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Published November 2009

JOURNAL

of the

: ENTOMOLOGICAL SOCIETY

ONTARIO

Volume One Hundred and Forty

2009

~ . ISSN 1713-7845

THE ENTOMOLOGICAL SOCIETY OF ONTARIO q

OFFICERS AND GOVERNORS e

2008-2009 |

President: C. SCOTT-DUPREE

Dept. of Environmental Biology

University of Guelph, Guelph, Ontario NIG 2W1

cscottdu@uoguelph.ca

President-Elect: G. UMPHREY

Department of Mathematics and Statistics

University of Guelph, Guelph, ON NIG 2W1

umphrey@uoguelph.ca

Past President: R. HALLETT

Dept. of Environmental Biology

University of Guelph, Guelph, ON NIG 2W1

rhallett@uoguelph.ca

Secretary: N. MCKENZIE

Vista Centre, 1830 Bank Street, P.O. Box 83025

Ottawa, ON K1V 1A3

nicole_mckenzie@hc-sc.ge.ca

Treasurer: K. BARBER

Natural Resources Canada, Canadian Forest Service

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

kbarber@nrcan.ge.ca

Directors:

D. CURRIE (2007-2009)

Dept. of Natural History, Royal Ontario Museum

Toronto, ON MSS 2C6

dcurrie@zoo.utoronto.ca

H. DOUGLAS

Canadian Food Inspection Agency

960 Carling Ave., Ottawa ON KIA 06C

douglash@inspection.ge.ca

S. KULLIK

Department of Environmental Biology

University of Guelph, Guelph, ON NIG 2W1

sigrun.kullik@sympatico.ca

K. RYALL (2009-2011)

Canadian Forest Service, Great Lakes Forestry Centre,

Natural Resources Canada, 1219 Queen Street East,

Sault Ste. Marie, ON P6A 2E5

Krista.Ryall@NRCan-RNCan.ge.ca

(2008-2010)

(2009-2011)

K. RYAN (2008-2010)

Faculty of Forestry, University of Toronto

Toronto, ON MSS 3B3

kathleen.ryan@utoronto.ca

J. SKEVINGTON (2007-2009)

Agriculture and Agri-Food Canada

960 Carling Ave., Ottawa, ON K1A 0C6

jeffrey.skevington@agr.gc.ca

St. Catharines, ON L2T 3M7

ESO Regional Rep to ESC: H. DOUGLAS

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Librarian: J. BRETT

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1 Stone Road West, Guelph, ON NIG 4Y2 a

jennifer.allen@ontario.ca a

Student Representative: J. GIBSON

Department of Biology, Carleton University a

1125 Colonel By Drive, Ottawa, ON K1S 5B6 ‘

jgibsonS5@connect.carleton.ca ™

Website: B. LYONS 4

Canadian Forest Service, Great Lakes Forestry Centre,

Natural Resources Canada, 1219 Queen St East, >

Sault Ste. Marie, ON P6A 2E5 ee

Barry.Lyons@NRCan-RNCan.ge.ca a

JESO Editor: M. RICHARDS ‘a

Dept. of Biological Sciences, Brock University

St. Catharines, ON L2S 3A1

miriam.richards@brocku.ca

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Dept. of Biological Sciences, Brock University .

St. Catharines, ON L2S 3A1 > e

sandra.rehan@brocku.ca a

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Agriculture and Agri-Food Canada

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N. CARTER

154 Riverview Blvd.

R. HARMSEN |

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Kingston, ON N7L 3N6 f

Y. MAUFFETTE .,

Faculté des sciences, Département des sciences biologiques 4

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

J. SKEVINGTON a

Agriculture and Agri-Food Canada :

Eastern Cereal and Oilseed Research Centre Aa

960 Carling Ave., Ottawa, ON K1A 0C6

FELLOWS OF THE ENTOMOLOGICAL SOCIETY OF ONTARIO A

W. W. BILL JUDD (2002) a

_C. RON HARRIS (2003) ‘a

GLENN WIGGINS (2006) aa

JESO Volume 140, 2009

MCZ

JOURNAL LIBRARY

of the DEC 14 2909

ENTOMOLOGICAL SOCIETY OF ONTARIO

HARVARD

UNIVERSITY

VOLUME 140 2009

With this volume, I am very pleased to report that the Journal of the Entomological

Society has more or less completed its transformation to electronic publication. Moving

JESO to this point has been my major objective as Editor, and it is very satisfying to

have achieved this goal. Perhaps you are already aware that pre-print PDF versions of

manuscripts are now being posted on the JESO website soon after final revisions are

accepted. We intend to keep publishing paper copies of the Journal once per year, and so

for the foreseeable future, this is the last change we are planning to make to our publication

process. Electronic pre-publication significantly speeds up the process of disseminating our

contributors’ research, which is one major benefit of this next step.

Another significant step forward is underway with updating of the JESO website.

For the past year or so, I have periodically tested Google’s ability to find published

manuscripts on the JESO website. The results were disappointing — until now. Recently,

JESO’s profile was updated in a web-based database of scientific journals, and an almost

immediate consequence was that JESO is now much more visible to electronic search

engines such as Google. In other words, not only are JESO manuscripts now quickly

available for dissemination, but they should become steadily more visible to entomologists

around the world.

As always, the research presented in Volume 140 (2009) covers a range of

entomological topics and methodologies. JESO is a particularly appropriate journal for

publication of new faunal records, and this volume continues that tradition. A theme of

several papers in this volume is predator-prey interactions, the prey comprising plants or

animals. It is interesting how much entomology focuses on the causes and consequences of

insect dining, for insects, for the natural world, and of course, for ourselves. This volume

also includes a special reviewed essay on the shared history of the ESO and the ESC, one

of a series of essays that we hope to publish in the several years leading up to the 150th

anniversary of the society.

Happy reading!

Miriam H. Richards

Editor

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Forest tent caterpillar outbreaks in east-central Canada JESO Volume 140, 2009

ON THE DURATION AND DISTRIBUTION OF FOREST TENT

CATERPILLAR OUTBREAKS IN EAST-CENTRAL CANADA

B. J. COOKE',", F. LORENZETTP, J. ROLAND?

Natural Resources Canada, Canadian Forest Service, Northern Forestry Centre,

5320 - 122nd Street, Edmonton, Alberta, Canada T6H 385

email: bcooke@nrcan.ge.ca

Abstract J. ent. Soc. Ont. 140: 3-18

‘An analysis of forest tent caterpillar (Malacosoma disstria Hbn.) defoliation

records from Ontario and Quebec indicates that outbreaks recur periodically

and somewhat synchronously (7 = 0.51) in the two provinces, with six inter-

provincial-scale cycles having been observed over the period 1938-2002.

When the entire spatiotemporal range of observed defoliation is considered, it

appears that, at the local stand level, individual outbreaks tend to last for less

than a year on average. Within the three core areas where all six cycles were

observed (Dryden, Sudbury, Temiscamingue), individual outbreaks tended

to last for 2.6 + 0.5 years. The seemingly small difference between two

versus three years of detectable defoliation at the local stand level appears

to be critical, as this determines whether annual rates of stem mortality are

sufficient to produce obvious signs of forest decline. Infestations lasting three

years or longer normally occur in ~45% of the stands within the relatively

small core outbreak areas. However not all infestations behave “normally”,

in the sense of being the product of a regionally synchronized population

cycle. For example, we show how a reversing, traveling wave of forest tent

caterpillar outbreaks in northern Ontario in the 1990s generated an unusually

long-lasting infestation along the Highway 11 corridor — an outbreak which

resulted in a regional-scale decline of trembling aspen. This demonstrates

how incomplete synchronization of forest insect population cycles can

lead to overlapping waves of outbreak that may result in large-scale forest

disturbance.

Published November 2009

' Author to whom all correspondence should be addressed.

Université du Québec en Outaouais, Département d’ informatique et d’ingéniérie, Gatineau,

QC and Institut Québecois d’ Aménagement de la Forét Feuillue, Ripon, QC.

3 Department of Biological Sciences, University of Alberta, Edmonton, AB.

Cooke et al. JESO Volume 140, 2009

‘Introduction

The forest tent caterpillar, Malacosoma disstria Hbn., is a voracious defoliator of

hardwood trees throughout North America, exhibiting large-scale, periodic outbreaks on

trembling aspen, Populus tremuloides Michx., in much of the boreal forest (Witter 1979).

During a typical outbreak, detectable defoliation persists for one to many years, with the

total length of outbreak varying both spatially, within an outbreak, and temporally, among

outbreaks. Different authorities, reporting from different areas and over different time

periods, have provided different estimates of the average duration of outbreak (Table 1);

however, it is not clear why these estimates vary.

Outbreak duration undoubtedly varies among outbreaks and among jurisdictions.

However estimates also vary depending on the way the subjective term ‘outbreak’ is defined.

Sippell (1962), for example, pointed out that although the province-wide outbreak of 1948-

56 in Ontario spanned “a period of nine years”, infestations within “individual stands”

tended to exhibit only “one or two years” of “population excess”. Because local infestations

do not all occur at exactly same time among stands across the province, “infestations” (i.e.

local-scale outbreaks), by definition, do not last as long.as landscape-scale “outbreaks”. At

the limit, when infestation occurrence is highly asynchronous, it becomes impossible to

discern individual outbreaks — a situation which caused Hildahl and Reeks (1960) to reject

the idea of forest tent caterpillar population cycling in west-central Canada.

The purpose of this paper is to provide a transparent and statistically robust answer

to the question: “how long do forest tent caterpillar outbreaks tend to last?” — a question that

is asked by thousands of communities each decade across the country. For example, this is

the question currently being asked in Georgetown, P.E.I., where, after two consecutive years

of heavy defoliation, local residents and authorities are seeking a precise answer as to the

expected termination date, along with some idea of the degree of uncertainty surrounding

this estimate.

To the individual on the ground who has already witnessed a year or two of severe

defoliation, there is a major difference between an expected duration of “one or two years” of

outbreak versus “three or more years”. The variability and lack of specified precision in the

estimates in Table | is therefore disconcerting. The tendency in the literature to characterize

insect disturbance regimes in terms of their long-term, regional-scale behaviour — though

understandable from a population dynamics perspective — is not particularly helpful to the

individual landowner or stand-level forester facing the “here and now” of an outbreak crisis

situation. Forest tent caterpillars are capable of bringing about the decline of trembling

aspen trees and stands over large areas (Churchill et al. 1964, Candau et al. 2002, Hogg

et al. 2002). So the penultimate question of interest to all parties concerned is how long

outbreaks tend to last at the level of individual trees and stands.

Of particular concern is the fact that the outbreak duration estimates in Table |

are all higher than the figures calculated by Simpson and Coy (1999), who summarized

defoliation records in the various forest regions of Canada over the relatively short time

frame 1980-1996 (Fig. 1). Their analysis suggested that the three major forest regions were

quite similar, in that 95% of all infestations last for three years or less — a result that seems

to be at odds with the much longer estimates suggested in Table 1. Is this discrepancy

just a function of Sippell’s (1962) stand vs. landscape scaling issue? Or is it because of a

Forest tent caterpillar outbreaks in east-central Canada JESO Volume 140, 2009

mismatch in the time scales of observation? Clearly, the issue of outbreak duration is one

that needs to be addressed using quantitative, scale-sensitive methods if these important

discrepancies are to be resolved.

TABLE 1. Outbreak duration, estimated in a variety of ways across a range of jurisdictions,

according to several authors. In some cases detailed estimation methods are given in the

original source. In others the estimate is based on informed opinion.

