxic to both susceptible

and resistant populations. Resistant populations were more susceptible to

acephate. All of these characteristics were similar to the resistance described

in previous reports. Chlorpyrifos, which is scheduled to be deregistered

in 2006, may be replaced by the ecdysone agonist methoxyfenozide or the

neonicotinoid acetamiprid. The data indicated low levels of resistance (1.7

fold at the LC,,) for methoxyfenozide associated with OP resistance, but

control of the first generation was achieved in both small plot and program

trials. Later applications were less effective. Acetamiprid was generally

effective throughout the season and was equally toxic to both OP resistant

and susceptible populations. In field trials over two seasons, neither of these

products was associated with outbreaks of phytophagous mites. However,

' Author to whom all correspondence should be addressed.

Pree et al. JESO Volume 136, 2005

the potential fit of these products into IPM programs for peach will need

further assessment.

Introduction

An integrated pest management (IPM) program for peach, introduced to growers in

the mid 1970’s, was the first widely used IPM program in Ontario. The program relied on the

use of pheromone trap catch data to time applications of the organophosphorus insecticides

azinphosmethyl! and phosmet to control the Oriental fruit moth (OMF), Grapholita molesta

(Busck) (Lepidoptera: Tortricidae) (Phillips 1973). This program remained effective for

approximately 20 years but resistance to these insecticides resulted in up to 45% fruit

infestations in 1993 and 1994 (Pree et al. 1998). This was the first documented occurrence

of resistance to pesticides in G. molesta worldwide. Tests with neonate larvae (the targeted

life stage in the field) indicated cross-resistance to most OP insecticides, except acephate

and chlorpyrifos. Acephate was more toxic to resistant larvae than to susceptible larvae,

and chlorpyrifos was equitoxic to both populations. Resistance was highest (>100 fold)

to methyl carbamates, carbaryl, and carbofuran. Cross-resistance to pyrethroids was not

observed. Based on these observations and additional small plot tests, growers switched

to programs of repeated applications of pyrethroids. Concerns were expressed (Pree et al.

1998) that a program of repeated pyrethroid use might accelerate the development of further

resistance. Therefore, an interim resistance management strategy was implemented which

consisted of 1-2 applications of chlorpyrifos for the first annual generation, followed by

pyrethroids for later generations. From 1996-1999, monitoring of resistance changes in

commercial orchards using this program showed that resistance to OP insecticides declined

from about 50% to 12%, while pyrethroid resistance was approximately 16% (Kanga et

al. 2003). While this program successfully reduced the risk that resistance to pyrethroids

might become common, registration of chlorpyrifos was granted only on a temporary

and annual basis, and deregistration is scheduled for 2006 (Pest Management Regulatory

Agency 2003).

We report on the current status of resistance to organophosphorus (OP) and

pyrethroid insecticides in both apple and peach plantings in southern Ontario, provide an

update on the mechanisms of OP resistance, and compare both of these findings to previous

studies. Further, with the impending removal of chlorpyrifos, we also report on the

effectiveness of potential alternative materials and how they might fit into both Integrated

Pest Management and resistance management programs for Oriental Fruit Moth populations

in tree fruit.

Methods

Oriental fruit moth populations

The population of Oriental fruit moth used as the standard susceptible population

was the same colony used in earlier studies (Pree et al. 1998; Kanga et al. 1999) and unless

otherwise indicated was maintained on small green apples as described by Pree (1985).

Insecticide resistance in oriental fruit moth JESO Volume 136, 2005

This colony has been maintained unselected with few infusions of field collected insects

(none since 1985) for approximately 50 years.

The standard resistant population in these tests was collected from a mixed

apple/peach/pear planting near Beamsville, ON in 2002. Initial tests using a standard field

resistance monitoring procedure (Kanga et al. 1999) indicated that approximately 75% of

the population was resistant to OP and carbamate insecticides. The colony was established

on apple from 200-300 larvae. Larvae of this population were selected in each generation

with carbaryl at 150 mg/L using a Potter spray tower using procedures described by Pree

(1979). Newly hatched larvae were held in Petri dishes on ice and sprayed with 5 ml of a

150 mg/L solution of carbaryl in analytical grade acetone. Sprayed larvae were transferred

to apples in a standard rearing container (Pree 1985). Laboratory bioassays with neonate

larvae were conducted with the 9-15" laboratory selected generations.

For tests with methoxyfenozide, which is more active when ingested, we developed

subcolonies of each population adapted to an artificial diet. The diet was modified from

that of Yokoyama et al. (1987) and initially resulted in some larvae mortality (previously

adapted to green apples), but this decreased after 3-4 generations on the diet. Larvae were

used for tests after at least 6 generations on the diet. Three or four Oriental fruit moth

neonate larvae were placed in plastic cups (souffles) (P100, 25 ml capacity, SOLO Cup

Company, Urbana, IL) each containing about 10 ml of diet. Pupae were removed after 3-4

weeks and held in rearing jars (Pree 1985) for adult emergence.

Preparation of artificial diet

The diet consisted of: 3.0 g methyl-p-hydroxybenzoate, 1.8 g sorbic acid, 7.0 g

L-ascorbic acid, 10.5 g fructose, 13.0 g Vanderzant vitamin mix, 17.5 g a-protein (soybean

protein), 35.0 g wheat germ, 70.0 g Brewers yeast, 350.0 g ground pinto beans (BioServ,

Frenchtown, NJ), and 1500 ml distilled water.

Unless indicated, all ingredients were from ICN Biomedical (Aurora, OH). Dry

ingredients were blended for approximately 30 seconds with 1500 ml distilled water

until a smooth consistency was obtained. A 1 L media bottle containing 500 ml distilled

water and 16 g of agar was autoclaved until the mixture boiled, and the warm agar was

thoroughly mixed with the aqueous nutrient mixture. Warm diet mix was transferred to

plastic squeeze bottles and dispensed into individual cups. Cups (25 ml capacity) were

filled to an approximate depth of 1.5 cm, allowed to cool at room temperature, capped when

condensation had disappeared, and stored at room temperature until needed. The quantities

listed here provided approximately 400 individual cups of diet.

BIOASSAYS

Determination of resistance frequencies in field population

Resistance frequencies for OP and pyrethroid insecticides in the various orchard

populations and locations were monitored as described by Kanga et al. (1999; 2003). Adult

males captured in pheromone-baited traps were brought to the laboratory and fed overnight

with a 10% sucrose solution. They were then exposed to insecticides in glass vials as

described by Kanga and Plapp (1995). For tests with the Niagara populations, one or two

moths were placed in each vial and held for 24 hours in a cabinet at 22 + 2°C, 60% relative

humidity (RH), and 16:8 Light:Dark (L:D) cycle. Assays with populations from Norfolk

Pree et al, JESO Volume 136, 2005

=f Se One wets ee

and Essex were conducted in the test areas on a laboratory bench where temperatures were

similar but REE values were lower (about 40%), Adults unable to fly when tossed into the

a were considered dead (Kanga et al, 1999), The diagnostic concentrations used in all

bioassays were those used by Kanga et al, (2003) but were verified as diagnostic prior to

use here, Concentrations used were 0.1 pg/vial for carbofuran (which indicates both OP

and carbamate resistance and used because the higher level of resistance results in a better

separation of resistant and susceptible populations) (Kanga et al, 2003) and 2.5 pg /vial

for eypermethrin (as diagnostic of pyrethroid resistance), Both of these concentrations

killed all of a susceptible population, The carbofuran treatment did not affect any of an

OP resistant population whereas in tests by Kanga et al, (1999), cypermethrin at 2.5 pup/

vial killed approximately 25% of a pyrethroid resistant population, We did not have a

pyrethroid resistant population for comparison, There were 100-250 male moths tested

for each compound per generation per site reported, Data from 3-4 days trapping were

combined and mean survival rate over the generation trapped is presented,

Contact toxicity tests

Tests with contact insecticides on neonate larvae were similar to those described

by Pree et al, (1998), Insecticides, technical or analytical grade, obtained either from the

manutweturers or from Chem Services (West Chester, PA), were applied to first instar larvae

with a Potter spray tower in 5 ml of analytical grade acetone, After treatment, larvae were

held in plastic Petri dishes (Falcon L006, Becton Dickinson, Lincoln Park, NJ) for 2 hours

at 22 4 22°C and 60% REL Larvae that were unable to crawl when prodded were considered

dead, Mortality data from six concentrations of each insecticide with 10 replications of 10

larvae were used to plot regression lines of concentration vs, mortality, Data were subjected

to probit analysis (POLO-PC, Le Ora Software, Berkeley, CA), Resistance ratios were

considered significantly different if the 95% confidence limits at the LC 4) did not overlap,

Insecticide-diet mixtures

For tests with artificial diets, measured amounts of commercial formulations of the

test chemicals, methoxyfenozide (Intrepid 240F, DowAgrosciences Canada Inc, Calgary,

AB) or acetamiprid (Assail 70WP, Dupont Canada tne, Mississauga, ON) were diluted in

10 ml of distilled water and added to 390 ml of treshly prepared diet to provide the desired

final concentration expressed as mg/l active ingredient (ai), The diet and test chemical were

mixed thoroughly ina Waring blender and distributed approximately evenly into 50 SOLO

cups (PLOO, SOLO Cup Company). Two neonate larvae were added to each cup and held

ina cabinet at 22 ¢ 2°C, 60% RH, and a 16:8 L:D regime, Five concentrations plus a water

treated control were used for each chemical and each population with 10 replicates of 5

cups each, Tests were set up over at least 2 days with fresh insecticide, diet preparations

and newly hatched larvae each day, Mortality was assessed after 4 and 6 weeks when

cups Were examined for pupae, Cups containing no pupae were rated as negative or dead,

Cups which held one or 2 pupae were classed as positive (alive), Data were expressed as

the proportion of cups with dead larvae based on 10 replicates of 10 cups each (percent

mortality), Mortality in controls (i.e, control cups which produced no pupae) was 4-6%.,

Concentration:mortality data were analyzed by probit analysis as described for contact

toxicity tests above, Differences between responses were considered significantly different

Insecticide resistance in oriental fruit moth JESO Volume 136, 2005

if the 95% confidence limits at the LC, did not overlap.