Authority _ Jurisdiction Duration Type of

(yrs) estimate

Cerezke & Volney 1995 Prairie provinces 3-6 qualitative

Witter 1979 Minnesota, USA 3-4 qualitative

Sippell 1962 Ontario 3-9 semi-quantitative

Roland 1993 eight districts in Ontario 1.7-3.3 quantitative

0.600

B boreal plains

@ boreal shield

D atlantic maritime

O PEl

0.500

0.400

0.300

0.200

that was defoliated for given # years

0.100

Proportion of total area defoliated 1980-1996

0.000

1 2 3 4 5 6

Cumulative number of years of defoliation

FIGURE 1. Outbreak duration in three major forest regions of Canada, according to

Simpson and Coy’s (1999) Table 4.

Cooke et al. JESO Volume 140, 2009

Materials and Methods

We analysed the duration of forest tent caterpillar outbreaks over the seven decades

for which Canadian Forest Insect and Disease Survey records exist for the provinces of

Ontario and Quebec, an area which corresponds roughly to the “boreal shield” region reported

on by Simpson and Coy (1999). The source data, described by Fleming et al. (2000) for

Ontario and Cooke and Lorenzetti (2006) for Quebec, consist of digitally rasterized aerial

survey sketch maps of areas exhibiting moderate to severe defoliation attributable to forest

tent caterpillar. Spanning 65 years (1938-2002) and two of the country’s largest provinces,

this is the largest-scale study to date of long-term tent caterpillar outbreak dynamics.

Since 1938 there have been six distinct inter-provincial-scale outbreak cycles in

east-central Canada, with moderate to severe defoliation occurring at periodic intervals

of 9-13 years (Cooke and Lorenzetti 2006, Cooke et al. 2007). For each outbreak cycle,

the number of consecutive years of moderate to severe defoliation at a given “point” was

summed, and plotted in a histogram. This variable is henceforth referred to as “local-scale

outbreak duration”, and is intended to represent the average duration of outbreaks at the

“stand” level. In actuality these “points” were cells in a data raster, each cell spanning |

km? in Ontario and ~58 km? in Quebec, the coarser resolution of the Quebec data being a

function of the way these defoliation maps were rasterized by the province at a resolution of

15 minutes of latitude and longitude.

Results

Forest tent caterpillar outbreaks in Ontario and Quebec tend to exhibit similar

periodic patterns of occurrence (rv = 0.51 between provincial time-series), with the extent

of annual defoliation being more variable in Quebec (C.V. = 216%) than in Ontario (C.V.

= 139%) (Fig. 2, top). In both provinces there are a few core locations where defoliation is

much more frequent than in surrounding areas (Fig. 2, bottom).

A map of local-scale outbreak duration during each of the six inter-provincial

outbreak cycles reveals that the number of consecutive years of defoliation is highly

spatially variable, lasting anywhere from 0 to 9 years depending on location (Fig. 3). A

duration of “zero years” may seem paradoxical. However this is a natural result of the fact

that individual outbreaks in Ontario and Quebec tend to span only 43 + 7% (s.e.) and 37

+ 13% (s.e.) of the insect’s total (i.e. 1938-2002) outbreak range (Fig. 3). In other words,

during a typical 12 year long population/outbreak cycle, 60% of the stands located within

the area amenable to outbreak, for some reason, will not experience moderate-to-severe

defoliation. It is in this sense that a regionally defined outbreak event can be said to have a

duration of zero years in some locales.

How, then, to characterize the distribution of the number of years of defoliation at

a given location during a typical outbreak cycle? In particular, should the zero values from

non-defoliated areas be included in the analysis, or should they be excluded, as in Simpson

and Coy’s (1999) analysis (e.g. Fig. 1)? Excluding them would clearly bias the outbreak

duration estimate upward.

Forest tent caterpillar outbreaks in east-central Canada JESO Volume 140, 2009

Ill IV

--<--- Ontario

—e— Quebec

Area defoliated (1000 sq. km)

Pes wie mieterce

---0”"

ia dow aaa

0 = eoses aan

1935 1955 1965 - +1975

FTC 1938-2002

years of defoliation

1 14 27

90° 85° 80° 75° 70°

FIGURE 2. The distribution of forest tent caterpillar defoliation during six outbreak cycles

in the provinces of Ontario and Quebec. Top: Outbreak cycles are fairly well synchronized

between provinces, although cycles III, IV, and V appear to have been interrupted in the

early stages of development in 1963, 1976, 1989 in Quebec, but not in Ontario. Bottom:

Note the fairly seamless gradient across the Ontario-Quebec border, despite the different

survey and data pre-processing methods. Road density (shown as dark lines) is broadly

indicative of the degree of human settlement and forest fragmentation. Rectangle indicates

area plotted in Fig. 6.

Cooke et al. ; JESO Volume 140, 2009

A second issue is spatial heterogeneity in outbreak frequency. Noting that

defoliation in the Fig. 3 maps is most frequent in rural areas characterized by disturbed,

semi-agricultural landscapes (Roland 1993), it would clearly be advantageous to distinguish

between core areas where outbreaks are frequent versus fringe areas where outbreaks are

infrequent.

The frequency distribution of the number of years that a given cell is defoliated

during an outbreak cycle reveals that this variable is not unimodally distributed (Fig. 4).

The number of zero values in these distributions is high, as expected for a random (i.e.

Poisson) process with a low mean; however the spatial distribution of defoliation is clearly

non-random, following a spatially autocorrelated gradient pattern (Fig. 3). Indeed, the

CY *.

be — ) n. y ‘ AF . Ke : S ly

7 2 , 7 <

years of defoliation | dey aii : years of defoliation | "y. :

ll: 1947-1958 oe ees V: 1984-1994 |< — ,

1, & # peers (oe + 4 88. Fo)

years of defoliation

Ill: 1959-1971

—— a a

FIGURE 3. The distribution of forest tent caterpillar defoliation during each of six outbreak

cycles in Ontario and Quebec. Thin and thick black outlines indicate (i) the entire outbreak

range over the period 1938-2002 and (ii) the core areas where at least one year of defoliation

occurred during each of all six cycles. Core areas labelled as “D” (Dryden), “S” (Sudbury),

and “T” (Temiscamingue). The area between the thin and thick black outlines is referred to

as the “fringe” area — the area where “zero values” for local outbreak duration are common.

Percentages indicate the mean percentage of the outbreak range defoliated in each province

during each cycle. Histograms of outbreak duration provided in Fig. 4.

Forest tent caterpillar outbreaks in east-central Canada JESO Volume 140, 2009

25000

Ontario FTC 1938-2002 Ontario core areas only

- Ml: 1938-1944 20000

x Mill: 1947-1958

cs Billi: 1960-1971

7) 15000

5 WIV: 1972-1983

8 BV: 1984-1994

rz OVI: 1995-2002 10000

Ss O I-VI average

5000

a 0 Tie =;

0 1 2 3 4 5 6 7 8 9 1 2 3 4 5 6 7 8

number of years a 1 sq. km cell is defoliated

Quebec core area only

77)

—

ro)

it | : 5 || D

0 1 2 3 4 5 6 7 8 9 1 2 3 4 5 6 7 8

Number of years a 58 sq. km cell is defoliated Number of years a 58 sq. km cell is defoliated

FIGURE 4. Frequency distribution of the number of years a given cell is defoliated during

each of six outbreak cycles in Ontario (a,b) and Quebec (c,d), both across each province

(a,c), and in “core” areas only (b,d), as defined in Fig. 3. The core area in northwestern

Quebec is smaller than the core area in northwestern Ontario, and outbreak duration is much

more variable among cycles.

TABLE 2. Average number of years of FTC defoliation expected during a “typical”

outbreak cycle in a given cell in east-central Canada (based on n=6 cycles, 1938-2002;

data in Fig. 3). Condition “d > 3” symbolizes the area experiencing three or more years of

defoliation during a 12y outbreak cycle. Its relevance will become clear in Fig. 4. Note

that these estimates include “zero values” — cells which were not defoliated during the

(regionally defined) outbreak cycle. Also note that the core areas are areas which, by

definition, did not exhibit any zero values during any outbreak cycle.

Entire outbreak range Core area only

meants.e. range %aread>3 mean+s.e. range %aread>3

Ontario 0.92+0.11 0-9 ia 2.58 + 0.47 1-8 45.6

Quebec 0.404 2.20 0-9 10.4 2.70 + 0.61 1-6 45.5

Cooke et al. | JESO Volume 140, 2009

bimodality in many of the Fig. 4 distributions suggests a composite distribution resulting

from non-stationarity in the spatial distribution of outbreak.

Summary statistics of the mean and range of local-scale outbreak durations for

the two provinces are provided in Table 2. Mean outbreak duration is comparable in

the two provinces, although Quebec appears to offer a more variable environment, with

outbreak duration exhibiting twice the variance as in Ontario. This is because the fringe

area is estimated to comprise a much larger portion of the insect’s range in Quebec than in

Ontario. The higher variability in outbreak duration in Quebec appears to be exacerbated

by the unusually large extent of defoliation during cycle Il. Excluding cycle II from the

calculation, the average extent of outbreaks relative to the total area amenable to outbreak

would be 39 + 6% (s.e.) in Ontario and 24 + 6% (s.e.) in Quebec.

In the three “core” areas of northwestern Ontario (near Dryden), northeastern

Ontario (near Sudbury), and northwestern Quebec (near L. Temiscamingue) outbreaks

tend to last for 2.6 + 0.6 (s.e.) years. These are rural, populated areas where forest tent

caterpillars are highly likely to encounter humans. In the “fringe” areas, which are more

conifer-dominated, more remote, and are dominated by forest industry activity, outbreaks

tend to last for only 0.8 + 0.1 (s.e.) years in Ontario, and less than this in Quebec. Reporting

bias may therefore help to explain why the literature tends to overestimate the duration

of outbreaks at something greater than two years: in conifer-dominated boreal landscapes

there are fewer observers making fewer reports to fewer readers.

A key question is the probability that a given outbreak will persist for three years

or longer. In both provinces, infestations lasting three years or longer will occur in ~11% of

the outbreak range. Within core areas where populations oscillate with regular periodicity,

this figure jumps to ~45% — still, less than half.

Discussion

1. Duration of Outbreaks

Despite the large extent of forest tent caterpillar outbreaks in east-central Canada,

60% of the area theoretically available for defoliation does not actually experience any

significant defoliation during a typical 12-year outbreak cycle. For the purposes of

computing an average outbreak duration, it matters a great deal whether one chooses to

include these “zero values” in the computation. In the “core” areas where all n=6 outbreak

cycles occurred this is a moot point because there are no such zero values. Beyond the

“fringe” area there are nothing but zero values. If tent caterpillars can be found there, their

populations never reach the level of causing aerially detectable defoliation. It is thus within

the transition region of the fringe area that this question becomes relevant.

The quantitative estimate of outbreak duration by Roland (1993) in Table |

included some of these zero values, in the sense that “if a specific township suffered no

defoliation during an outbreak, this was included in the estimate of mean outbreak duration”.

However not all zero values were included because “populations were considered to be in

the outbreak phase if there was moderate to severe defoliation recorded in at least one-third

of the township”, which means that the time-frame for summation was defined locally,

not globally. Consequently there were many instances where the lack of defoliation in a

Forest tent caterpillar outbreaks in east-central Canada JESO Volume 140, 2009

district prevented local zero values within a township (~10 x 10 km) from being included

in the district sum, despite the possible presence of significant and extensive defoliation in

neighbouring districts. The qualitative estimates of outbreak duration provided in Table |

probably tacitly exclude such zero values. If that is the case, it may help explain why these

estimates appear to be biased high.