BIOCHEMICAL ASSAYS

General esterase

Esterase activity in the susceptible and OP-resistant populations was measured

using a procedure adapted from Herath et al. (1987) that used a-naphthyl acetate as a

substrate. The reaction mixture consisted of 800 ul of a-naphthyl acetate (0.3 mM) in 0.1

M, pH 7.2 phosphate buffer, and 100 ul of insect homogenate. One adult abdomen/ml was

homogenized in ice-cold phosphate buffer and the homogenate centrifuged at 10,000 g for

10 minutes at 4°C. The supernatant was used in assays. The reaction was run in a 1.5 ml

Eppendorf tube in an Eppendorf Thermomixer at 37°C and 450 rpm. The reaction was

stopped after 15 minutes with 80 ul of a solution of 100 mg of Fast Blue B salt in 50 ml of a

5% solution of sodium dodecyl sulfate. The change in absorbance at 450 nm was measured

on an Ultraspec 3100 pro spectrophotometer (Biochrom Ltd, Cambridge, UK). There were

15 replications over 12 different days using 112-121 insects for each population. Protein

concentrations in tissue homogenates were determined by the method of Bradford (1976).

Mean esterase activities were compared using an unpaired t-test (P<0.05) (Sigmastat

Version 2.0, SPSS Inc, Chicago, IL). -

Acetylcholinesterase assays

Acetylcholinesterase (AChE) activity was measured using acetylthiocholine as a

substrate (Ellman et al. 1961). Inhibition of AChE was determined using methods adapted

from Moores et al. (1988) and Pree et al. (2003). For assays, moth heads were frozen at

—70°C for at least 30 minutes and each head was placed into a 1.5 ml Eppendorf tube;

50 ul of 0.1 M phosphate buffer (pH 7.5) was added, and the head was ground for 10-15

seconds. This homogenate was held on ice until used. The reaction mixture was 25 ul of

homogenate, 50 ul of 1 mM 5,5-dithio-bis (2-nitrobenzoic acid) (DTNB) in 0.1 M phosphate

buffer (pH 7.5), and 100 ul of 0.1 M phosphate buffer containing Triton X-100 (10 g/l).

This mixture was equilibrated for 2 minutes in an Eppendorf tube and the reaction started

with the addition of 20 yl of the substrate (1mM acetyl thiocholine iodide). For inhibition

tests, 20 ul of 10% M carbaryl in ethanol was placed in tubes prior to the addition of the

reaction mixture and the ethanol evaporated. The rate of change at OD 405, was measured

for the initial 10 minutes of the reaction in an Ultraspec 3100 Pro spectrophotometer. For

protein determinations, one adult Oriental fruit moth head was ground in 50 ul of 0.1 M

phosphate buffer with no Triton X-100 and 10 ul was used in the Bradford (1976) assay

for protein with bovine serum albumin as the standard. Activity (and inhibition at 10° M

carbaryl) was determined for males and females of both populations. Data were based on

12-14 replications of 96-101 individuals. Differences between means were identified by an

analysis of variance and a Tukey test (P<0.05) (Sigmastat Version 2.0 SPSS, Chicago, IL).

FIELD TESTS OF ALTERNATIVE INSECTICIDES

Trials were conducted at the Jordan Station Experimental Farm of Agriculture and

Agri-Food Canada (AAFC), Jordan, ON. For tests in small plots, treatments were replicated

4 times, assigned to 2 tree plots arranged according to a randomized complete block

design. Based on pheromone trap catches of male moths in adjacent or nearby plantings,

Pree et al, JESO Volume 136, 2005

applications were timed for egg hatch of the first or second generations of Oriental ‘fruit

moths using standard methods (Pree et al. 1983) and a phenology model (Rice et al. 1982)

that used 7.2°C as a base temperature. Trees were spaced 4.6 x 5.5 m and were 3-6 years

old as indicated. Insecticides were diluted to a rate comparable to 3,000 L/ha and trees

sprayed to runoff with a truck-mounted sprayer (Rittenhouse Sprayers Ltd., St. Catharines,

ON) equipped with a Spraying Systems handgun (Spraying Systems Co., Wheaton, IL)

fitted with a D-6 orifice plate. Pressure was set at 2,000 kPa. Nine to thirteen L of spray

mix was applied per plot. Plots were assessed 10-19 days post-spray when all twig and fruit

damage was removed and counted. Data were analyzed using an analysis of variance and

a Tukey test (P<0.05),

In larger scale trials, insecticides were applied to approximately 0.50 ha blocks of

mature cv, Loring peach trees at the AAFC Jordan Station Experimental Farm in season-long

programs. All plots were treated with superior oil (60 L/ha) as a dilute spray (20 L/1000

L water) in April for control of overwintered eggs of European red mite (Panonychus ulmi

(Koch), Acari: Tetranychidae).

In 2003, we assessed 3 programs. The most widely used commercial program

for control of Oriental fruit moth consists of 1.7 kg ai/ha chlorpyrifos (Lorsban 50W, Dow

AgroSciences, Calgary, AB), for generation |, followed by pyrethroids for generations

2 and 3 (program 1). In our tests we used 10 g ai/ha deltamethrin (Decis 5 EC, Bayer

CropSciences, Calgary, AB). Program 2 was 360 g ai/ha methoxyfenozide (Intrepid 240F,

Dow AgroSciences) for generation |, followed by deltamethrin for generations 2 and 3.

Program 3 was chlorpyrifos for generation | followed by methoxyfenozide for generations

2 and 3. Treatments were applied using a Rittenhouse GB Laser P20 sprayer (Rittenhouse

Sprayers Ltd., St. Catharines, ON) set to deliver 840 L/ha and were timed using standard

procedures (Pree et al. 1983) based on data from pheromone-baited traps placed in test

plots or in nearby peach plantings. Insecticides were applied 30 May for generation 1, 16

July and | August for generation 2 and 30 August for generation 3 (and as a preharvest

treatment). Infested terminals were assessed 17 June and 30 July when all of the terminals

on 10% of the trees in each plot were examined for damage by larvae. At harvest, on 8

and 10 September 2003, we examined 10-12 of the ripest fruit on each tree for Oriental

fruit moth damage. Further, 20% of these fruit were cut apart and checked for damage

not visible from surface assessments. Data from twig damage and fruit assessments did

not fit a normal distribution and attempts to transform data were unsuccessful (based on

Kolmogorov-Smirnovy test with Lilliefors’ correction), so were analyzed using a Kruskal-

Wallis test (Sigmastat Version 2.0).

Mite populations were assessed 25 August when 3 replicate samples of 100 leaves

were collected from each plot. Leaves were examined for numbers of European red mite

Panonychus ulmi (Koch) (Acari:Tetranychidae), peach silver mite Acu/us cornutus Banks

(Acari: Eriophyidae), and for predaceous mites (Acari: Phytoseidae). For each sample, 20

leaves were examined under a binocular microscope and an additional 80 leaves brushed

with a Henderson-McBurnie mite brushing machine (Henderson and McBurnie 1943).

Peach silver mite infestations were assessed on 20 leaves/sample. Infestations were rated

on a scale of 0-5: 0 = 0 mites/leaf; 1 = 1-10 mites/leaf; 2 = 11-25 mites/leaf; 3 = 26-50

mites/leaf; 4 = 51-100 mites/leaf; 5 = 101+ mites/leaf. After testing for fit in a normal

distribution, data were analyzed by analysis of variance with differences between treatment

Insecticide resistance in oriental fruit moth JESO Volume 136, 2005

means identified using a Tukey test (P<0.05).

In 2004, 4 programs were assessed: Program 1-chlorpyrifos for generation |

followed by deltamethrin for generations 2 and 3; Program 2-methoxyfenozide for generation

1, with deltamethrin for generations 2 and 3; Program 3-acetamiprid for generation 1,

methoxyfenozide for generations 2 and 3; and Program 4-methoxyfenozide for generation

1, acetamiprid for generations 2 and 3. Rates used were as in 2003 with acetamiprid (Assail

7OWP) applied at 168 g ai/ha. Treatments were timed as described by Pree et al. (1983)

with applications on 21 May for generation 1, 9 July for generation 2, and 6 and 24 August

(preharvest) for generation 3. On 8 June and 28 July, infested terminals were counted and

removed on 10% of the trees, selected randomly, in each block. At harvest, 27 and 30

August, we examined 10-12 of the ripest fruit on each tree for damage by Oriental fruit

moth larvae. As in 2003, we cut apart 20% of these fruit and checked for damage not

visible from surface assessments. Data from assessment of damaged twigs by generation

2 larvae did not fit a normal distribution until transformed (log x +1). Data were analyzed

by an analysis of variance and differences between means were separated by a Tukey test

(P<0.05). Mites were sampled August 25 as described above for 2003. Data were analyzed

as described for 2003.

Results

Status of resistance in field populations

The occurrence of resistance to organophosphorus (OP) and carbamate insecticides

at the Jordan Station site ranged from 31% in generation | in 2004 to 75% in generation 2

in 2003 (Table 1). The diagnostic concentrations used for tests allowed survival of resistant

insects only. Over the three seasons at the Jordan Station site, resistance rates were similar

in generation | (from 31-39%), but were generally higher than reported for the same location

in 1999 by Kanga et al. (2003) who found a decline to <20% resistance for OP insecticides.

Resistance to OP insecticides was usually highest at the end of each season in generation 3.

Resistance to OP insecticides in the populations from the Niagara peninsula were always

lowest in the first or overwintered generation and lower than in the third generation of the

previous year at the Jordan site where observations were made over 3 years. The Grimsby

site, largely planted with apples (no peach) and the Beamsville site, a mixture of peach,

apple, and pear, showed similar patterns of increased frequencies of OP resistance over the

season. However, OP resistance did not continue to increase over the 3 years of sampling

at the Jordan Station site, nor did resistance levels in generation 3.

Control programs for Oriental fruit moth on peach were chlorpyrifos for the first

generation followed by up to four applications of a pyrethroid (cypermethrin, deltamethrin,

or, lambda cyhalothrin) over the rest of the season. Programs on apple were variable but most

included at least one application of azinphosmethyl] or phosmet. Resistance to pyrethroids

did not increase over the three seasons sampled at the Jordan site and was generally lower

than OP resistance at all locations tested in Niagara. Resistance to pyrethroids was similar

or lower than reported in previous studies by Kanga et al. (2003).

Sites sampled in Norfolk (Table 2) were all apple and all showed the occurrence of

resistance to OP insecticides at frequencies up to 54%. The percent identified as resistant

Pree et al. | JESO Volume 136, 2005

TABLE |. Occurrence of resistance in Oriental fruit moth populations with organophosphorus

(OP) and pyrethroid insecticides in the Niagara peninsula, ON, 2002-2004. The survival rate

is based on 100-120 adult males/ generation for each insecticide. OP resistance determined

with carbofuran, pyrethroid resistance with cypermethrin. Diagnostic concentrations used

killed all susceptible mortality data indicate percent resistant moths.

Survival Rate (%) + SE

OP Pyrethroid

Generation Generation

Site | 2 3 | 2 3

2002

Jordan Station 39.0+17.3 53.04 6.0 59.9+ 7.7 (0) 102 °2.0° 10O+#1.9

Grimsby 29.32 2.9 35.82£13.2 713+ 47 0.92419 28+ 3.5 0

Beamsville 72.2+ 7.5 743+411.6 839+ 4.6 0.8241.7 30+ 3.8 8826.0

2003

Jordan Station 34.84+14.0 75.4+ 5.2 n.a. 0) 12+ 13.9 n.a.

2004

Jordan Station 31.0+ 6.8 380+ 6.9 53.0+ 12.8 0) 10+ 2.0 0

Vineland 45.0.+:19.7'\ 64.04:22:0'7 S8.0% 106 8h3225" htheols 0

declined over the 3 seasons of the test at the Simcoe site but this was the only location

sampled each season and initial samples (2001) were from generation 3 when resistance

was generally higher than in generation | (Kanga et al. 2003). Pyrethroid resistance was

detected at most of the sites.

In Essex county (Table 2), samples were largely from peach plantings and both

the Oxley and Varner populations showed OP resistance but not pyrethroid resistance.

Whether resistance to pyrethroids declined (despite up to 4 applications/season) and OP

resistance increased relative to levels indicated in earlier reports (Kanga et al. 2003) or

whether these results are an expression of fluctuations in resistance frequencies and do

not necessarily represent trends, is not clear. Most of the sites trapped in these studies

had not been previously tested but the Jordan Station site had been extensively tested by

Kanga et al. (2003). It seems unlikely that changes in bioassay techniques were responsible

for these observed changes because concentration: response regressions were redeveloped

for these tests and produced results similar to those used in earlier assays by Kanga et al.

(2003). Diagnostic concentrations were the same as in their earlier report. These data may

support the argument of Tabashnik et al. (2000) that the frequency of resistance does not

necessarily increase each season despite considerable selection pressures. They reported

that Bt resistance frequencies in pink bollworm Pectinophora gossypiella (Saunders)

(Lepidoptera: Gelechiidae) did not increase as expected over three seasons (1997-1999)

Insecticide resistance in oriental fruit moth JESO Volume 136, 2005

TABLE 2. Occurrence of resistance to organophosphorus (OP) and pyrethroid insecticides

in Oriental fruit moth populations from Norfolk and Essex counties, ON, 2001-2003.

Survival rate was based on the responses of 100-250 males.

Survival Rate (%) + SE

Site Generation OP Pyrethroid

Norfolk County

2001

Walsh 3 49.1+ 4.9 3.9 22.7

Simcoe 3 29.129.5 11 1.3

Vittoria 3 29.8+ 5.0 2-6 3:3

2002

Walsh ] 54.0 + 16.0 5.04 3:4

Simcoe bisecy 20.0+ 9.9 24448

Vittoria sl 36.3 + 20.1 TA+47

2003

Renton | 24 T2283 0

Simcoe | 13:7 4° 63 0

Essex County

2003

Oxley l 60.0 + 10.9 0

Varner l 41.9 + 14.7 0

despite high levels of selection from Bt cotton.

Laboratory tests

Laboratory bioassays of neonate larvae with the population collected from

Beamsville and selected in the laboratory (Table 3) indicated that the characteristics of the

resistance levels expressed were similar to those reported for populations collected in 1994

(Pree et al. 1998). Resistance to the OP insecticides azinphosmethyl and phosmet was

expressed at low levels, chlorpyrifos was equally toxic to both resistant and susceptible

populations, and acephate was more toxic to the resistant population than the susceptible

population. Resistance to the methyl carbamates carbaryl and carbofuran was expressed

at high levels and could not be quantified. There was no cross resistance to the pyrethroid

cypermethrin. All of these observations indicated that the resistance was not different

from that determined in the initial report (Pree et al. 1998). Additional tests with the

neonicotinoids, imidacloprid, and acetamiprid, indicated these were equally toxic to both

susceptible and resistant populations.

, 2005

JESO Volume 136

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Insecticide resistance in oriental fruit moth JESO Volume 136, 2005

Tests with insecticides incorporated into an artificial diet indicated that acetamiprid

was equally toxic to both susceptible and resistant populations but that methoxyfenozide

was slightly more toxic (1.7 fold) to the susceptible population (Table 4). By this procedure,

methoxyfenozide was more toxic (to both susceptible and resistant populations) than

acetamiprid. Resistance to methoxyfenozide, and to its analog tebufenozide, has been

shown in populations of the obliquebanded leafroller Choristoneura rosaceana (Harris)

(Lepidoptera: Tortricidae) that expressed resistance to OP insecticides (Waldstein et al.

1999; Pree et al. 2003).

Resistance mechanisms

Esterase activity in adult abdomens was higher by a factor of 1.6 in the resistant

population (20.0 + 2.8 umoles min! mg" protein versus 12.5 + 1.1 for the susceptible

population). Differences between resistant and susceptible populations in 1994 (Kanga

et al. 1997) were 3.9 fold. AChE activity was higher in both adult males and females

of the resistant population (Table 5) but was not different between the sexes for either

population. Measurements of AChE inhibition with carbaryl at 10°*M indicated >90%

inhibition in both sexes for the susceptible population and <20% inhibition of AChE in the

resistant population. In earlier studies Kanga et al. (1997) reported no differences between

populations in total AChE activity but did report large differences in inhibition between

susceptible and resistant populations. Both elevated general esterases and the presence

of an AChE insensitive to inhibition remain likely important factors in this resistance to

OP and carbamate insecticides. We did not assay other possible resistance mechanisms.

Elevated levels of oxidases and glutathione S-transferases were not shown to be involved in

the resistance in earlier studies (Kanga et al. 1997).

Field tests of alternative insecticides

In small plot trials in 2002, methoxyfenozide applied at 360 g ai/ha was as effective

as the standard chlorpyrifos or deltamethrin (Table 6). Lower rates of methoxyfenozide were

less effective than the standards. Populations of Oriental fruit moth at this site were 40-55%

TABLE 4. Toxicity of insecticides incorporated into diet fed to susceptible and resistant

populations of Oriental fruit moth larvae. Resistance ratio was calculated as LC,, resistant

strain divided by LC, susceptible strain.

Insecticide Population Slope + SE LC, x? Resistance

(n=500) mg/kg Ratio

methoxyfenozide Susceptible 4.1+0.41 0.028 (0.024-0.032) 9.3

Resistant Sob0:23 0.049 (0.042-0.057) 5.8 1.7

acetamiprid Susceptible 6.9 + 0.73 0.45 (0.41-0.48) 6.6

Resistant 627-41.07 0.48 (0.45-0.52) 12.2 1.1

Pree et al. JESO Volume 136, 2005

TABLE 5. Inhibition of acetylcholinesterases in susceptible and resistant populations of

Oriental fruit moth.

Population Sex Mean Rate + SE Mean % Inhibition + SE

10* M Carbaryl . (n = 95-134 heads)

umoles/min/mg protein

(n = 96-101)

Susceptible Male 28.1 +2.3 a! 91.0+2.7

Female 29.4+3.8a 91.7+2.6

Resistant Male 52.3+4.8b 17.2+7.4

Female 429+43b 15.8 + 6.0

'Same letters are not significantly different (Tukey test (P<0.05)).

TABLE 6. Control of Oriental fruit moth damage on peach in small field plots, Jordan

Station, ON, 2002. OFM Damage/Plot includes damage to twigs and fruit.

Treatment Formulation Rate OFM Damage/Plot

a.i./ha

Generation!

chlorpyrifos Lorsban SOWP 1700 g 28. <

methoxyfenozide Intrepid 2F 120g 12.85 b

methoxyfenozide Intrepid 2F 240 g | 9.1 be

methoxyfenozide Intrepid 2F 360 g 8.5 be

Control - 24.0 a

Generation 2°

deltamethrin Decis SEC 10g 19:21.eF

methoxyfenozide Intrepid 2F 120 g 127.0 b

methoxyfenozide Intrepid 2F 240 g 127.0 b

methoxyfenozide Intrepid 2F 360 g 101.5 Se

Control - 239.5 a

' Applied 3 June 2002, to cv. Loring, Damage assessment 27 June 2002.

> Same letters are not significantly different (Tukey test, P<0.05).

> Applied 10 and 23 July 2002, to cv. Loring, Damage assessment 2 August 2002.

Insecticide resistance in oriental fruit moth JESO Volume 136, 2005

TABLE 7. Control of Oriental fruit moth damage on peach in small field plots, Jordan

Station, ON, 2004. OFM Damage/Plot includes damage to twigs and fruit.

Treatment Formulation Rate Total OFM Damage

a.i./ha

Generation!

deltamethrin Decis EC 10g 1.0 b?

acetamiprid Assail 70WP 47.2 ¢ 13.8b

acetamiprid Assail 70WP 168.8 g 11.5b

acetamiprid Assail 70WP 176g 4.3b

Control - 30.3 a

Generation 2?

deltamethrin Decis 5EC 10g 1.3b

acetamiprid Assail 70WP 47.2 ¢ 3.0b

acetamiprid Assail 70WP 168.8 g 1.8b

acetamiprid Assail 70WP. 172 ¢g 15b

Control - 98a

' Applied 21 May 2004, 124 DD base 7.2 °C after first male capture, cv. Elberta, Damage

assessment 9 June 2004.

* Same letters are not significantly different (Tukey test, P<0.05).

> Applied 9 July 2004, 617 DD, base 7.2°C after first male capture, cv. Elberta, Damage

assessment 22 July 2004.

resistant to OP insecticides and >5% were resistant to pyrethroids (Table 1). Infestations

were higher in tests with the second generation in 2002. In similar tests in 2004 (Table 7),

acetamiprid at three different rates was as effective as the standard deltamethrin.

In season long tests of various Oriental fruit moth control programs in 2003 and

2004 (Table 8) all of the programs effectively prevented damage to twigs by first generation

larvae. As in most seasons, damage was less in generation | than later in the season. In

2003, in generation 2, damage to terminals was higher in methoxyfenozide-treated plots.

This did not result in significantly higher damage at harvest although damage to fruit was

slightly higher than the grower accepted threshold of 1%. In 2004 (Table 8) in generation 2,

damage to terminals was again higher in methoxyfenozide-treated plots than in deltamethrin

or acetamiprid-treated plots. At harvest, damage to fruit was higher in plots treated with

methoxyfenozide in generations 2 and 3. All other programs had <1% fruit damaged at

harvest.