Given the contrasting data in Tables 1 and 2, we surmise that the estimates presented

in Table | are descriptions of the dynamics of outbreaks in core areas where (1) outbreaks

occur more frequently and more regularly, (2) the probability of people encountering mass

aggregations of crawling larvae is highest, (3) forests are more fragmented and the local

infestations that comprise the outbreak are not particularly well-synchronized across the

landscape, and (4) forest entomologists interested in quantifying hardwood timber impacts

were historically most likely to focus their attention.

Given the more objective and comprehensive analysis represented in Table 2, it

would clearly be a distortion to suggest that infestations of three or more years in duration

are in any way normal in east-central Canada. Authors who report an average outbreak

duration of anything greater than three years — as in Table 1 — therefore must be reporting on

the basis of individual infestations summed across a larger regional extent, which harkens

back to Sippell’s (1962) original comments on the relatively short duration of local-scale

infestations compared to landscape-scale outbreaks, when individual infestations occur

somewhat asynchronously.

In our case, choosing to focus on local-scale infestation dynamics means that our

estimates of outbreak duration are not only bias-free, they also relate more closely to (1)

the locally-acting processes that are thought to govern cycling (e.g. parasitism, predation,

starvation, host-plant effects, disease) and (11) the critical outcomes of concern (e.g.

probability of permanent tree damage). Our estimates are thus useful to both the small

private landowner and the large forest company.

Finally, the estimates reported here may well turn out to fit other regions, such

as west-central Canada and the Atlantic maritime region, because they correspond well

with the larger-scale, shorter-term estimates reported by Simpson and Coy (1999) in Fig.

1. Had we focused on landscape-level outbreak duration, this might not be the case, for

it is well established that forest tent caterpillar outbreaks are less well synchronized in the

prairie provinces (Hildahl and Reeks 1960) than in Ontario (Sippell 1962). By focusing

on the duration of local-scale infestations, we effectively avoid the issue of the degree of

synchrony among infestations within the area (and time-frame) of outbreak.

2. Forest-Insect Feedbacks

Roland (1993) was the first to attempt a quantitative analysis of the Ontario tent

caterpillar data, and what he showed (using a smaller-scale, abbreviated dataset spanning

cycles II-IV from 1948 to 1984) was that forest tent caterpillar outbreaks in eight major

forest districts tended to last for 2.2 years on average, consistent with what is reported here

for core areas of outbreak. He further showed that there tended to be a split in outbreak

duration, with outbreaks in districts where forests were heavily fragmented lasting “4 to 6

years” and outbreaks in districts where forests were intact lasting only “one or two years”.

A formal analysis indicated that just a single km of edge per square kilometre of forest

area would increase the expected duration of outbreaks from 1.8 years to 2.7 years (see

Cookeret als JESO Volume 140, 2009

his Fig. 2). Consistent with the Ontario data, where there is a strong association between

aspen defoliation and the presence of major roads (Cooke and Roland 2000), we see in Fig.

2 a similar association in the province of Quebec — especially in the northwestern region

around L. Temiscamingue. Moreover, the association between disturbance and prolonged

outbreaks during cycles II-IV (Roland 1993) also appears to be present during cycles I, V,

and VI (Fig. 3). The relationship between forest fragmentation and outbreak duration thus

appears to be quite robust.

From a forestry perspective, the foregoing analysis becomes highly significant

when one considers the result of Churchill et al. (1964), who showed that among dominant,

co-dominant and intermediate (i.e. non-suppressed) trees, mortality due to “an unidentifiable

agent” tended to increase sharply (from 10% to 30%) as the number of years of defoliation by

forest tent caterpillars increased from two to three years of heavy defoliation (Fig. 5). These

authors concluded that the unidentified killing agent must have been the delayed action of

forest tent caterpillar defoliation occurring during the 1950s. Notably, caterpillar-caused

mortality did not happen immediately after the outbreak had started or ended (Duncan and

O suppressed

@ not suppressed

70.0

@ mechanical

0 wind

@ woodborers

OC Nectria

® Hypoxolon

& Fomes

60.0

50.0

% Stem mortality (1955-61)

L HL LHL HHL HHH L HL LHL HHL

Defoliation history Defoliation history

% Stem mortality due to unknown causes (1955-61)

HHH

FIGURE 5. Aspen mortality in Minnesota occurring as a result of the 1951-59 forest tent

caterpillar outbreak cycle (original data in Churchill et al. 1964). ‘L’ indicates a single year

of light defoliation. ‘HHH’ indicates three consecutive years of heavy defoliation. (a) 73%

of all mortality in the ‘HHH’ category is a result of “unknown” causes (i.e. delayed effects

of forest tent caterpillar defoliation). (b) Trees that were “not suppressed” (all dominant, co-

dominant and intermediate trees in the stand) show a clear response to defoliation intensity

over time.

Forest tent caterpillar outbreaks in east-central Canada JESO Volume 140, 2009

Hodson 1958), but occurred gradually, and in association with the growing abundance of

a number of ancillary secondary agents. As time passed, the level of mortality became

increasingly statistically significant and increasingly visually detectable. This is a pattern

that has also been observed in western Canada (Hogg et al. 2002).

Putting the Roland (1993) and Churchill et al. (1964) results together, one may

conclude that a single unit of forest fragmentation (one km forest edge per square km of

forest area) can increase the probability that defoliation will intensify from 1.8 years of

outbreak to 2.7 years of outbreak, which, based on Fig. 5b, would imply a two-fold increase

in mortality among dominant stems, from ~12% to ~30%. In summary, although it is

extremely uncommon for moderate-to-severe defoliation to last as long as 3 years or more

in a given stand, (1) it clearly can happen, (2) forest fragmentation significantly increases

the probability that the three-year threshold is crossed, and (3) the crossing of the three-

year threshold implies significant tree mortality’. From this we conclude that not only are

forest tent caterpillars quite capable of killing their primary host, trembling aspen, but the

probability of heavy mortality increases with forest fragmentation. Notably, this implies

a closed feedback loop between the effect of forest structure on insect dynamics, and the

reciprocal impact of insects on the forest — a relationship that has been confirmed for two

other major Canadian defoliators: the jack pine budworm (Nealis et al. 2003) and the spruce

budworm (Nealis and Régniére 2004).

3. Overlapping Traveling Waves of Outbreak

Candau et al. (2002) suggested that forest tent caterpillars may have been the primary

cause of more than 500 000 hectares of declining aspen along Trans-Canada Highway 11

in northern Ontario — an area where defoliation historically occurs rather frequently (Fig.

2, bottom). These authors showed that outbreak cycles V and VI in this region happened

to occur in very close temporal proximity to one another, with consecutive outbreak peaks

separated by six years, instead of ten years, which is the provincial norm (Fig. 2, top). What

they did not show, however, is that the compression of these cycles in time was associated

with a curious epidemiological phenomenon: a reversing traveling wave of outbreak along

the corridor of Highway 11. The first wave traveled eastward from Hearst to Cochrane

1989-1995, and the second wave traveled westward from Cochrane to Hearst 1996-2004

(Fig. 6, top). Between these two locations, in the zone of overlap at Kapuskasing-Smooth

Rock Falls, trembling aspen host trees, having very little respite from defoliation during the

middle years 1993-1996, were exceptionally vulnerable to sudden dieback and decline (Fig.

6, bottom).

Traveling waves of insect outbreak are of interest to population ecologists because

they are one of the dynamic features predicted by theoreticians to occur in spatially extended

predator-prey systems (Hassell et al. 1994, Bjornstad et al. 2002). However, this particular

traveling wave appears to be different from those that occur in simple theoretical models

in that it reversed direction very suddenly. It is not yet clear why this outbreak progressed

in the unusual way that it did, but this question is being investigated through population

' Note we are not suggesting that 3 years of defoliation is an ecological threshold parameter in a nonlinear mortality

function. On the contrary, we expect the mortality function is a smooth linear function of the degree and duration

of defoliation, and that three years is merely the amount of defoliation required to surpass an arithmetic impact

detectability threshold (unpublished data, D. Marchand, F. Lorenzetti, Y. Mauffette, Y. Bergeron).

Cooke'et al. JESO Volume 140, 2009

provincial outbreak cycle V: 1989-1995 provincial outbreak cycle V1:1996-2004

expansion E. from Hearst translocation E. to Cochrane expansion W. from Cochrane translocation W. to Hearst

a _—_—_—_—_—_—_—_— - a i 7 _ a = —= — a hy =

bOe ac

; a Kaguskasings :

smooth RocksRalls

yrs defoliation 1989-2003

0 5 10

a oe eo

FIGURE 6. Progression of defoliation during outbreak cycles V (1989-1995) and VI

(1996-2004) in northern Ontario. Top: annual displacement of defoliation between years.

Outbreak V, originating at Hearst, expanded and shifted eastward toward Cochrane, while

outbreak VI, originating at Cochrane, expanded and shifted westward toward Hearst.

Bottom: cumulative distribution of defoliation, 1989-2003. Although Hearst and Smooth

Rock Falls both experienced ~9 years of defoliation over the two outbreak cycles, it was at

Smooth Rock Falls where the two population cycles occurred in such rapid succession that

there was little or no respite in defoliation. This is where the highest levels of aspen decline

were observed (Candau et al. 2002).

Forest tent caterpillar outbreaks in east-central Canada JESO Volume 140, 2009

studies and simulation modeling. What we can state, however, is a clear prediction that a

complex dynamic of this type can be expected to be replayed in the future. Meanwhile, it

would be worthwhile trying to determine how much aspen decline might have happened in

response to the overlapping waves of tent caterpillar outbreak that occurred in the boreal

and aspen parkland regions of Alberta in the early and late 1980s, respectively (Cooke

2001).

Finally, this exposé-reveals a demarcation problem in our attempts to quantify

outbreak duration. Recalling that 1994 was the year between cycles V and VI where

province-wide defoliation reached a minimum (Fig. 2), we see now that this was actually

the peak year of defoliation in the out-of-phase regional oscillation at Smooth Rock Falls

(Fig. 6, top). Thus our provincially defined time-frame led to a regional-scale truncation

of the out-of-phase outbreak at Smooth Rock Falls, such that this single regional outbreak

was treated as two separate provincial outbreaks. Outbreak duration in this instance

was therefore underestimated. Estimation error due to imperfect demarcation (deciding

where one outbreak cycle stops and another one starts) is clearly unavoidable when cycle

synchronization is imperfect.

4. Variability in Outbreak Duration, Extent and Timing

Outbreaks appear to be more variable in extent in Quebec than in Ontario, although

this inference is based on a limited sample of only six cycles. Excluding the unusually

extensive outbreak cycle II from the Quebec data, it would appear that the two provinces

exhibit similar levels of variability. However it is not clear that such dismissal is warranted.

Although cycle II was unusually extensive in Quebec, it was also the most extensive

outbreak on record in Ontario. Before discounting cycle II in Quebec as an outlier, it is

important to know if this anomaly might be explained by some persistent feature of the

environment, such as a more variable climate in Quebec.