In 2003, populations of European red mite did not reach threshold or action levels

(5-10 mites/leaf in July) (Anonymous 2004) under any of the programs tested in 2003 (Table

9). Numbers of European red mites were higher in 2004 but did not exceed the economic

threshold. However, in 2003, populations of the peach silver mite and numbers of beneficial

mites were higher in all plots treated with deltamethrin. In 2004, numbers of peach silver

JESO Volume 136, 2005

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Pree et al. JESO Volume 136, 2005

mites did not increase as in 2003 and remained at low numbers in all plots. Beneficial mites

were found in all plots but numbers were generally higher where phytophagous mites (either

P.ulmi or A. cornutus) were available as a food source. In 2003, numbers of beneficial mites

were higher in plots treated with deltamethrin in generations 2 and 3 of Oriental fruit moth,

plots which held large numbers of peach silver mites. In 2004, numbers of beneficial mites

were highest in plots treated with acetamiprid in generations 2 and 3.

Discussion

The initial goal of the resistance management strategy for Oriental fruit moth

established after the episode of resistance to OP insecticides in 1993-1994 was to maintain

susceptibility to pyrethroids, the only effective alternative at that time. That program

appears to have been successful. The percentage of the population expressing resistance

to pyrethroids declined from levels reported in earlier studies and resistance to pyrethroids

was not found at all sites. Resistance to OP insecticides was found at all sites and was at a

higher frequency than previously reported at one extensively monitored site. This occurred

despite avoidance of the OP insecticides, azinphosmethyl and phosmet, that had previously

been used on peach for control of Oriental fruit moth. Chlorpyrifos was equally toxic, as

in previous studies, to both resistant and susceptible populations. In the last 10 years, the

Oriental fruit moth has become a pest on apples where OP insecticides have continued to be

used for other pests and this may be the source of OP resistant insects. Pyrethroids are not

used extensively on apples.

The potential impact of the impending removal of chlorpyrifos after 2006 may be

ameliorated by the use of either methoxyfenozide or the neonicotinoid acetamiprid. The

cross resistance to methoxyfenozide identified here was expressed at low levels (1.7 fold at

the LC,,) and, at higher rates, this compound effectively controlled Oriental fruit moth in

both small plot and program trials against the first generation. Later applications against

the second and third generations, especially in the program trials, were less effective.

Acetamiprid was effective throughout the season. If methoxyfenozide were reserved as a

replacement for chlopyrifos for use against generation |, this would hold the neonicotinoid

acetimiprid and/or pyrethroids or rotations of these two groups for the rest of the season.

The use of pyrethroids in IPM programs has often been discouraged because of their impact

on beneficial mites and the associated outbreaks of phytophagous mites (Croft 1990). In

the program trials reported here, numbers of European red mites did not exceed acceptable

thresholds but high populations of peach silver mites were associated with pyrethroid use

in 2003. Further evaluation of the impact of these products on beneficial mite populations

in peach and apple ecosystems is necessary but Beers et al. (2005) have shown increased

populations of phytophagous mites following repeated applications of various neonicotinoids.

In any case, the addition of these two new insecticides will provide an opportunity not only

to manage or prevent resistance to pyrethroids, but if all three groups of chemicals are

utilized, should also delay the development of resistance to these new products. There

is also an alternative control program (Trimble et al. 2001) that involves the integration

of insecticides for the first generation with mating disruption for later generations. That

program would likely provide the best long-term resistance management strategy for

Insecticide resistance in oriental fruit moth JESO Volume 136, 2005

Oriental fruit moth on peach in Ontario.

Acknowledgements

We thank Jaqueline Bacsek, Leah Hamilton, Kirstin Weerdenberg, Jill Shupe, and

Wayne P. Roberts for assistance in bioassays of field populations and laboratory rearing. We

thank Kathleen Jensen for assistance in organizing the tables and standardizing the text.

References

Anonymous. 2004. Fruit production recommendations 2004. Ontario Ministry of Agriculture,

Food and Rural Affairs. Publication 360. 294 pp.

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Microclimate and mite predation and oviposition JESO Volume 136, 2005

INFLUENCE OF GREENHOUSE MICROCLIMATE ON

NEOSEIULUS (AMBLYSEIUS) CUCUMERIS

(ACARI: PHYTOSEIIDAE) PREDATION ON

FRANKLINIELLA OCCIDENTALIS (THYSANOPTERA:

THRIPIDAE) AND OVIPOSITION ON GREENHOUSE

CUCUMBER

T. JONES, J. L. SHIPP'’, C. D. SCOTT-DUPREE, C. R. HARRIS

Department of Environmental Biology, University of Guelph,

Guelph, Ontario, Canada, NIG 2W1

email: shippl@agr.gc.ca

Abstract J. ent. Soc. Ont. 136: 71—83

The influence of leaf boundary layer vapor pressure deficit (VPD) and leaf

temperature on the predation.rate by Neoseiulus (Amblyseius) cucumeris

(Oudemans) (Acari: Phytoseiidae) on Frankliniella occidentalis (Pergande)

(Thysanoptera: Thripidae) and on the oviposition rate by N. cucumeris on

cucumber leaves were determined for greenhouse cucumber grown under

semi-commercial production conditions. Vapor pressure deficit did not affect

either the predation or oviposition rates by N. cucumeris. Examination of

ambient and boundary layer VPDs revealed that it was difficult to produce

substantial changes in boundary layer VPD in high-gutter greenhouses.

Therefore, the relatively steady state of humid conditions at the leaf boundary

layer resulted in no significant differences in predation and oviposition rates

despite changes in ambient VPD. However, leaf temperature did influence

the predation and oviposition rates by N. cucumeris: both rates increased at

the higher temperature. This suggests that establishing seasonal release rates

should result in increased efficiency of this predator during the cooler periods

of the year.

Introduction

The predatory mite Neoseiulus (Amblyseius) cucumeris (Oudemans) (Acari:

Phytoseiidae) is an important biological control agent used to control Frankliniella

occidentalis (Pergande) (Thysanoptera: Thripidae) on greenhouse cucumbers worldwide

(Ramakers and O’Neill 1999; Shipp and Ramakers 2004). Neoseiulus cucumeris feeds

primarily on first instar F. occidentalis, and adult female mites have been reported in

' Author to whom all correspondence should be addressed.

? Agriculture and Agri-Food Canada, Greenhouse and Processing Crops Research Centre,

Harrow, Ontario, Canada NOR 1G0

jones eial: : JESO Volume 136, 2005

laboratory trials at 25-26°C to reach a predation plateau of between 4.4 and 6.9 first instars

per day on cucumber leaf disks (Shipp and Whitfield 1991; van Houten et al. 1995a; 1995b).

However, the predation rate of the nymphal stages of N. cucumeris on F. occidentalis is

more complex and often depends on the presence of an adult NV. cucumeris to assist in killing

the larger prey host or to share a killed first instar (Cloutier and Johnson 1993). Control of

F: occidentalis at the recommended rates for release of N. cucumeris is often not achieved

until 5-6 weeks after the mites are released into the greenhouse (Shipp et al. 1996).

Improving the predation efficiency of N. cucumeris should result in faster, more

effective control of F. occidentalis populations. Abiotic conditions such as temperature and

humidity influence the rate of predation by mites (Stenseth 1979; Ball 1980; Everson 1980;

Hardman and Rogers 1991). Higher temperatures are believed to cause greater predation

rates as a result of the increased energy demand of the predator, which translates behaviorally

into hunger and its associated activities, such as foraging (Everson 1980). Laboratory

evidence indicates that vapor pressure deficit (VPD) (i.e., the difference between saturate

and actual water vapor pressure at a specific air temperature) affects predation rates as well.

Stenseth (1979) reported that Phytoseiulus persimilis Athias-Henriot (Acari: Phytoseiidae)

provides more effective control of Tetranychus urticae (Koch) (Acari: Tetranychidae) at

higher temperatures and humidities (27°C and 70-85% RH). The predation rate by N.

cucumeris on F: occidentalis was strongly influenced by VPD in laboratory trials (Shipp et

al. 1996). A quadratic model fitted to the predation responses of adult N. cucumeris on first

instar F. occidentalis over a VPD range of 0.4-3.94 kPa showed that the greatest predation

rates occurred at either end of the VPD range.

The efficacy of a biological control agent also depends on its ability to reproduce in

its environment. Several factors can affect the ovipositional behaviour of predatory mites.

A mated N. cucumeris can oviposit 1.3 to 2.5 eggs per day depending on plant host, climate,

and food (Castagnoli and Simoni 1990; Castagnoli and Liguori 1991; van Houten et al.

1995a; 1995b).

In the past, the majority of studies examining the influence of temperature and humidity

on predation and oviposition rates were conducted in controlled environment chambers in

order to precisely control the climatic parameters being investigated. However, insects and

mites usually spend most of their time within the boundary layer of plant leaves (i.e., 0.5-5.0

mm from the leaf surface depending on wind speed, leaf shape, size, and hairiness), which

can have temperature and humidity conditions that are quite different from greenhouse

ambient conditions (Nobel 1974; Ferro and Southwick 1984). Ferro et al. (1979) showed

that leaf temperatures on apple trees could reach 25°C on cool, clear days while ambient

air temperature was only 15°C. Conversely, on hot sunny days when the air temperature

was 39°C, leaf temperature was only 26°C. The changes between greenhouse macroclimate

and microclimate at the plant surface are not as great as in the field situation since the

climate in the greenhouse can be controlled using computerized climate control systems.

However, the macroclimate in the greenhouse can be very different from the microclimate

at the phyllopane (leaf surface) due to thermal, concentration, and velocity boundary layers

that can result in steep gradients in temperature, VPD, and CO, (Jewett and Jarvis 2001).

Boulard et al. (2002), in studying the influence of tomato leaf boundary layer climate on

the implications of microbial control of whiteflies, found that relative humidity measured at

5 mm from the leaf surface in a tomato greenhouse could be 20-30% greater than ambient

Microclimate and mite predation and oviposition JESO Volume 136, 2005

measurements. The objectives of the present study were to determine the influence of

boundary layer VPD and leaf temperature on the rate of predation by N. cucumeris on F.

occidentalis, and on the oviposition rate by N. cucumeris on greenhouse cucumber.

Methods

Experimental treatments

Trials were conducted at the Greenhouse and Processing Crops Research Centre,

Harrow, Ontario on greenhouse cucumber (Cucumis sativus L.) (cv. Bodega) in three

glasshouse compartments (7x13 m) from May 2000 to September 2001. A planting density

of 1.5 plants/m* (seven double rows with 10 plants/row/compartment) was used for all

trials. The outside rows and end plants for each row served as guard plants and were not

used in any measurements. The Harrow Fertigation Manager (Climate Control Systems

Inc., Leamington, ON) was used to irrigate and fertilize the plants according to commercial

recommendations (Ontario Ministry of Agriculture and Food 2001).