There does not appear to be any evidence that the range of forest tent caterpillar

outbreaks in east-central Canada is shifting gradually northward in response to a climate

warming trend (Fig. 3). Thus it would be premature to suggest that the decline of aspen in

northern Ontario in the late 1990s was facilitated by climate warming. This system does

not appear to be responding as strongly to climate change as, say, mountain pine beetle

in western Canada (Carroll et al. 2004). On the other hand, given that (1) weather is not

the only driver of the system’s dynamics, and (ii) the 20th century global warming trend

has been punctuated by brief cooling phases (Smith and Reynolds 2005), it may be quite

difficult to estimate the marginal effects of climate change, especially with such a short,

stochastic time-series. Indeed, one of the reasons we have tried to be as quantitative as

possible in estimating outbreak duration is so that future studies looking at this question

will have a solid baseline from which to start. Although tent caterpillar outbreaks may last

as long as 3-6 years in some areas, this is neither precise enough nor accurate enough an

estimate to serve as a baseline for future studies looking at potential shifts in dynamics in

response to climatic and landscape change.

The reason we are keen to continue pursuing this hypothesis is because of regional

differences in outbreak occurrence, with outbreak duration being twice as variable in

Quebec as in Ontario. Looking back at the provincial defoliation time-series of Fig. 2., it

is striking how cycles III, IV, and V appear to have been interrupted in the early stages of

Cooke et al. JESO Volume 140, 2009

development in Quebec, but not in Ontario, hence the asynchronous pattern of outbreak

between the two regions during that time-period. In fact, the years of cycle interruption can

be identified with some precision: 1963, 1976, 1989. It would not surprise us if it should

turn out that these cycles were interrupted by cold spring or winter weather, as described by

Cooke and Roland (2003), for it certainly appears that the insect’s distribution in Quebec

may be strongly limited by a combination of climate and topography (Cooke and Lorenzetti

2006). It is for this reason that we expect climatic change may eventually be found to have

some influence on long-term tent caterpillar dynamics. However additional research on the

relationship between insect survival and weather is required before the hypothesis can be

refined to the point of a specific prediction.

Conclusion

This note is not intended to discount other figures published in the literature,

but merely to put them in context. We want to emphasize that although most forest tent

caterpillar outbreaks do not last longer than 1-2 years, those rare ones that do last longer than

2 years tend to result in “significant” (i.e. readily detectable and/or economically important)

mortality. The reason that forest tent caterpillars are generally thought of as benign insects

is not because they are incapable of destroying a forest. Rather, it is because outbreaks are

typically terminated before they reach their third year. As our analysis indicates, there are

always small areas where outbreaks linger on for 4 years or longer.

Our second major point is that although it is desirable to be able to forecast

population oscillations in time and space, from a forestry perspective it is not particularly

useful to be able to predict cycle timing across the bulk of the outbreak range, when it

is the number of years of defoliation in excess of three that determines whether or not

forests survive. The real challenge lies in predicting precisely when, where and under what

circumstances the number of years of defoliation will exceed the three-year threshold.

Just as meteorologists have difficulty in predicting extreme weather events, so

entomologists are likely to find it challenging to obtain any success in predicting extreme

entomological events. Predicting animal population fluctuations is an imprecise science.

Predicting which of these fluctuations are likely to result in anomalously severe and

prolonged population eruptions is going to require continuing research into the fundamentals

of population dynamics. Understanding the forces that lead to imperfect synchronization

of cyclic population fluctuations is one promising avenue for determining when and where

waves of outbreak may overlap to produce unusually long-lasting infestations capable of

causing large-scale forest decline.

Acknowledgements

Ronald Fournier (Canadian Forest Service) and Bruno Boulet (Ministere des

Ressources Naturelles et Faune du Québec) kindly provided access to forest tent caterpillar

defoliation data from Ontario and Quebec, respectively.

Forest tent caterpillar outbreaks in east-central Canada JESO Volume 140, 2009

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Pipunculidae attacking proconiine sharpshooters JESO Volume 140, 2009

NEW RECORDS OF PIPUNCULIDAE ATTACKING PROCONIINE

SHARPSHOOTERS (AUCHENORRHYNCHA: CICADELLIDAE:

PROCONIIND)

J. H. SKEVINGTON', J. A. GOOLSBY?

Agriculture and Agri-Food Canada, Canadian National Collection of Insects, Arachnids

and Nematodes, 960 Carling Avenue, K.W. Neatby Building, Ottawa, ON, K1A 0C6

email: jeffrey.skevington@agr.gc.ca

Abstract . J. ent. Soc. Ont. 140: 19-26

Five records of Pipunculidae (Diptera) attacking proconiine sharpshooters

(Auchenorrhyncha: .Cicadellidae) are documented here for the first time.

Eudorylas alternatus (Cresson) is documented as a parasitoid of Cuerna

obtusa Oman and Beamer and Oncometopia orbona (Fabricius) is recorded

as being attacked by an apparently-undescribed species of Eudorylas

(Pipunculidae). Records of unidentified pipunculid larvae are also recorded

from Cuerna kaloostiani Nielson, Cuerna curvata Oman & Beamer, and

Cuerna sp. near striata (Walker) — septentrionalis (Walker). We describe

these observations, summarize the data for them and explore the potential of

Pipunculidae as biological control agents for pest proconiines such as glassy-

winged sharpshooter (Homalodisca vitripennis (Germar)). We also reveal the

utility of DNA barcoding for identifying pipunculid larvae.

Published November 2009

Introduction

With the exception of the big-headed fly genus Nephrocerus Zetterstedt which

attack crane fly adults (Tipulidae), pipunculids are parasitoids of leafhoppers and

planthoppers (Hemiptera: Auchenorrhyncha). They typically attack second instar larvae,

although some parasitize adults (Waloff and Jervis 1987). Big-headed flies are found in

almost every terrestrial habitat world-wide including agricultural ecosystems. Their larvae

develop fully within their host, typically emerging from the dorsum of the abdomen of adult

hosts after a rapid development. Hosts are usually rendered sterile or are killed by these

parasitoids. Larvae normally pupariate in the leaf litter or soil. Development is variable with

multivoltine species typically eclosing from the puparium within a few days to weeks and

univoltine species overwintering in the substrate (Waloff 1980; Skevington and Marshall

' Author to whom all correspondence should be addressed.

? United States Department of Agriculture, Agricultural Research Service, Kika de la Garza

Subtropical Agricultural Research Center, Beneficial Insects Research Unit, Weslaco, TX,

USA

Skevington and Goolsby JESO Volume 140, 2009

1997). The effects of pipunculid parasitization on planthoppers and leafhoppers have been

documented by numerous scientists, most recently by May (1979), Chandra (1980), Waloff

(1980), Lauterer (1981), Hug (1984, 1986a, 1986b), Ylonen and Raatikainen (1984), Yano

(1985), and Skevington and Marshall (1997). Parasitized hosts are sometimes recognizable

by their swollen abdomen and sluggish movements.

Recorded rates of parasitism vary from fractions of a percent to nearly 100 percent

in local populations. For example, Hartung and Severin (1915) found Circulifer tenellus

(Baker) (beet leafhopper, Cicadellidae) with up to 47% parasitism by two pipunculid

species and Skevington and Marshall (1997) recorded parasitism rates of Cuerna striata by

Eudorylas sp. near alternatus to be as high as 89%. Despite the importance of pipunculids

as parasitoids, few rearing records exist for Pipunculidae, particularly in North America

(Skevington and Marshall 1997). Data on host ranges are available for more than 52

European species of Pipunculidae (Skevington and Marshall 1997) while in the Nearctic

Region only 16 species have received such documentation (Skevington and Marshall 1997;

Moya-Raygoza et al. 2004; Koenig and Young 2007).

The potential value of Pipunculidae for biological control has stimulated some

work on the bionomics of this family. For example, research into the control of the potato

leafhopper, Empoasca fabae (Harris), a major pest of alfalfa in mid-western and eastern

USA and Canada, involved exploration within Europe for natural enemies to be introduced

to the United States (Jervis 1992). Chalarus specimens were reared for this effort but

apparently were never released. Similarly, European species of Chalarus were considered

for introduction into New Zealand for control of Frogatt’s apple leafhopper, Edwardsiana

crataegi (Douglas), populations of which are insecticide resistant (Jervis 1992). A release

was never made because of concerns about adding yet another foreign species to the New

Zealand fauna (pers. comm. M. De Meyer).

We decided to investigate the potential of these flies as parasitoids of Glassy-winged

Sharpshooter (GWSS, Homalodisca vitripennis (Germar) (Cicadellidae, Proconiini)) in

2005. This species is native to the southeastern USA and northeastern Mexico, from Augusta,

Georgia to Leesburg, Florida, west to ValVerde and Edwards counties in Texas, south to

Mexico (Turner and Pollard 1959; Triapitsyn and Phillips 2000). It has become a serious pest

of grapes in California where it was introduced in 1989 (Sorensen and Gill 1996; Hoddle

2004). Glassy-winged sharpshooters are effective vectors of Xylella fastidiosa Wells et al.

(Eubacteria), the causative agent of Pierce’s Disease in grapes, which has severely damaged

vineyards in southern and central California (Hoddle 2004). Considerable effort has been

expended to find egg parasitoids of GWSS and other pest leafhoppers, but little effort to

date has been made to study their nymphal parasitoids (Goolsby and Setamou 2005; Irwin

and Hoddle 2005; Pilkington et al. 2005). Finding a larval parasitoid for GWSS would be

a great advance in potential biological control programs for the species. Although we have

not discovered such a parasitoid, the discovery of several pipunculid parasitoids (described

below) attacking related proconiine species is encouraging.

Pipunculidae attacking proconiine sharpshooters JESO Volume 140, 2009

Methods and Materials

Adult pipunculids and leafhoppers were either killed with cyanide and pinned or

collected into 100% alcohol. Specimens are deposited in the Canadian National Collection

of Insects, Arachnids and Nematodes (CNC) and the Illinois Natural History Survey

Collection (INHS). The CNC specimens are all labelled with a unique number (either in

the format JSS # n or CNCD # n). Pipunculid larvae were collected into 70% alcohol (RR)

or 100% alcohol (JHS). Voucher data for the material used in this study are available in

Appendix 1. — : :

Field work contributing to this study was conducted by two teams. Roman

Rakitov collected the Arizona specimens while conducting general fieldwork there in

2003. John Goolsby coordinated fieldwork in Texas where his team was searching for

potential biological control candidates for GWSS. When possible, leafhoppers were killed

and dissected in the lab to search for parasitoids. When no lab facilities were available,

leafhoppers were examined in the field for evidence of parasitism. Although leafhoppers

that are parasitized by third instar pipunculids may be recognized in the field by their

sluggish behaviour and swollen abdomens, we found no behavioural changes in cicadellids

parasitized by first instar larvae. Dissection of a random series of leafhoppers in the field

(by removing their abdomens and squeezing out the contents) thus allowed discovery of

parasitized populations of leafhoppers. Even though very small, first instar pipunculids are

easy to see as they crawl around.

Pipunculid larvae and adults collected in the survey were sequenced in an effort to

match the identity of the immatures with the adult specimens. DNA was extracted and a 658

base pair fragment of the COI gene (now referred to as cox1 in the ‘barcoding’ literature) was

amplified using the primer pair LCO1490 (5’-GGTCAACAAATCATAAAGATATTGG-3 ’)

and HCO2198 (5’-TAAACTTCAGGGTGACCAAAAAATCA-3’) (Folmer et al. 1994).

Methods used follow Hebert et al. (2003). Relevant sequences were deposited in GenBank

(Appendix 1).

Parsimony and neighbour-joining analyses were performed with PAUP* (Swofford

2001). Chalarus sp. was defined as the outgroup for all analyses, as this is the putative

basal genus of Pipunculidae (Rafael and De Meyer 1996; Skevington and Yeates 2000). The

heuristic search procedure was used with stepwise-addition and 100 random replications.