Three ranges of ambient VPD treatments were evaluated at each of two ambient

temperatures regimes to simulate winter crop production conditions (21°C day and 20°C

night) and summer conditions (25°C day and 22°C night) as measured at top canopy height

(2.2 m). It was not possible to use exactly the same ambient VPD values for the range of

humidity treatments for the summer and winter trials because greenhouse ambient VPD is

directly affected by outside humidity conditions. However, the differences among ambient

VPD treatment values at the top canopy among the three greenhouses for the summer

and winter trials were essentially similar (0.26-0.65 kPa) over all trials. The three VPD

treatments were achieved by randomly assigning one of three humidity settings to each

of the three greenhouses that were set at the same temperature regime (summer or winter

production conditions). Thus, each trial consisted of three greenhouses at the same ambient

temperature (21 or 25°C), but with each greenhouse at a different humidity. The trials were

replicated over time. An Argus Greenhouse Climate Management System (Argus Control

System Ltd., White Rock, BC) was used to maintain set point temperature and humidity

conditions at the top canopy height.

All predation trials were conducted on the undersurface of leaves at two heights in

the canopy (middle: 1.5 m, top: 2.2 m) and oviposition trials were conducted at one height

in the canopy (middle: 1.5 m). VPD values at middle canopy were always higher than

at top canopy. This relationship is common in greenhouse vegetables due to the lower

light intensity and wind velocity within the canopy and has been shown for both low and

high-gutter greenhouses (Jewett and Jarvis 2001; Zhang and Shipp 2002). At mid-canopy

height in each greenhouse, ambient temperature and humidity were measured with a Hycal

temperature/humidity probe (Hycal Co., El Monte, CA). Leaf temperatures were monitored

using infra-red thermocouple (IRt) sensors (Omega, Laval, PQ), that were placed 1-2 cm

from the leaf surface. These climate parameters (ambient temperature and humidity, and

leaf surface temperature) in combination with the greenhouse cucumber plant surface

climate model (PSCLIMATE) developed by Zhang et al. (2002), were used to calculate

ambient and boundary layer VPDs.

Jones et al. JESO Volume 136, 2005

Predation trials with Neoseiulus cucumeris

A single, 1-2 day old, mated female mite and 15 first instar / occidentalis were

placed on the undersurface of a cucumber leaf in a plexiglass clip cage (0.7x4.0 cm) (Fig.

1). The cage had thrips-proof screening on one end on the lower surface of the leaf and a

padded plastic plate on the top of the leaf. A fold back clip was used to hold together both

pieces of the cage. To ensure that all test mites were at the same age, eggs of N. cucumeris

were collected by sifting commercially purchased cultures of Thripex-plus (Koppert Canada,

Leamington, ON) using a fine mesh screen. Eggs that passed through the screen were

collected on the bottom of a 9 cm Petri dish and were placed using a moistened camel’s hair

brush on the ventral side of a kidney bean (Phaseolus vulgaris L.) leaf. The leaf was then

floated on distilled water on the bottom of a 14 cm Petri dish to maintain a high humidity

and to prevent the mites from leaving the leaf. The leaf was held in the centre of the dish

by placing TangleFoot (Adhesive Pest Management and Tree Protection Products, Grand

Rapids, MI) between the leaf and the bottom of the dish. Cohorts of 30 eggs were placed

beneath a small piece of leaf (20 mm*’) that was placed at the centre of each larger leaf.

The dishes were incubated at 25 + I1°C and L12:D12. The mites were fed frozen first and

second instar /) occidentalis daily and were transferred to a new leaf after each molt. Adults

appeared approximately 5-6 days after hatching. To ensure mating and starvation, the mites

FIGURE 1. Screen leaf cage used in the predation and oviposition trials as viewed on the

undersurface of a cucumber leaf.

Microclimate and mite predation and oviposition JESO Volume 136, 2005

were placed in a vial (1 female: 1 male) without food for 24 h before a trial.

First instar F. occidentalis were used in the predation trials, as this is the preferred

prey stage for N. cucumeris (Shipp and Whitfield 1991). To obtain first instars, adult F

occidentalis were placed on the ventral side of kidney bean leaves that were placed on a

piece of filter paper and cotton, saturated with distilled water, on the bottom of a 9 cm Petri

dish. A Petri dish cover with thrips-proof screening on one end was placed over the Petri

dish and secured with a large, fold-back clip to confine the F) occidentalis. The Petri dish

was placed in a controlled environmental chamber at 27 + 1°C and 80% RH. The adults

were removed after 24 h and first instar F. occidentalis were removed with a moistened

camel’s hair brush approximately 3-4 days later.

The predation trials were conducted for 24 h, after which the cage and leaf area inside

the cage were removed and examined using a dissecting microscope (50X magnification)

for the number of dead and live thrips. A thrips was considered dead, if it was shriveled

or did not move when touched with a probe. The status of the mite (live or dead) also was

noted and any cages in which the mite was dead were not included in the data analysis. For

each trial, a leaf cage was also set up with 15 first instar thrips and no predatory mites at

each canopy height for each VPD treatment to determine the survival rate of F. occidentalis

over the 24 h predation period. Predation trials were replicated six to nine times for each

temperature and VPD treatment over the summer and winter crop production periods with

two to four cages per plant height (middle and top canopy) in each trial. For each predation

trial, middle and top canopy leaf cages were paired together on the same plant. Different

cucumber plants were used for each pairing and for each trial.

Oviposition trials with Neoseiulus cucumeris

A single 1-2 day old, mated female N. cucumeris which was starved for 24 h was

transferred to the undersurface of a cucumber leaf at middle canopy in each of the treatment

greenhouses. The same cages as those used in the predation trials were set up with one

N. cucumeris per cage per treatment. Frozen second instar F- occidentalis, in excess of

what a mite would consume (>10 thrips/mite/day), were placed in the cage as food. Every

24 h for 7 days, the leaf and cage were examined using a dissecting microscope (50X

magnification), and the number of oviposited eggs and status of the mites (live, dead, or

missing) was recorded. For examination of the leaf cage, the leaf area around the cage

was cut and the cage and excised leaf area were returned to the laboratory for observation.

After checking the leaf and cage, the cage and mite were placed on a new leaf in the same

greenhouse. Cages that had missing mites were discarded. This procedure was replicated

three times at each temperature and VPD treatment with 8-10 cages per replication.

Data analysis

All count data were square root transformed before analysis; untransformed data

are reported in the tables and graphs. The impact of leaf surface temperature, crop canopy

height, and leaf boundary layer VPD on the daily predation rate (prey/predator/day) by N.

cucumeris on F. occidentalis, and on the oviposition rate (eggs/female/day) by N. cucumeris

was analyzed using an ANCOVA. The ANCOVA was conducted with temperature and

canopy height as the main factors for the predation trials, with temperature as the main

factor for the ovipositional trials, and VPD as the covariate factor for both experiments

Jones et al. JESO Volume 136, 2005

(PROC GLM, SAS Institute 1995).

Results and Discussion

Influence of vapor pressure deficit on predation and oviposition rates of Neoseiulus

cucumeris

Mean (+ SE) ambient temperatures and corresponding leaf boundary layer VPDs

and leaf temperatures for the predation and oviposition trials are presented in Table 1. The

survival rate for first instar F’ occidentalis that were placed in the control leaf cages was

always greater than 97.5%, indicating essentially zero mortality of the first instars when N.

cucumeris was not present in the leaf cages. ANCOVA showed that the predation rate of

N. cucumeris at the top and middle canopy heights was not significantly different (F, =

1.76, P= 0.19 ) (Table 2). The mean numbers of first instar F’ occidentalis killed by N.

cucumeris were not significantly affected by the leaf boundary layer VPDs (F’, , = 1.42, P=

0.24) (Table 2). All first and second order interactions were also not significant.

Previous studies found that mite predation rates can be affected by different air

humidity regimes (Mori and Chant 1966a; Shipp et al. 1996; Rott and Ponsonby 2000).

At high VPDs (low humidities), mites become dehydrated and as a result, feed more to

compensate for this loss of water (Boudreaux 1958). Mori and Chant (1966b) found that

the predatory mite, P persimilis, and its prey, 7: urticae, are more active at higher VPDs

which results in more frequent encounters between predator and prey and thus higher rates

of predation. However, Shipp et al. (1996) reported that first instars of F) occidentalis

were less active at high VPDs, while N. cucumeris remained active. Neoseiulus cucumeris

ceased moving after 12 h at VPDs > 2.12 kPa. In the present study, leaf surface VPDs were

always low (< 0.69 kPa) and thus, the mites or thrips were not exposed to dehydrating water

stress conditions. In addition, the thrips could obtain water by feeding on the cucumber

leaves.

The results from Shipp et al. (1996) indicated that even over the limited VPD range

tested in the present greenhouse trials, predation rates should have increased with decreased

VPD. In the controlled environmental chamber trials, temperature and VPD were constant,

but in the greenhouse trials, VPDs fluctuated slightly (up to+ 7%). Kramer and Hain (1989)

and van Houten and van Lier (1995) reported that mite survival increased when mites

were exposed to fluctuating versus constant humidity conditions. Also, due to reduced air

movement in the cage compared to an open leaf, the boundary layer in the leaf cages may

be slightly greater than would be predicted using the PSCLIMATE model to determine

VPD at the leaf surface. However, the basic premise for air movement in a boundary layer

is “still” air.

The effect of leaf boundary layer VPDs on the number of eggs oviposited daily

by N. cucumeris was also not significant (F | a l.21, P=:0.29) (Table 3). There is no

published information on the effect of VPD on the oviposition rate of predatory mites,

although laboratory trials have found that Tetranychus spp. have an increased oviposition

rate under “dry” conditions (Boudreaux 1958). The range of boundary layer VPDs in our

study was probably too narrow to detect any influence of VPD on oviposition rates by N.

cucumeris.

JESO Volume 136, 2005

Microclimate and mite predation and oviposition

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Jones et al. | JESO Volume 136, 2005

TABLE 2. ANCOVA for the effect of temperature and canopy height (main factors) and

leaf bounder layer VPD (covariate) on the daily predation rate (number of first instar

Frankliniella occidentalis/female/day) by Neoseiulus cucumeris.

Effect df F Probability

Temperature 25.87 < 0.0001

Height l 1.76 n.s.

Temp* Height l 0.33 n.s.

VPD l 1.42 n.s.

VPD*Temp l 0.05 n.s.

VPD* Height l 1.39 n.s.

VPD*Temp*Height l 0.67 n.s.

Error 81

TABLE 3. ANCOVA for the effect of temperature (main factor) and leaf boundry layer

VPD (covariate) on the daily oviposition rate (number of eggs/female/day) by Neoseiulus

cucumeris.

Effect df F Probability

Temperature | 12.18 0.0036

VPD l 1.21 n.s.

VPD*Temperature l 0.18 n.s.

Error 14

Examination of VPD data during the trials reveals how stable boundary layer VPD

is without extreme changes in the ambient conditions (Table 1). Extreme climate changes

are detrimental to greenhouse crop production, and it is believed that the use of high-gutter

(4.2-5.4 m) greenhouses have substantially reduced the occurrence of extreme fluctuation

in greenhouse climate due to a “buffer” layer of air between the crop and the outside of

the greenhouse (Jewett and Jarvis 2001; Hao et al. 2005). Thus, in high-gutter greenhouse

vegetable production systems as used in this study, ambient VPDs for recommended

commercial production practices (0.4-0.8 kPa [Ontario Ministry of Agriculture and Food

2001]) have a minimal impact on plant boundary layer VPD, which is usually in a range that

seems to be too small to have a significant impact on predation or oviposition rates by N.

cucumeris. However in low-gutter greenhouses (< 2.5 m), especially with side ventilation,

boundary layer VPD is much more impacted by ambient VPD and can even approach

ambient conditions depending on wind speed (Boulard et al. 2004).

Microclimate and mite predation and oviposition JESO Volume 136, 2005

Influence of temperature on predation and oviposition rates of Neoseiulus cucumeris

ANCOVA of the predation data showed that the mean number of first instar F.

occidentalis killed by N. cucumeris was significantly influenced by leaf temperature

ip 7125.87, P< 0.0001) (Table 2). A predation rate of 8.5-8.7 thrips/day at 24°C was

approximately double the rate at 20°C. The increase in predation rate with increased

temperature corresponds with laboratory trials conducted on different leaf surfaces with

other predatory mite species (Stenseth 1979; Ball 1980; Everson 1980; Hardman and Rogers

1991). Leaf temperature influences the body temperature of the predatory mite as well as

the food conversion rate (Sabelis 1981). Temperature has been shown to affect the rate

of gut emptying, the attack rate, and handling time of the prey (Thompson 1978, Everson

1980; Sabelis 1981). Higher temperatures increase the metabolic rate of predators and

thereby decrease the digestive pause between prey (Nakamaru 1977). Hungry predators

have more successful capture rates, are more active, and search more vigorously for prey

(Sandness and McMurtry 1972). Under higher temperatures, N. cucumeris is more active

than F. occidentalis (Shipp et al. 1996; T. Jones, unpublished data) and would probably have

more frequent encounters with F. occidentalis, resulting in an increased predation rate.

The mean number of eggs oviposited by N. cucumeris was also influenced by

temperature (Table 3). Neoseiulus cucumeris oviposited a significantly greater number of

eggs at the higher temperature when compared to the lower temperature range (F, ,,= 12.18,

P= 0.0036) (Table 4). van Houten et al. (1995a) and Gillespie and Ramey (1988) observed

oviposition rates of 2.2 and 1.5 eggs/day at 25 and 20°C respectively, for N. cucumeris. This

relationship is the result of proportionately decreased digestion time and increased predation

rate at higher temperatures. The increased predation at higher temperatures provides the

mite with the increased energy required for the higher egg production.

TABLE 4. Number of first instar Frankliniella occidentalis (mean + SE) killed over a 24

h period by Neoseiulus cucumeris and the number of eggs laid by female NV. cucumeris at

two leaf temperatures.

Leaf Canopy Predation rate n! Oviposition rate n

temperature height (prey/day) (eggs/female/day)

(°C)

24 Top 8.5+0.30a 21 --

Mid 8.7+0.46a 19 2.14+0.05 a 9

20 Top 4.3+0.26b DS --

Mid 4.4+0.24b 24 1.37 + 0.05 b 9

' Number of replicates over time with two to four cage observations each trial per

greenhouse.

The initial prey density is 15 thrips per cage.

? Number of replicates over time with eight to ten cage observations each trial.

Within each column, means followed by different letters are significantly different at

P< 0.05.

Jones et al. | | JESO Volume 136, 2005

Previous studies investigating the interactions between climate (temperature

and humidity) and predation and oviposition rates by greenhouse predatory mites were

all conducted under controlled conditions in the laboratory. It is important to test these

relationships under conditions that are more similar to commercial production conditions

to ensure that the relationships are still valid. The present experiment is the first study

to evaluate the influence of leaf temperature and boundary layer VPD on predation and

Oviposition rates by NV. cucumeris on a greenhouse crop (cucumber) under semi-commercial

production conditions, Under high-gutter greenhouse production conditions, boundary

layer VPD varied very little from 0.1-0.7 kPa. At this range, VPD did not have a significant

impact on predation or oviposition rates by N. cucumeris, The range for leaf boundary layer

VPD can be much greater under low-gutter greenhouse production conditions, especially

with side ventilation (Boulard et al. 2004). Leaf temperature did have a significant impact

on predation and oviposition rates by NV. cucumeris.

In summary, the current introduction rates of N. cucumeris tor F. occidentalis

do not consider greenhouse climatic or plant surface microclimatic conditions when

recommendations are made to growers. Usually an introduction rate is recommended

depending on the crop and/or the level of thrips infestation, irrespective of the time of

year. This study demonstrated that plant surface microclimate can have a significant impact

on the effectiveness of predatory mites. Leaf surface temperatures of cucumber plants

were different from ambient air temperatures, but were within about | + 0.5°C of ambient

temperature (Table |). Greenhouse climate is accurately controlled using computerized

climate control systems and can be maintained within narrow limits (41°C) within

commercial greenhouses. The 24 h temperature regimes used in our study corresponds to

climate conditions in greenhouse cucumber crops during the winter and summer in Ontario.

However, similar seasonal differences occur for other greenhouse cucumber production

areas in temperate climate regions.

Based on the results from this study, V. cucumeris will provide the most effective

control of - occidentalis when conditions are near the higher end of recommended

production temperatures for cucumbers (17-25°C (Ontario Ministry of Agriculture and Food

2001)). Growers often state that thrips control during the winter months is not as effective as

during summer, or that it takes too long (Shipp, unpublished data). Therefore, during winter

conditions when temperatures are lower, growers should introduce mites more frequently

into the greenhouse. Increased knowledge and understanding of greenhouse climate and

plant surface microclimate, and their effect on insect and mite biology/behaviour will result

in improved effectiveness of biological control programs for greenhouse crops.

Acknowledgements

Authors thank Dr. T. Gillespie (University of Guelph), K. Wang, and Y. Zhang for

their helpful suggestions during the study and K. Rusk for her technical support. This study

was supported in part by Ontario Ministry of Agriculture and Food, Food Systems 2002,

Ontario Greenhouse Vegetable Growers and Agriculture and Agri-Food Canada, Matching

Initiatives Investment.

Microclimate and mite predation and oviposition JESO Volume 136, 2005

References

Ball, J.C. 1980. Development, fecundity, and prey consumption of four species of predacious

mites (Phytoseiidae) at two constant temperatures. Environmental Entomology 9:

298-303.

Boudreaux, H. B. 1958. The effect of relative humidity on egg-laying, hatching, and survival

in various spider mites. Journal of Insect Physiology 2: 65-72.

Boulard, T., M. Mermier, J. Fargues, N. Smits, M. Rougier, and J.C. Roy. 2002. Tomato

leaf boundary layer climate: implications for microbiological whitefly control in

greenhouses. Agriculture and Forest Meteorology 110: 159-176.

Boulard, T., H. Fatnassi, J. C. Roy, J. Lagier, J. Fargues, N. Smits, M. Rougier, and B.

Jeannequin. 2004. Effect of greenhouse ventilation on humidity of inside air and in

leaf boundary-layer. Agriculture and Forest Meteorology 125: 225-239.

Castagnoli, M. and S. Simoni. 1990. Biological observations and life table parameters of

Amblyseius cucumeris (Oudemans) (Acarina: Phytosetidae) reared on different

diets. Redia 73: 569-583.

Castagnoli, M. and M. Liguori. 1991. Laboratory observations on duration of copulation

and egg production of three Phytoseiid species fed on pollen. pp. 231-239 In The

Acari: Reproduction, Development and Life History Strategies. Schuster R. and

Murphy P.W. (eds.) Chapman and Hall, London. 554 pp.

Cloutier, C. and S. G. Johnson. 1993. Interaction between life stages in a phytoseiid predator:

western flower thrips prey killed by adults as food for photonymphs of Amblyseius

cucumeris. Experimental and Applied Acarology 17: 441-449.

Everson, P. 1980. The relative activity and functional response of Phytoseiulus persimilis

(Acarina: Phytoseiidae) and Tetranychus urticae (Acarina: Tetranychidae): The

effect of temperature. The Canadian Entomologist 112: 17-24.

Ferro, D, R. B. Chapman, and D. R. Penman. 1979. Observations on insect microclimate

and insect pest management. Environmental Entomology 8: 1000-1003.

Ferro, D. N. and E. E. Southwick. 1984. Microclimate of small arthropods: estimating

humidity within the leaf boundary layer. Environmental Entomology 13: 926-929.

Gillespie, D. R. and C. A. Ramey. 1988. Life history and cold storage of Ambiyseius

cucumeris (Acarina: Phytoseiidae). Journal of Entomological Society of British

Columbia 85: 71-76.

Hao, X., T. Jewett, J. Zheng, and S. Khosla. 2005. Microclimate and energy consumption in

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Hardman, J. M. and M. L. Rogers. 1991. Effects of temperature and prey density on

survival, development, and feeding rates of immature Zyphlodromus pyri (Acari:

Phytoseiidae). Environmental Entomology 20: 1089-1096.

Jewett, T. J. and W. R. Jarvis. 2001. Management of the greenhouse microclimate in relation

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Kramer, D. A. and F. P. Hain. 1989. Effect of constant- and variable-humidity and

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ununguis (Acarina: Tetranychidae) and Neoseiulus fallacis (Acarina: Phytoseiidae).

Environmental Entomology 18: 741-746.

Jones et al. | JESO Volume 136, 2005

Mori, H. and D. A. Chant. 1966a. The influence of prey density, relative humidity, and

starvation on the predaceous behaviour of Phytoseiulus persimilis Athias-Henriot

(Acarina: Phytosetidae). Canadian Journal of Zoology 44: 483-491.

Mori, H. and D. A. Chant. 1966b. The influence of humidity on the activity of Phytoseiulus

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Nakamura, K. 1977. A model for the functional response of a predator to varying prey

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Nobel, P. S. 1974. Introduction to Biophysical Plant Physiology. W. H. Freeman and

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Ontario Ministry of Agriculture and Food. 2001. Growing Greenhouse Vegetables. Ontario

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Toronto, Ontario, Canada. 116 pp.