The heuristic search option was used with tree bisection-reconnection branch swapping,

MULPARS, and random addition of taxa. Multistate characters were treated as non-

additive.

Skevington and Goolsby JESO Volume 140, 2009

Results and Discussion

Arizona

Between 13 and 18 April 2003, 33 Eudorylas alternatus puparia were obtained by R.

Rakitov from pipunculid larvae developing within Cuerna obtusa in Arizona (Appendix 1).

From these puparia, 19 adult pipunculids (10 females, 9 males) were reared. The leafhoppers

were collected in forests of Pinus edulis and P. ponderosa. Note that the identification of

these flies is tentative, despite being based on examination of the FE. a/ternatus holotype.

Confirmation will only be possible in the context of a complete revision of Eudorylas. The

best current key to Nearctic eudorylines (Hardy 1943) does not work and over half of the

species in the genus are undescribed (Skevington unpublished data). These flies appear to

be conspecific with the flies reared from Cuerna striata in Ontario, Canada (Skevington and

Marshall 1997). Although there is minor genitalic variation, their coxl sequences differ by

only 0.5%. This is typical of genetic distances among species of Pipunculidae (Skevington

et al. 2007).

Rakitov (personal communication) also reports records of pipunculized specimens

of Cuerna kaloostiani from Arizona, Cuerna curvata from California, and Cuerna sp. near

striata — septentrionalis from Utah. The parasitized cicadellids and extracted pipunculid

larvae supporting these records are in the INHS collection. These pipunculids are likely also

species of Eudorylini, but further research is needed to corroborate this hypothesis.

Texas

On 20 October 2005, we dissected two first instar pipunculid larvae out of adult

Oncometopia orbona at Yegua Creek, Texas (from ten O. orbona that were dissected). All

efforts to rear this species of pipunculid from additional leafhoppers failed. Larval pipunculids

are unidentifiable to species and in most cases, even to genus. In an effort to identify the

larvae, we extracted DNA from one specimen and sequenced cox1. The generic identity

of this larva was hypothesized based on phylogenetic placement of this sequence within a

large matrix being prepared for a paper on the phylogeny of Pipuncultdae (Skevington et al .

unpublished data). Parsimony analysis using this dataset supported the placement of the larva

as a member of the genus Eudorylas (the closest relative, E. alternatus, was 14.2% different

based on pairwise analysis). This generic identification was expected, given that the other

two identified pipunculids recorded as attacking proconiines were species of Eudorylas.

Based on this discovery, we added 54 morphospecies of Eudorylini from the southern USA

to the cox! dataset and found a match (specimen CNCD3333) — the uncorrected pairwise

distance between the two specimens is 0.6%, within the range of typical intraspecific genetic

distances for pipunculids (Skevington et al. 2007). Assigning a name to this fly continues

to be a problem. It cannot be identified with existing keys and will only be named in the

context of a planned revision of the Eudorylini (Skevington, in prep). What we have learned

though is where this species is likely to occur. Comparing CNCD3333 with other female

pipunculids in the Canadian National Collection of Insects and the United States National

Museum collection, turned up five specimens of this species (listed as Eudorylas sp. TX8 in

Appendix 1). As a result, we now know that this species occurs from College Station and

Yegua Creek, Texas (Houston area) to Greenville, Mississippi, and appears to be at least

bivoltine. Flight times are from April to May and September.

Pipunculidae attacking proconiine sharpshooters JESO Volume 140, 2009

This example illustrates the power of DNA barcoding to associate immature stages

with adults. It also illustrates how important it is to continue to work towards modern

revisions of these flies. One of us (JHS) has been routinely DNA barcoding all of the

species that he includes in revisions for five years (Skevington 2005b; Skevington 2006;

Skevington and Féldvari 2007; Skevington and Kehlmaier 2008), but a concerted effort is

clearly needed to barcode as many species of adult pipunculids as possible. Doing so will

open up research on biological control and facilitate ecological studies of these important

flies. |

Given the oligophagous nature of most pipunculids, we speculate that the species

attacking O. orbona will also be found in H. vitripennis as both of these proconiines occur

in the same habitats at the same time of year. Further research is warranted to collect,

rear and evaluate this species of pipunculid as a potential biological control agent of H.

vitripennis where it is invasive in California. Revision of Nearctic Eudorylini is also clearly

a priority. It is likely that over 200 species occur in the Nearctic Region and only 38 valid

species are currently described (Skevington 2005a). Most of these are not identifiable using

current resources.

Acknowledgments

Thanks to Roman Rakitov (Illinois Natural History Museum) for providing the

host and parasitoid data for the southwestern Cuerna species, identifying species of Cuerna

and Oncometopia, and commenting on the manuscript. This work was supported by funding

from Agriculture and Agri-Food Canada and the United States Department of Agriculture,

Agricultural Research Service. Assistance with the DNA barcode analysis was provided by

J. deWaard and P. Hebert (Canadian Centre for DNA Barcoding).

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Skevington and Goolsby JESO Volume 140, 2009

Appendix 1 — Material Examined (Voucher data)

Pipunculidae: Pipunculinae: Eudorylini: Eudorylas alternatus (Cresson): USA, AZ,

Coconino Co., 2.5 miles S Tusayan, “10X” Campground, 35°56’16.3” N, 112°07°48.7” W,

R. Rakitov, 9, 10), 11 puparia, 3 third instar larvae, collected in Pinus edulis & Pinus

ponderosa forest, host collection date 11.iv.2003, pupation dates 13-18.iv.2003, adult

emergence dates 9-13.v.2003, host Cuerna obtusa Oman and Beamer, JSS# 13848-13849

(CNC), 13850 (INHS), 13851 — 3 legs removed for sequencing — GenBank # DQ349219,

13852-13854 (CNC), 13855 (INHS), 13856-13869, 13871-13881 (CNC).

Eudorylas sp. nr. alternatus (Cresson) Canada, ON, Sideroad 25, 5 km SE Arkell, 1 |, host

collection date 27.iv.1993, pupation dates |.v.1993, adult emergence date 20.v.2003, host

Cuerna striata Walker, JSS#12590 (CNC) — 3 legs removed for sequencing — GenBank #

DQ349219.

Eudorylas sp. TX8: larvae: USA, TX, Lee Co., Yegua Creek, 30°17°28” N, 96°15°39”

W, 82 m, J. Skevington, 20.x.2005, 2 first instar larvae (one per host), host Oncometopia

orbona (Fabricius) adults (one voucher JSS#16947 listed below), JSS#16853, one larva

destroyed for sequencing — GenBank # DQ337627 (CNC); adult females: USA, TX,

Brazos Co., College Station, Lick Creek Park, 30°38’ N, 96°20’ W, 17. Iv. 2006, Malaise

trap, R. A. Wharton, CNCD3333 — GenBank # FJ860147 (CNC); USA, MS, Lafayete Co.,

F. M. Hull, v.1951, CNCD4914, iv.-v.1946, CNCD4914 (CNC); MS, Greenville, 11.ix.1922

(2 specimens), CNCD4916-7 (CNC).

Cicadellidae: Cicadellinae: Proconiini: Oncometopia orbona (Fabricius): USA, TX, Lee

Co., Yegua Creek, 30°17°28” N, 96°15°39” W, 82 m, J. Skevington, 20.x.2005, host of first

instar Eudorylini larva (larva destroyed for sequencing), | adult |, JSS#16947 (CNC).

Population dynamics of Asian lady beetles and aphids JESO Volume 140, 2009

POPULATION DYNAMICS OF HARMONIA AXYRIDIS AND APHIS

GLYCINES IN NIAGARA PENINSULA SOYBEAN FIELDS AND

VINEYARDS

C. A. BAHLAIT’, M. K. SEARS

School of Environmental Sciences, University of Guelph

Guelph, Ontario, Canada NIG 2W1

~ email: cbahlai@uoguelph.ca

Abstract | J. ent. Soc. Ont. 140: 27-39

Multicoloured Asian lady beetle (Harmonia axyridis) is an occasional

pest of wine and juice grapes in vineyards throughout northeastern North

America. In late season, beetles aggregate on grape clusters immediately

before harvest, and are difficult and expensive to remove before processing.

Outbreaks of H. axyridis are thought to be related to soybean aphid (Aphis

glycines) populations. Heavy infestations of aphids occur late in the season

on soybeans and can sustain large numbers of H. axyridis. Each summer

from 2004 to 2006, 23-29 soybean fields along the escarpment of the Niagara

Peninsula were monitored each week for soybean aphid infestation, and all

life stages of H. axyridis were recorded. Where substantial populations of A.

glycines were found, larvae and adults of H. axyridis soon followed. Severity

of H. axyridis infestation in vineyards was still high even when A. glycines

populations were reduced by insecticides in soybean fields in 2005. Outbreaks

of H. axyridis in vineyards are correlated with substantial populations of

soybean aphid that occur early in the season. Outbreak populations of H.

axyridis in vineyards were observed in years where A. glycines eggs were

not abundant on overwintering hosts, thus H. axyridis density appears to be

negatively correlated with numbers of overwintering A. glycines eggs on its

primary host, Rhamnus cathartica. A model of interaction between the two

species is proposed.

Published November 2009

Introduction

Multicoloured Asian lady beetle (Harmonia axyridis (Pallas), Coleoptera:

Coccinellidae) is an alien invasive predator important in southern Ontario agro-ecosystems.

Harmonia axyridis is an occasional pest of wine and juice grapes in vineyards throughout

northeastern North America (Ker and Carter 2004). Like most coccinellids, adults and

larvae of H. axyridis are predacious, with a diet consisting primarily of aphids and other

' Author to whom all correspondence should be addressed.

Bahlai and Sears ; JESO Volume 140, 2009

soft-bodied insects, supplemented by small amounts of plant material (Hodek 1973). In

the Niagara Peninsula region of southern Ontario, late season aggregations of adults have

been observed on ripening grape clusters immediately before harvest (Ker and Carter 2004).

During processing, beetles may be crushed into the slurry of skins and stems (Pickering

2004). Beetles are difficult and expensive to remove from grape clusters before processing,

and if they are not removed before processing the grapes, alkaloids secreted by beetles as

defensive chemicals affect the flavour of wines and juices (Koch 2003, Pickering 2004).

Insects inevitably are present at grape harvest, but usually in low enough numbers

that their presence does not affect quality or flavour of wine. Harmonia axyridis presents

a problem because the beetle itself has a very unpleasant taste and odour, due to a bitter

defensive chemical, 2-isopropyl-3-methoxypyrazine (IPMP), present in its haemolymph

(Pickering 2004). The limit of detection by humans of IPMP in water is in the range of two

parts per trillion (Pickering 2004).

The Niagara Peninsula region of Ontario is an intensely cultivated area home to

94% of Ontario’s grape industry, with an annual farm gate value of $60 million (Gardner

et al. 2006). Approximately 40 million litres of wine are produced in Ontario each year,

generating $438 million in retail sales (Grape Growers of Ontario 2007). This region also

has large areas devoted to field crops located above the escarpment and south of grape

growing areas. Most vineyards have fields of soybeans planted within 1-2 km of their

location (Fig. 1).

Outbreaks of H. axyridis in grapes may be related to infestations of soybean aphid

(Aphis glycines Matsumura). Though H. axyridis has been present since 1994 in southern

Ontario, large populations of H. axyridis were not observed in Ontario vineyards until 2001,

coinciding with the arrival of A. glycines (Ker and Carter 2004). Harmonia axyridis is an

important natural predator of A. g/ycines in its native range (Koch 2003, Wu et al. 2004).

First identified in North America in 2000 in Wisconsin, A. glycines is now a severe pest

of cultivated soybean (Glycine max Merrill) in 21 American states and three Canadian

provinces (Ragsdale et al. 2004). Aphis glycines undergoes a heteroecious, holocyclic life

cycle, alternating between parthenogenic reproduction on its secondary summer host, G.

max, and sexual reproduction and overwintering on a primary woody winter host, buckthorn

(Rhamnus spp.) (Ragsdale et al. 2004, Voegtlin et al. 2004) . Typically, A. glycines occurs

as a sexual morph on foliage of Rhamnus spp. in autumn, as an egg on buds of Rhamnus

spp. in winter, as an asexually reproducing female on Rhamnus spp. in spring, and as

an asexually reproducing female in cultivated soybean in summer (Ragsdale et al. 2004,

Voegtlin et al. 2004).

Ample populations of aphids can support large numbers of H. axyridis (Fox et

al. 2004), and there is anecdotal evidence that in years favouring heavy infestation of A.

glycines, heavy infestations of H. axyridis occur in vineyards. A biennial cyclical pattern

of outbreak years seems to be emerging for both H. axyridis in grapes and A. glycines in

soybeans, in which economically damaging infestations of both species occurred in 2001,

2003, 2005, and 2007, but only spot infestations were observed in 2002, 2004, 2006, and

2008 (Bahlai 2007, Glemser, E. et al., unpub. data)

It is possible that soybean fields near to Niagara vineyards serve as a reservoir for

H. axyridis. Starting near the middle of the growing season, H. axyridis might reproduce in

Population dynamics of Asian lady beetles and aphids JESO Volume 140, 2009

soybean fields, and feed as adults and larvae on aphids. When aphids move to overwintering

sites, beetles seeking alternate food sources in the absence of aphids, might move directly

to nearby vineyards filled with ripening grapes. If this relationship holds true, numbers of

H. axyridis observed in vineyards should correspond to numbers of beetles observed in

soybean fields just before soybean leaf senescence occurs.

From June to September in 2004, 2005 and 2006, we monitored 23-29 soybean

fields weekly for A. g/ycines infestation and for all life stages of H. axyridis in the Regional

Municipality of Niagara, ON. The purpose of this study was threefold:

1) to provide scouting information for Niagara area soybean growers regarding

A. glycines infestation levels, and to estimate numbers of H. axyridis for Ontario grape

growers and vintners,

2) to examine whether population assessments of H. axyridis in soybean fields

correspond with infestations of the beetle in nearby vineyards,

3) to test the hypothesis that soybean fields act as reservoirs for H. axyridis before

beetles infest vineyards.

Methods and Materials

During the growing seasons of 2004-2006, soybean fields were selected along the

edge of the Niagara Escarpment in proximity to vineyards from Grimsby (43.2°N, 79.7°W)

to Niagara-on-the-Lake (43.2°N, 79.1°W )(Fig. 1). Nearby vineyards generally were

located on the “bench” below the escarpment, in the northern portion of the peninsula, 2-8

km from the shore of Lake Ontario. Soybean fields selected were generally located to the

immediate southwest of vineyards, and within a 5 km radius. In 2004, 23 soybean fields

were monitored each week; in 2005 and 2006, 29 and 28 fields were monitored each week,

respectively.

Monitoring consisted of sampling 10 sites randomly selected within a soybean

field. At each site, three soybean plants were assessed for soybean aphid populations using

the following rating scale (after Difonzo and Hines 2002). Aphid populations were assessed

on stems and on the middle leaflets of the lowest, middle and top trifoliate leaves. The

following rating system was applied to each part of the plant: 0 = No aphids present (not

infested), 1 = 1-10 aphids present (low infestation), 2 = 11-25 aphids present (moderate

infestation), 3 = 26-100 aphids present (high infestation), and 4 = 100+ aphids present

(extreme infestation). Ratings for all plant parts were averaged, providing a total infestation

rating out of four for each plant. These ratings were averaged by field and provided an

average infestation score for the fields in a particular area.

For each of the plants assessed, the number of larval, pupal and adult H. axyridis

present on the plant were counted. These numbers were averaged by field and geographical

region, and reported in units of average number of individuals per soybean plant.

Each site was monitored once weekly, commencing on July 13 in 2004, June 22

in 2005, and June 23 in 2006. Monitoring continued until soybean leaf drop occurred

in all observation fields in September. Population data for fields in particular areas were

compared with H. axyridis infestation levels in nearby corresponding vineyards and with

counts of overwintering eggs of A. glycines as described in Welsman et al. (2007).

Bahlai and Sears JESO Volume 140, 2009

Data for the regression analyses were organized by observations in a given week.

Regression analyses were performed on population data using PROC GLM (SAS Institute,

Cary, NC) to determine whether counts of larvae, pupae and adults of H. axyridis would

correlate over time with infestations of A. glycines, or if a one or two week delay in interval

would provide a better statistical relationship. A significance level of |'=0.05 was used for

all analyses.

Results

In 2004, A. glycines populations reached moderate infestation levels in soybean

fields across Niagara in late August (Fig. 2). None of the observation fields had insecticides

applied at this time, because at the time this study was performed treatment was not

recommended for soybean aphid control after the R5 (‘beginning seed’) plant stage is

reached (Baute 2007). Infestation rankings reached an average of 0.25 in the week of

August 12, 2004 (Table 1). Sharp increases in aphid infestation occurred in the two weeks

following August 24, with populations peaking by September 7 in all observation fields.

Numbers of H. axyridis larvae followed a similar growth and peak pattern, with jumps

in their population growth correlating with increases in aphid infestation (R* = 0.88, p <

0.0001)(Table 2, Fig. 2). Abundance of pupae correlated significantly with aphid infestation

after one week (R? = 0.53, p = 0.006), with adult beetles following at two weeks after aphid

infestation increase (R*? = 0.70, p = 0.002) (Table 2, Fig. 2).

In 2005, an average rating of 0.25 was first recorded on July 19, over three weeks

earlier than was observed in 2004 (Table 1). Earlier infestation of soybean fields by A.

glycines and rapid increasing severity of infestation in early summer (Fig. 2) resulted in 28

of 29 observation fields being sprayed with cyhalothrin-lambda (Matador 120E*, Syngenta

Crop Protection Canada) or dimethoate (Cygon 480° and Lagon 480EC*, Cheminova

Canada) (OMAFRA 2005) to control populations in the weeks of August 9 and 16, 2005.

Aphid infestation across the peninsula peaked in these weeks, and subsequently decreased

for the rest of the season in most of the observation fields (Fig. 2). Populations of H.

axyridis began to increase in observation fields early in the season, correlating with aphid

infestation levels (Table 2), but sharply declined after the application of insecticides (Fig.

2). The relationship between aphid infestation and larvae or pupae counts after a delay of

one week in 2005 was weaker than in 2004 (R? = 0.42, p = 0.020 for larvae, R?= 0.44, p =

0.012 for pupae) (Table 2).

In 2006, very low aphid infestation and very few H. axyridis were observed in

Niagara Peninsula soybean fields (Fig. 2). Aphid infestation density did not reach a rating

of 0.25 until August 22, 2006 (Table 1). Counts of larvae of H. axyridis were observed

to correlate well with aphid infestations in soybean fields, but occurred one week and

two weeks after the corresponding aphid population estimate (Table 2). The relationship

between beetle and aphid densities was significant for all three temporal relationships

examined (Table 2).

JESO Volume 140, 2009

Population dynamics of Asian lady beetles and aphids

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Jun Jun Jul Jul Jul Jul Aug Aug Aug Aug Aug Sep Sep Sep Sep

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23- 27- CO 11- 17- 28 O03 10 1& 22- 30 O7- 12- 19 26

Jun Jun Jul Jul Jul Jul Aug Aug Aug Aug Aug Sep Sep Sep Sep

Date

FIGURE 2: Soybean aphid (SBA) infestation rating and multicoloured Asian ladybeetle

(MALB) counts for soybean fields in the Niagara Peninsula region of southern Ontario in

2004, 2005 and 2006. Area-wide averages are shown. A sample of thirty soybean plants

in observations field were monitored weekly. SBA ratings were performed by examining

the stem and upper, middle and lower trifoliates of each plant and rating each portion of

the plant on a scale of 0 to 4, where 0=No aphids present, 1=1-10 aphids present, 2=11-25

aphids present, 3=26-100 aphids present, 4=100+ aphids present. Raw counts of MALB

adults, pupae and larvae were performed for each plant. Note: scales of graphs differ to

preserve detail when average aphid infestation levels and ladybeetle counts are lower in

2005 and 2006.

Population dynamics of Asian lady beetles and aphids JESO Volume 140, 2009

TABLE 1: Soybean aphid (SBA) populations in soybean fields in the Niagara Peninsula

region of southern Ontario, 2004-2006. Dates when an average of one aphid colony per

plant was first observed and when soybean leaf drop occurred, a description of the peak

aphid population, is provided. Counts of aphid eggs subsequently observed in overwintering

sites and infestation levels of multicoloured Asian ladybeetle (MALB) in vineyards for each

year are included.

Total

Date rating Date of SBA eggs

of 0.25 soybean observedon MALB infestation in

Year —_ reached SBA peak leaf drop R. cathartica' vineyards?

2004 12-Aug-04 Moderate, 28-Sep-04 5585 Low: spot

after pod set, infestations (raw

no chemical data not available)

control required

2005 19-Jul-05 Moderate, 21-Sep-05 4 High: widespread

before pod infestation (896 adult

set, chemical MALB observed in

control widely sampling period)

applied

2006 22-Aug-06 Low, afterpod 12-Sep-06 250 Low: spot

set, no chemical infestations (105

control required adult MALB

observed in sampling

period)

' Welsman et al. (2007): soybean aphid eggs collected from 10 cm buckthorn twig segments (N=1200) in autumn

near Guelph, ON.

* Kevin Ker, Ker Crop Management Services, personal communication. Assessments completed by counting

number of MALB observed per meter of grape vine in commercial vineyards.

TABLE 2: Linear regression of observed populations of various life stages of multicoloured

Asian ladybeetle (MALB) on soybean aphid infestation scores in soybean fields in the

Niagara peninsula region of southern Ontario, 2004-2006. Counts of each life stage of

MALB were correlated to soybean aphid infestation observed concurrently, one, and two

weeks before.

MALB life stages

Larvae Pupae Adults

Year Week rR’ R? P R? P

2004 0 0.88 <0.0001 * 0.09 0.340 0.06 0.420

| R56 nce OO0T 3% 0:53 0.006 * 0.30 0.080

2 0.33 0.080 0.91 <0.0001 * 0.70 0.002 *

2005 0 0.18 0.130 0.12 0.230 0.40 0.020

| 0.42 0.020 * 0.44 OTT wie tiny Meo 0h Fate

2 0.19 0.150 0.33 0.050 0.56 0,005, .*

2006 0 0.41 0.200 0.45 0.012 * 084 <0.0001 *

] 0.70 0.001 * 082 <0.0001 * 0.86 <0.0001

ye u98 ~~ '=00001 * 0.85" ~ <0.0001 “* 0.71 aa.