Ramakers, P. M. J. and T. M. O’Neill. 1999. Implementation of IPM: Case Studies -

Cucurbits. pp. 435-453 /n Integrated Pest and Disease Management in Greenhouse

Crops. Albajes R., M. Lodovica Gullino, J. C. van Lenteren, and Y. Elad (eds.)

Kluwer Academic Publishers, London. 545 pp.

Rott, A. S. and D. J. Ponsonby. 2000. Improving the control of Tetranychus urticae on

edible glasshouse crops using a specialist coccinellid (Stethorus punctillum Weise)

and a generalist mite (Amblyseius californicus McGregor) as biocontrol agents.

Biocontrol Science and Technology 10: 487-498.

Sabelis, M. W. 1981. Biological control of two-spotted spider mites using phytosetid

predators. Part I. Modelling the predator-prey interaction at the individual level.

Agricultural Research Reports, Agricultural University, Wageningen. Netherlands.

242 pp.

Sandness, J. N. and J. A. McMurtry. 1972. Prey consumption behavior of Amblyseius

largoensis in relation to hunger. The Canadian Entomologist 104: 461-470.

SAS Institute, 1995. SAS/STAT users guide. SAS Institute. Cary, NC. 1606 pp.

Shipp, J. L. and G. H. Whitfield. 1991. Functional response of the predatory mite, Amblyseius

cucumeris (Acari: Phytosetidae), on western flower thrips, Frankliniella occidentalis

(Thysanoptera: Thripidae). Environmental Entomology 20: 694-699.

Shipp, J. L., K. I. Ward, and T. J. Gillespie. 1996. Influence of temperature and vapor pressure

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Frankliniella occidentalis. Entomologia Experimentalis et Applicata 78: 31-38.

Shipp, J. L. and P. M. J. Ramakers. 2004. Biological control of thrips on vegetable crops. pp.

265-276 In Biocontrol in protected culture. Heinz K. M., R. G. Van Driesche, and

M. P. Parrella (eds.) Ball Publishing, Batavia, Illinois. 552 pp.

Stenseth, C. 1979. Effect of temperature and humidity on the development of Phytoseiulus

persimilis and its ability to regulate populations of Tetranychus urticae (Acarina:

Phytoseiidae, Tetranychidae). Entomophaga 24: 311-317.

Thompson, D. J. 1978. Towards a realistic predator-prey model: The effect of temperature

on the functional response and life history of larvae of the damselfly, /schnura

elegans. Journal of Animal Ecology 47: 757-767.

van Houten, Y. M. and A. M. M. van Lier. 1995. Influence of temperature and humidity on

Microclimate and mite predation and oviposition JESO Volume 136, 2005

the survival of eggs of the thrips predator Amblyseius cucumeris. Mededelingen van

de Faculteit Landbouwwetenschappen Rijksuniversiteit Gent 60/3a: 879-884.

van Houten, Y. M., P. C. J. van Rijn, L. K. Tanigoshi, P. van Stratum, and J. Bruin. 1995a.

Preselection of predatory mites to improve year-round biological control of western

flower thrips in greenhouse crops. Entomologia Experimentalis et Applicata 74:

225-234. ;

van Houten, Y. M., P. van Stratum, J. Bruin, and A. Veerman. 1995b. Selection for non-

diapause in Amblyseius cucumeris and Amblyseius barkeri and exploration of the

effectiveness of strains for thrips control. Entomologia Experimentalis et Applicata

77: 289-295.

Zhang, Y, T. J. Jewett, and J. L. Shipp. 2002. A dynamic model to estimate in-canopy and

leaf surface microclimate of greenhouse cucumber crops. Transactions of American

Society of Agricultural Engineers 45: 179-192.

Zhang, Y. and J. L. Shipp. 2002. Manipulating plant moisture conditions using greenhouse

high-pressure fogging. HorTechnology 12: 261-267.

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New records for Rhopalosiphum rufiabdominale JESO Volume 136, 2005

NEW RECORDS FOR RHOPALOSIPHUM RUFIABDOMINALE

(SASAKT) (HEMIPTERA: APHIDIDAE) ON GREENHOUSE

TOMATOES AND PEPPERS

G. M. G. ZILAHI-BALOGH'”, R. G. FOOTTIT?, G. FERGUSON‘, J. L. SHIPP

_ Agriculture & Agri-Food Canada,

Greenhouse & Processing Crops Research Centre,

Harrow, Ontario, Canada NOR 1G0

The rice-root aphid, Rhopalosiphum rufiabdominale (Sasaki) (Hemiptera:

Aphididae), described from upland rice in 1899 (Doncaster 1956), is known to have a

worldwide distribution (Blackman and Eastop 2000). Primary hosts are Prunus spp., while

secondary hosts are monocotyledonous plants in the families Poaceae and Cyperaceae

but include some dicotyledons, especially Solanaceae (Blackman and Eastop 2000). The

heteroecious holocycle between Prunus and roots of secondary hosts was reported from

Japan (Yano et al. 1983; Torikura 1991). Elsewhere, R. rufiabdominale is thought to be

anholocyclic on roots of secondary hosts. In many parts of the world it is a pest of rice and

cereals (Yano et al. 1983; Chapin et al. 2001). Rhopalosiphum rufiabdominale is not known

to overwinter in Ontario, but migrates annually from the southern United States (Paliwal

1980).

In October 2004, numerous aphids were observed on the roots of greenhouse

sweet pepper, Capsicum annuum L. (Solanaceae) at the Agriculture and Agri-Food Canada

Greenhouse and Processing Crops Research Centre in Harrow, ON. Specimens were

identified as R. rufiabdominale by E. Maw (Agriculture and Agri-Food Canada, Eastern

Cereal and Oilseed Research Centre, Ottawa). The following spring in April 2005, large

populations of R. rufiabdominale were again observed at the same location, but on this

occasion on the lower stem of greenhouse grown tomatoes, Lycopersicon esculentum Mill.

(cv Rhapsody) (Solanaceae). Identification was confirmed by R. G. Foottit. All plants

were infested. The aphids were observed moving up the stem from the root zone and were

alate viviparae. Aphids dispersed and disappeared as the season progressed. We believe

that the aphids may have overwintered in the greenhouse on secondary hosts as April is too

early in the season in Ontario for winged migrants to have arrived from the southern United

States. Paliwal (1980) reported R. rufiabdominale on cereals from mid-July onwards in

Ontario. In late October 2005, large populations of alate R. rufiabdominale were observed

again on greenhouse tomatoes on the lower stems of the plants at the same location. By late

November populations declined and few alates were observed alive on the above ground

' Author to whom all correspondence should be addressed.

? Department of Environmental Sciences, Nova Scotia Agricultural College, P.O. Box 550,

Truro, Nova Scotia, B2N 5E3, gzilahi@nsac.ca

> Agriculture and Agri-Food Canada, Eastern Cereal and Oilseed Research Centre, Ottawa,

ON, K1A 0C6

* Ontario Ministry of Agriculture and Rural Affairs, Harrow, ON, NOR 1G0

Zilahi-Balogh et al. | | JESO Volume 136, 2005

portions of the plants. In this instance, it is probable that winged aphids moved into the

greenhouse from outside.

This is the first record of R. rufiabdominale on greenhouse tomatoes and sweet

pepper in Canada. It has been previously reported causing damage on greenhouse grown

zucchini in Italy (Ciampolini et al. 1993). Earlier host records of R. rufiabdominale in the

Canadian National Collection include Picea glauca (roots of spruce seedlings in nursery;

Ladner, British Columbia), Gardenia spp. (roots; Surrey, British Columbia), Triticum spp.

(Ottawa, Ontario), and Sparangium spp. (Ottawa, Ontario).

Little information is known about the potential damage and yield loss due to R.

rufiabdominale on greenhouse tomatoes and sweet pepper. However, its overwintering

presence in Ontario greenhouses has implications outside the scope of protected cultivation,

specifically on cereal crops where its impact has been documented (Jedlinski 1981; Riedell

et al. 2003). This aphid is an effective vector of barley yellow dwarf virus (BYDV) in

Canada (Paliwal 1980). BYDV is distributed worldwide and is considered one of the most

economically important diseases of cereals in the world (Riedell et al. 2003), and can persist

in volunteer cereals and wild grasses (Paliwal 1982). BYDYV is present in cereal crops

in Ontario every year but the incidence varies from one region to another (Paliwal and

Comeau 1987). In Illinois, Jedlinski (1981) reported that subterranean R. rufiabdominale

apterae are capable of overwintering on seedling wheat and transmitting BYDV, and

suggested that undetected aphid colonies may explain BYDV outbreaks in the absence

of conspicuous aphid populations. However, Chapin et al. (2001) found no correlation

between R. rufiabdominale abundance and BYDV incidence and yield loss on the coastal

plains of south Carolina.

Important aphid vectors of BYDV in Ontario are R. padi L., R. maidis (Fitch),

and Sitobion avenae (Fabricius) (Hemiptera: Aphididae), which migrate each year from

the United States (Paliwal and Comeau 1987). However, the potential importance of R.

rufiabdominale should not be overlooked since the transmissibility of BYDV isolates to

cereal hosts by R. rufiabdominale is similar to R. padi (Jedlinski 1981). If R. rufiabdominale

becomes prevalent in greenhouses, it has the potential to move into cereal fields earlier than

other aphid vectors more commonly associated with BYDV in Ontario.

Acknowledgements

We thank E. Maw (Agriculture and Agri-Food Canada, Eastern Cereal and Oilseed

Research Centre, Ottawa, ON) for species identification, and two anonymous reviewers for

their comments that have improved the quality of the manuscript.

References

Blackman, R. L. and V. F. Eastop. 2000. Aphids on the World’s Crops: an identification and

information guide. 2™ edition. John Wiley & Sons, Ltd., Chinchester, UK.

Chapin, J. W., J. S. Thomas, S. M. Gray, D. M. Smith, and S. E. Halbert. 2001. Seasonal

abundance of aphids (Homoptera: Aphididae) in wheat and their role as barley

New records for Rhopalosiphum rufiabdominale JESO Volume 136, 2005

yellow dwarf virus vectors in the South Carolina coastal plain. Journal of Economic

Entomology 94: 410-421.

Ciampolini, M., V. di Perna, and C. Maiulini. 1993. Damage by root aphids to vegetable

crops in greenhouses in Lazio. Informatore-Agrario 49: 59-63.

Doncaster, J. P. 1956. The rice root aphid. Bulletin of Entomological Research, 47: 741-

747.