* Significant at (1=0.05

Bahlai and Sears JESO Volume 140, 2009

Discussion

To date, a biennial cycle of outbreak years of both H. axyridis and A. glycines has

consistently occurred in the Niagara (Glemser, E. et al, unpub. data). As was observed in

2001 and 2003, in the present study, high numbers of aphids appeared early in the 2005

growing season, and high lady beetle numbers appeared in vineyards later in the season

(Table 1). In 2004 and 2006, as in 2002, low or moderate late-season soybean aphid

infestations occurred, and only low infestations. of H. axyridis were observed in vineyards

(Table 1). Infestation by H. axyridis in vineyards in a given year did not necessarily

correlate with observed numbers of ladybeetles in soybean fields immediately before leaf

senescence. More H. axyridis individuals in total were observed in soybeans in 2004, when

only spot infestations of the beetle were observed in vineyards, than in 2005, when vineyard

infestation by H. axyridis was reported to be much higher. This provides evidence against

the hypothesis that abundance of ladybeetles in vineyards results entirely from abundance

of A. glycines and that beetles move directly from soybeans to ripening grapes.

The application of insecticides to most of our observation soybean fields in 2005

confounded our results considerably. The insecticides cyhalothrin-lambda (Matador 120E*,

Syngenta Crop Protection Canada), a pyrethroid, and two formulations of dimethoate

(Cygon 480® and Lagon 480EC*, Cheminova Canada), an organophosphate, are registered

for use in controlling soybean aphid in Ontario soybeans (QMAFRA 2005). Pyrethroids

are extremely toxic to larvae of H. axyridis (Youn et al. 2003) and unpublished field trials

suggest they have a repellent effect on adults (K. Ker, personal communication). In leaf-dip

trials, organophosphorous pesticides applied at normal rates resulted in low survivorship of

all life stages of H. axyridis (Youn et al. 2003)

This decline in abundance of H. axyridis in soybean fields observed after insecticide

application may occur for several reasons: 1) the insecticide is toxic to H. axyridis, 2) the

insecticide may act as a repellent to adults of H. axyridis, so that they disperse from the

field and new migrants avoid the field, or 3) the sudden drop in aphid abundance results

in insufficient aphid populations for the induction of oviposition by H. axyridis, so that

beetles disperse to locate other populations of insects on which to feed. A combination of

these explanations likely leads to the observed population decline of H. axyridis. By early

August, when insecticides are applied if needed to soybeans for control of A. glycines in

the Niagara region, we have observed other aphid species supporting feeding populations

of H. axyridis on common weeds such as lamb’s quarters (Chenopodium album L.) or milk

vetch (Vicia cracca L.). If insecticides repel surviving adults of H. axyridis, and there are

not sufficient aphid populations in soybean fields, beetles will move out of soybean and

forage on abundant populations of other aphid species occurring in weedy areas, woodlots

or orchards. This dispersal of H. axyridis from soybean fields confounds monitoring of

ladybeetle population numbers because large numbers of H. axyridis are likely present

outside soybean fields in late summer, and at that time of year, populations of A. glycines

may no longer have as much influence on the population growth of H. axyridis.

The exact relationship between outbreaks of H. axyridis in vineyards and

outbreaks of A. g/ycines in soybeans can only be speculated upon at this time, but the two

may be related. Grape harvest in Ontario usually begins in the last week of September, and

continues until the middle of October, except for vineyards where grapes are destined for

Population dynamics of Asian lady beetles and aphids JESO Volume 140, 2009

use in late-harvest or ice wines. In most years, there is a two to three week difference in

time between soybean leaf senescence and the beginning of grape harvest. It is unknown

where H. axyridis populations which were previously residing in soybean are located during

this two to three week period. It is possible that H. axyridis simply uses grapes as a food

source immediately prior to overwintering, as sugars in grapes may be more efficiently

converted to stored energy in the fat body of the insect than proteins from aphids (Hodek

1973, Denlinger 2005). In this scenario, grapes may be a preferred food of H. axyridis.

However, if ripe grapes are preferred over aphids by H. axyridis, beetles would be observed

in vineyards in every year, and not just when aphids are scarce.

A possible explanation may be found in the overwintering habits of A. g/ycines. As

day length decreases and soybean leaves senesce, A. glycines migrates to the overwintering

host, buckthorn (Rhamnus spp.) (Voegtlin et al. 2004). Mating occurs on this host and eggs

are laid on buckthorn buds (Ragsdale et al. 2004). In Ontario, oviposition by A. glycines

typically occurs by late October (Welsman et al. 2007). In this study, during years when

A. glycines infestation remained below economic threshold (i.e. 2004 and 2006), moderate

populations of aphids were observed in soybean fields immediately prior to soybean plant

senescence. In a companion study in the same years Welsman et al. (2007), found that high

numbers of overwintering eggs of A. glycines were found in Rhamnus cathartica stands in

Ontario (Table 1). Conversely, in 2005, when soybean fields monitored in this study had

heavy, early infestations of A. glycines, very few overwintering eggs were observed using

the same protocols as in 2004 and 2006 (Table 1).

We propose that interactions between 4. glycines and H. axyridis on the primary

host of A. glycines in spring and again after soybean senescence on the overwintering host

of A. glycines, play a larger role in dictating the abundance of H. axyridis in vineyards than

do late summer interactions in soybean, as previously speculated. Large populations of A.

glycines on its overwintering host may “kick-start” or “distract” H. axyridis, depending on

the time of year at which it occurs. Abundance of A. g/ycines early in the season initiates

(kick-starts) population growth of H. axyridis. Abundances of A. glycines on R. cathartica

in autumn function to draw H. axyridis away (distract) from vineyards in the fall (Fig. 3).

In spring, H. axyridis are usually found on Rhamnus cathartica almost immediately

after bud swell in mid to late April, feeding on aphids, and mating (Bahlai et al. 2007, Bahlai

et al. 2008). Rhamnus cathartica leaves begins to grow earlier than most other woody

plants in southern Ontario agroecosystems and egg hatch of A. gl/ycines coincides with this

event (Bahlai et al. 2007), so it is likely that A. glycines on R. cathartica represent one of

the earliest abundant food sources for H. axyridis (Fig. 3A iii). Predation by coccinellids,

predatory bugs, and syrphid larvae and parasitism by braconid and aphilinid wasps have

been shown to affect the population dynamics of soybean aphid in soybean fields (eg: Fox

et al. 2004, Heimpel et al. 2004, Fox et al. 2005, Desneux et al. 2006, Brosius et al. 2007).

Welsman et al. (2007) found that predation, rather than parasitism, slows the growth of these

early-season populations of A. glycines occurring on buckthorn, and attributed most of the

mortality to coccinellids. Oviposition among coccinellids typically occurs in the presence

of food (Hodek 1973) so it is reasonable to speculate that abundance of A. glycines in April

may allow H. axyridis to oviposit earlier in the season than would have occurred otherwise,

effectively ‘kick-starting’ the population growth of H. axyridis.

In early summer, A. glycines migrates to its summer host, soybean. When large

Baldaivausd.Seate JESO Volume 140, 2009

numbers of aphids are observed in soybean, increasing numbers of larvae and adults of H.

axyridis are observed soon after. Predation can cause a crash in aphid populations by the

end of the season (Fig. 3A ii, shaded curve) (Fox et al. 2004) . Alternatively, insecticides

may be applied to soybean fields for aphid control, causing aphid numbers to decline in

soybean fields (Fig. 3A ii, solid curve).

In mid-September, A. glycines migrates back to its overwintering host, R.

cathartica, where it remains feeding on foliage until oviposition occurs, usually around the

time the shrub drops its leaves in late October (Welsman et al. 2007). Rhamnus cathartica

retains its leaves later than many other plants in southern Ontario agroecosystems, so this

A) High year (kick-start)

Relative abundance

Soybean

Buckthorn

Spring Summer Autumn

B) Moderate-low year (distract)

Relative abundance

Buckthorn Soybean

Spring Summer Autumn

FIGURE 3: Hypothetical “kick-start- distract’ model of interaction between Harmonia

axyridis and Aphis glycines. In this scenario, early season abundances of A. gilycines on

buckthorn ‘kick-start’ population growth of H. axyridis, and late season abundances of A.

glycines ‘distract’ H. axyridis from grapes until after harvest. A) Kick-start year, B) Distract

year. Illustrated for each year are hypothetical abundances of 1) H. axyridis, ii) A. glycines

on soybean and iti) A. glycines on buckthorn. Dotted vertical lines represent grape harvest.

Population dynamics of Asian lady beetles and aphids JESO Volume 140, 2009

host may represent the last reservoir of aphid populations before winter within the Niagara

region agroecosystem. In years when A. glycines is abundant on its overwintering host, H.

axyridis typically is observed with the aphid (Bahlai et al. 2008), and is ‘distracted’ from

ripening grapes in vineyards (Fig. 3 B iii).

In years when A. glycines is not abundant on buckthorn, H. axyridis aggregates in

large numbers in Niagara Peninsula area vineyards (Fig. 3A 1, 111) (Welsman et al. 2007).

When aphids are scarce, beetles may move to ripening grapes because volatiles released by

fermentation of fruit may be similar to volatiles associated with aphid honeydew (Bahlai

et al. 2008). If this is the case, ripe grape odour could ‘trick’ H. axyridis into foraging in

vineyards for aphids, or simply act as a cue for the location of a ‘next best’ food source.

In years when a high number of aphid eggs had been observed in the previous

winter, both A. glycines and H. axyridis were observed at higher numbers in soybean in

July. Higher counts of H. axyridis were observed in late July in 2005 than in 2004 and

2006. However, when insecticides were applied to these fields in August 2005, numbers of

H. axyridis, like aphid infestations, decreased immediately, and persisted at low levels for

the remainder of the season (Fig. 2). Yet we observed substantial numbers of H. axyridis

feeding on aphids living on weeds in naturalized and semi-naturalized areas adjacent to

our observation fields in mid to late August of 2005 and large numbers of H. axyridis were

observed in vineyards that year. In years where only spot infestations of H. axyridis were

observed in vineyards (i.e. 2004 and 2006), abundance of A. g/ycines on the overwintering

host was observed. In these years, lower numbers of H. axyridis could have been sated by

large populations of aphids preparing to mate and oviposit on buckthorn.

This kick-start/distract model for the interaction of H. axyridis with A. glycines,

combined with insecticide application practices, may help to explain the biennial cycle of

infestation for both A. glycines and H. axyridis. To develop an effective integrated pest

management strategy to control vineyard infestations of H. axyridis, several specific areas

of inquiry should be pursued. Population monitoring of these beetles and their prey should

be continued to gain data regarding numbers and distribution; monitoring of A. glycines

and H. axyridis should continue in Niagara Peninsula area soybean fields, and should be

expanded to include populations of aphids in other crops and weeds in late summer. This

monitoring could provide information about agroecosystems in which H. axyridis occurs

in late summer. This may provide an early warning for potential vineyard infestations.

Because the interactions between 4. glycines and H. axyridis appear to be consistently

following a biennial cycle, additional population data can be used to refine predictions of

when and where outbreaks of these two species will occur, and under what conditions.

Acknowledgements

The authors would like to sincerely thank Zack Peters, Caitlin Smith, Kristen

Eddington, Emily MacLeod, Lisa Daoust, Chris Martin, Tracey Mancini (Sitek) and Erin

Jones for their help in the field, Neil Carter for his expertise and the advice of Rebecca

Hallett and Art Schaafsma. The authors would also like to acknowledge the funding

contributions of the Ontario Ministry of Agriculture, Food and Rural Affairs, the Grape

Growers of Ontario, the Wine Council of Ontario, and the Ontario Soybean Growers.

Bahlai and Sears JESO Volume 140, 2009

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Coccinellidae). Biological Control 28: 164-170.