Jedlinski, H. 1981. Rice root aphid, Rhopalosiphum rufiabdominalis, a vector of barley

yellow dwarf virus in Illinois, and the disease complex. Plant Disease 65: 975-

978. |

Paliwal, Y. C. 1980. Transmission of barley yellow dwarf virus isolates by the cereal root

aphid Rhopalosiphum rufiabdominalis. Canadian Journal of Plant Pathology 2:

90-92.

Paliwal, Y. C. 1982. Role of perennial grasses, winter wheat, and aphid vectors in the disease

cycle and epidemiology of barley yellow dwarf virus. Canadian Journal of Plant

Pathology 4: 367-374.

Paliwal, Y.C and A. Comeau. 1987. Yellow dwarf of cereals. Ontario Ministry of Agriculture

Food and Rural Affairs Factsheet 87-074. http://www.omafra.gov.on.ca/english/

crops/facts/87-074.htm. Last-accessed 6 January 2006.

Riedell, W. E., R. W. Kieckhefer, M. A. C. Langham, and L. S. Hesler. 2003. Root and shoot

responses to bird cherry-oat aphids and barley yellow dwarf virus in spring wheat.

Crop Science 43: 1380-1386.

Torikura, H. 1991. Revisional notes on Japanese Rhopalosiphum with keys to species based

on morphs on the primary host. Japanese Journal of Entomology 59: 257-273.

Yano, K., T. Miyake and V. F. Eastop. 1983. The biology and economic importance of rice

aphids (Hemiptera: Aphididae): a review. Bulletin of Entomological Research 73:

539-566.

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Book review JESO Volume 136, 2005

BOOK REVIEW

For Love of Insects. 2003. by T. Eisner. Harvard University Press, Cambridge, USA.

448 pp + xi. ISBN 0-674-01827-3 (paperback edition: 2005). $19.95 US.

“This book is about the thrill of discovery.” Thus begins For Love of Insects. It

is a book on the wonder of insects, focusing primarily on insect chemicals in an ecological

context. Eisner has strung together stories from a career’s worth of research. The stories

are divided by subject into chapters such as, “Tales from the Website,” on spider webs and

“The Circumventers,” on defensive chemicals. By reading this collection of related stories,

the reader not only learns about the specific systems discussed, but also gains a general

appreciation for the role of chemicals in insect ecology. What is most impressive is that

Eisner has made the book accessible, thought provoking, and a true joy to read without

sacrificing any scientific accuracy.

Eisner is a rare breed, becoming increasingly rarer: a highly successful naturalist

who is equally skilled in the field and laboratory. Exploratory walks in the field draw his

attention to curiosities: Why don’t fish eat whirligig beetles? What kind of insect eats

carnivorous plants? Inspired, he then conducts experiments in the laboratory to answer his

questions, often using elegantly simple methods. He is quick to give credit to colleagues

contributing to discoveries. Even his pet bird, Phogel, gets credit for telling Eisner which

insects were palatable.

Eisner’s insect stories are fascinating, thanks in part to his storytelling ability and

of course in large part to the insects themselves. There are walking sticks that preemptively

spray avian predators before being pecked. There are Eleodes beetles that assume a

handstand defensive posture, exposing their chemical-packed abdomen, and Moneilema

beetles that mimic this posture. There is a dizzying complex of six beetle and four moth

species, all mimicking various morphs of each other, varying in habitat, and even preying

upon each other in some cases. More questions than answers are available for this system

in particular, and Eisner encourages future research with the words “doctoral students take

note.”

The most interesting section outlines Eisner’s research on moths of the genus

Utetheisa. Over the course of five PhD students’ research programs, Eisner’s lab has

been able to compile a thorough account of their life history, from larval food choice to

defensive chemicals to sexual selection to cannibalism. Each time I thought the story was

as intricate as it could be, more discoveries were revealed that were even more exciting.

Eisner anticipates the reader’s mind throughout, and asks aloud the very questions you want

to ask.

One of the best aspects of this book is its photographs. They often tell stories

better than any text can. The spray of bombardier beetles, for instance, can be summed up

in just a few pictures. Small diagrams of relevant chemical structures are provided, but are

bonus material, not necessary for a reader’s understanding of a story. Photographs augment

almost every subject of this book, from scanning electron micrographs of ant appendages

tangled up with polyxenid millipede tufts, to dissection images that illustrate how antlions

avoid eating ant acid sacs. Not only are the photographs beautiful and informative, but they

also allow the reader to simply flip through the pictures in the book years after reading to

Fitzsimmons JESO Volume 136, 2005

jog memory of the book’s content. If your memory is as poor as mine, you will find this a

great benefit.

Although Eisner cares deeply about conservation, he does not focus on it directly.

Instead he endeavours to share his wonder of insects with an implied message: if you

understand the wonder of nature you will care enough to protect it. It is no coincidence that

the chemical structure of Mexican bean beetle defense appears in the final few pages of the

book; its elaborate ring structure will impress anyone regardless of chemistry knowledge.

These ant-deterrent compounds were previously undiscovered, and Eisner drives home the

point that much knowledge can be gained from nature.

My only criticism of the book is a small one. Although the order and family is

mentioned for many of the species in this book, it is not given for all. A standard “(Order,

Family)” accompanying the first mention of each species would encumber readability little,

and would help those readers interested in taxonomic classification.

I strongly recommend this book to any reader curious about insect adaptations.

The writing is accessible enough that inquisitive members of the public can enjoy the book.

Researchers young and old will appreciate not only the scientific content of the book, but

also Eisner’s approach to science itself. He combines non-hypothesis-driven observations

with “biorationality” logical deductions to derive hypotheses, and then tests them with

elegantly simple experiments. He freely offers suggestions of promising areas for future

research. He includes some unpublished results, such as how stink bugs’ saliva weakens

spider webs to facilitate escape. If there is any one thing that encompasses the sentiment

of this book, it is punctuation. Question marks are abundant. Wonder is abundant in this

book, as in nature.

JAY M. FITZSIMMONS

Department of Biological Sciences,

University of Windsor,

Windsor, Ontario, Canada, N9B 3P4

fitzsimj@uwindsor.ca

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’ *

2005 ANNUAL MEETING

The 142™* Annual Meeting of the Entomological Society of Ontario was held at the

University of Toronto on October 21-23, 2005. Approximately 80 people attended the

meeting, which had the theme of “Insects in the Urban Environment”. The Entomological

Society of Ontario thanks all the speakers, participants, and organizers who helped to

make the meeting such a success. The Entomological Society of Ontario is grateful for the

support received from Ontario Ministry of Natural Resources (Forest Health & Silviculture

Section), Engage Agro, Landscape Ontario, Lareseco, Faculty of Forestry, University of

Toronto, BASF, N.M. Bartlett Inc., Syngenta Crop Protection Canada, and the Toronto

Zoo.

ENTOMOLOGICAL SOCIETY OF ONTARIO

The Society founded in 1863, is the second oldest Entomological Society in North America

and among the nine oldest, existing entomological societies in the world. It serves as an

association of persons interested in entomology and is dedicated to the furtherance of

the science by holding meetings and publication of the Journal of the Entomological

Society of Ontario. The Journal publishes fully refereed scientific papers, and has a

world-wide circulation. The Society headquarters are at the University of Guelph. The

Society’s library is housed in the McLaughlin Library of the University and is available

to all members. .

An annual fee of $30 provides membership in the Society, the right to publish in the

Journal, and receive the Newsletter and the Journal. Students, amateurs and retired

entomologists can join free of charge but do not receive the Journal.

A World Wide Web home page for the Society is available at the following URL:

http://www.entsocont.com

FELLOWS OF THE ENTOMOLOCIAL SOCIETY OF ONTARIO

W. W. Bill Judd 2002

C. Ron Harris 2003

Edward C. Becker 2003

APPLICATION FOR MEMBERSHIP

Name:

Address:

Postal Code:

Please send cheque or money order to:

D. Hunt, Secretary, Entomological Society of Ontario

c/o Agriculture and Agri-Food Canada G.P.C.R.C.

2585 County Road 20, Harrow, ON, NOR 1G0

NOTICE TO CONTRIBUTORS

Please refer to the Society web site (http://www.entsocont.com/pub.htm) for current

instructions to authors, which were last printed in Volume 131 (2000), pages 145-147 and

can be updated at any time. Copies of those instructions are available from the Editor.

CONTENTS

L FROM THE EDITOR oe ee er

II. SUBMITTED MANUSCRIPTS

BOUCHARD, P., T. A WHEELER, and H. GOULET. — Ground beetles (Coleoptera:

Carabidae) from alvar habitats in Ontario. .......................cccceseeeeececeeeeeeeeereeseeeneneeens D729

MARTIN, A. D., R. S. VERNON, and R. H. HALLETT — Influence of colour and —

trap height on captures of adult pea leafminer, Liriomyza huidobrensis (Blanchard)

(Diptera: Agromyzidae), in celery. ....::6..0....).016,c:s Aicckecpaeseeeees ee 25-35

BUCK, M., S. M. PAIERO, and S. A. MARSHALL. — New records of native and

introduced aculeate Hymenoptera from Ontario, with keys to eastern Canadian

species of Cerceris (Crabronidae) and eastern Nearctic species of Chelostoma

(Megachilidae). ...............:.0:sessshesennsadnsbvesdadesensuslaqedesesiesiien ey. ieee aan nan 37-52

PREE, D. J., K. J. WHITTY, M. K. POGODA, and L. A. BITTNER. — Status of

resistance to insecticides in populations of the Oriental fruit moth Grapholita molesta

(Busck) (Lepidoptera: Tortricidae) in southern Ontario. ...................:0:cceeeee 53-70

JONES, T., J. L. SHIPP, C. D. SCOTT-DUPREE, and C. R. HARRIS. — Influence of

greenhouse microclimate on Neoseiulus (Amblyseius) cucumeris (Acari: Phytoseiidae)

predation on Frankliniella occidentalis (Thysanoptera: Thripidae) and oviposition

on greemhouse CUCUAMDET. .<........<...cecgscasertdesaseoesuaesbaduseuekeheseeessennugnanienae ann 71-83

ZILAHI-BALOGH, G. M. G., R. G. FOOTTIT, G. FERGUSON, and J. L. SHIPP

— New records for Rhopalosiphum rufiabdominale (Sasaki) (Hemiptera: Aphididae)

on greenhouse tomatoes and peppe’..........:.0:..<<s<esss-reonsceosvesnssandenvancunshe en 85-87

Il. BOOK REVIEW

FITZSIMMONS, J. M. — For love of insects. 2003. by T. Eisner..................... 89-90

IV. ANNUAL MEETING inside back cover

V. ENTOMOLOGICAL SOCIETY OF ONTARIO inside back cover

VI. APPLICATION FOR MEMBERSHIP inside back bowen

VII. NOTICE TO CONTRIBUTORS inside back cover

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