Cutler and Rogers JESO Volume 140, 2009

NEW RECORD OF THE ASIATIC GARDEN BEETLE, MALADERA

CASTANEA (ARROW), IN ATLANTIC CANADA

G. C. Cutler', R. E. L. Rogers?

Department of Environmental Sciences, Nova Scotia Agricultural College

Truro, NS, Canada B2N 5E3

email: ccutler@nsac.ca

Scientific Note J. ent. Soc. Ont. 140: 40-45

The Asiatic garden beetle, Maladera castanea (Arrow) (Coleoptera: Scarabaeidae),

was first named by Arrow in the genus Autoserica in 1913, moved to the genus Aserica in

1927 by Arrow, and then moved by Pope to genus Maladera in 1961 (Evans and Smith

2005). Maladera castanea is endemic to the Russian Far East, Japan, North Korea, and

South Korea (Ahrens 2006). It was first collected in North America near Rutherford, New

Jersey in 1921 (Hallock 1929, 1930, 1936) but has been studied sporadically since 1927.

It is known to have established along the eastern seaboard from Massachusetts to South

Carolina, west to Pennsylvania and Ohio (Hallock 1936; Potter 1998). In those regions

it is generally a minor pest of turfgrass, ornamentals and some vegetables. However, M.

castanea may cause serious economic damage, is known to feed on more than 100 host

plants, and may be locally abundant, particularly in weedy or abandoned areas (Hallock

1936; Koppenhofer and Fuzy 2003; Tashiro 1987).

Maladera castanea appears to have been first collected in Canada in Saint-

Armand, Québec, in 1996 (Chantal 2003). Specimens were subsequently found in multiple

locations of southern Québec (Bostanian et al. 2003; Chantal 2003). Here, we document

the collection of M. castanea from Cumberland County, Nova Scotia, which we believe is

the first record of this insect in Atlantic Canada.

Collections occurred in a commercial, wild (syn. “lowbush”) blueberry (Vaccinium

angustifolium Ait.) growing area near Fox River, Cumberland Co., Nova Scotia (N45o0

40.83’, W640 53.43’). One particular field was described by the producer as a “flag-ship”

field, historically producing high numbers of berries. For reasons that were unknown to the

grower, production in the field had decreased and attempts to rejuvenate the field through

conventional fertilizer, pesticide and irrigation practices were unsuccessful. Soil samples

(approximately 20 x 20 x 20 cm) were collected with a spade shovel on 21 May 2003, 17

July 2007, and in early June 2008, from areas in fields with poor plant growth, near the farm

road and along a hedgerow of trees that separated fields. Samples were sifted through in the

field or later in the laboratory and collected larvae were stored in 70% ethanol.

Published November 2009

' Author to whom all correspondence should be addressed.

? Wildwood Labs Inc. 53 Blossom Drive, Kentville, NS, Canada B4N 3Z1

New record of the Asiatic garden beetle in Atlantic Canada JESO Volume 140, 2009

An unexpectedly large number of Scarabaeidae larvae were present in several of

the soil samples, particularly those adjacent to patches of grass and sedge, common weeds

in wild blueberry fields. Formal counts of larvae from each sample were not conducted,

but several grubs were collected in 2003, around 40 in 2007, and several in 2008. We

also observed that many blueberry plant roots from samples containing these Scarabaeidae

larvae had suffered feeding damage, with extensive girdling and destruction of fibrous roots

and root hairs, as well as root necrosis as a result of this feeding (Fig. 1).

FIGURE 1. Damage to wild (lowbush) blueberry roots where M. castanea larvae were

found in Fox River, Nova Scotia, 2003. Photo: R.E.L. Rogers, Wildwood Labs Inc.

Cutler and Rogers JESO Volume 140, 2009

Anal slit

Pali (spines)

9th abdominal

segment

FIGURE 2. Rastral pattern on the 10th abdominal segment of a M. castanea larva, illustrating

the characteristic longitudinal anal slit and crescent-shaped transverse row of spines (adapted

from Tashiro 1987; with permission, NY State Agricultural Experiment Station).

Larval specimens were confirmed as M. castanea by R.E.L.R and G.C.C. Voucher

specimens have been deposited in the A. D. Pickett Entomology Museum at the Nova

Scotia Agricultural College and the Canadian National Collection of Insects, Arachnids and

Nematodes, Ottawa, ON. Maladera castanea \arvae can be most easily distinguished from

other scarabaeid turfgrass feeders such as the Japanese beetle, Popillia japonica Newman,

the oriental beetle, Exomala (syn. Anomala) orientalis Waterhouse, masked chafers,

Cyclocephala spp., and European chafer, Rhizotrogus majalis (Rhazoumowsky), by the

characteristic positioning of the anal slit and arrangement of spines, hairs and bare spaces

on the raster of the terminal (tenth) abdominal segment. A single, transverse row of spines

in a crescent shape is the most noticeable character (Fig. 2, 3a), and whereas the anal slit

may be transverse or Y-shaped in related species, it is essentially longitudinal in M. castanea

(Reding and Klein 2006; Tashiro 1987). Other distinguishing larval characters include very .

small claws of the metathoracic legs, as compared to the pro- and mesothoracic legs, and a

light-coloured, enlarged bulbous stipe of the maxilla (Fig. 3b). Maladera castanea larvae

are smaller than those of P. japonica, E. orientalis, and R. majalis, with full-grown third

instars being approximately 19 mm long. Maladera castanea adults were not collected,

but they are 8-11 mm long, dull chestnut-brown, with a velvety, slight iridescent sheen

(Tashiro 1987). Adult beetles generally conceal themselves in moist soil at the base of food

plants and grasses during the day. They fly only at night, but are highly attracted to lights,

a behaviour that has proved useful in collecting or monitoring for M. castanea (Tashiro

1987).

Soil samples were not collected throughout the blueberry fields in question, and

therefore it is not possible to correlate M. castanea with the progressively poorer berry yields

generated. However, white grubs, including M. castanea, are increasingly important pests

of highbush blueberries (Alm et al. 1999; Cowles 2005; Wise et al. 2007), other Vaccinium

spp. (Koppenhofer et al. 2008; Wenninger and Averill 2006), strawberries (LaMondia et

New record of the Asiatic garden beetle in Atlantic Canada JESO Volume 140, 2009

FIGURE 3. Maladera castanea larva from Fox River, Nova Scotia, 2008, (a) rastral pattern

on the 10th abdominal segment and (b) head illustrating the enlarged bulbous stipe of the

maxilla. Photos: R.E.L. Rogers, Wildwood Labs Inc.

Cutler and Rogers JESO Volume 140, 2009

al. 2002), turf (Koppenhofer and Fuzy 2003), and other crops, ornamentals and perennials

(Tashiro 1987). Hallock (1936) reported that adults and/or larvae may cause considerable

injury to many vegetables, including beets, carrots, corn, parsnips, peppers, and turnips,

but that larvae were almost always more numerous in grassy areas overgrown with weeds,

particularly in the presence of hawkweed (the preferred oviposition site), goldenrod,

wild asters and, to a lesser extent, sorrel. Indeed, we found M. castanea feeding on V.

angustifolium in patches next to a high density of grasses and other weeds. Further, being

an unfamiliar, subterranean root feeder with few natural enemies (Tashiro 1987), there is

potential for undetected population growth.

Although it is unknown how M. castanea became established in the Fox River area,

the producer revealed that there is occasional back-and-forth transport of farm machinery

(e.g. tractors, harvesters) from operations in the state of Maine where the beetle is known

to exist, suggesting cross-border transport. Alternatively, this M4. castanea record could

simply be the product of natural expansion throughout North America. With recent intensive

efforts of C.G. Majka and colleagues to document Coleoptera occurrence in the Maritimes

(http://www.chebucto.ns.ca/Environment/NHR/PDF/index.html), it is somewhat surprising

that M. castanea has not been found earlier or elsewhere. Whether the geographic range of

M. castanea in this region is poorly understood or if the beetle is of sporadic occurrence is

unclear. Future work will attempt to map the distribution of M. castanea throughout Nova

Scotia.

Acknowledgements

We thank L. Babineau, K. Ramanaidu and D. Mclsaac for their help during collection

of M. castanea larvae, R. Delbridge, G. Brown and K. MacKenzie for their assistance, and

G. Williams for reviewing an early version of the manuscript. Partial funding for this work

was provided by the Wild Blueberry Producers Association of Nova Scotia, Agriculture and

Agri-Food Canada through Agri-Futures Nova Scotia, Conseil pour le développement de

agriculture du Québec, the New Brunswick Agricultural Council/Counseil Agricole due

Nouveau-Brunswick, the PEI Adapt Council, Agri-Adapt Council Inc. and the Nova Scotra

Department of Agriculture Technology Development Program.

References

Ahrens, D. 2006. Subfamily Sericinae Kirby, 1837, p. 229-248, Jn I. L6bl and A. Smetana,

eds. Catalogue of Palaearctic Coleoptera. Apollo Books, Stenstrup, Denmark.

Alm, S.R., M.G. Villani and W. Roelofs. 1999. Oriental beetles (Coleoptera: Scarabaeidae):

Current distribution in the United States and optimization of monitoring traps.

Journal of Economic Entomology 92: 931-935.

Bostanian, N.J., C. Vincent, H. Goulet, L. Lesage, J. Lasnier, J. Bellemare and Y. Mauffette.

2003. The arthropod fauna of Québec vineyards with particular reference to

phytophagous arthropods. Journal of Economic Entomology 96: 1221-1229.

New record of the Asiatic garden beetle in Atlantic Canada JESO Volume 140, 2009

Chantal, C. 2003. Six nouvelles mentions québécoises de coléoptéres. Fabreries 28: 25-

30.

Cowles, R.S. 2005. White grub management in blueberries. Proceedings of the New England

Vegetable & Fruit Conference. Manchester, New Hampshire.

Evans, A.V. and A.B.T. Smith. 2005. An electronic checklist of the New World chafers

(Coleoptera: Scarabaeidae: Melolonthinae). DigitalCommons@University of

Nebraska - Lincoln. http://digitalcommons.unl.edu/entomologypapers/2.

Hallock, H.C. 1929. Known distribution and abundance of Anomala orientalis Waterhouse,

Aserica castanea Arrow, and Serica similis Lewis in New York. Journal of

Economic Entomology 22: 293-299.

Hallock, H.C. 1930. Some observations upon the biology and control of Aserica castanea

Arrow. Journal of Economic Entomology 23: 281-286.

Hallock, H.C. 1936. Notes on biology and control of the Asiatic garden beetle. Journal of

Economic Entomology 29: 348-356.

Koppenhofer, A.M. and E.M. Fuzy. 2003. Biological and chemical control of the Asiatic

garden beetle, Maladera castanea (Coleoptera: Scarabaeidae). Journal of Economic

Entomology 96: 1076-1082.

Koppenhofer, A.M., C.R. Rodriguez-Saona, S. Polavarapu and R.J. Holdcraft. 2008.

Entomopathogenic nematodes for control of Phyllophaga georgiana (Coleoptera:

Scarabaeidae) in cranberries. Biocontrol Science and Technology 18: 21-31.

LaMondia, J.A., W.H. Elmer, T.L. Mervosh and R.S. Cowles. 2002. Integrated management

of strawberry pests by rotation and intercropping. Crop Protection 21: 837-846.

Potter, D.A. 1998. Destructive turfgrass insects: Biology, diagnosis, and control. Ann Arbor

Press, Chelsea, MI.

Reding, M.E. and M.G. Klein. 2006. Common white grubs of northeast Ohio

nurseries. Government Publication/Report. www.ars.usda.gov/sp2UserFiles/

Place/36071000/Publications/Reding192314 2006 CommonWhiteGrubs.pdf.

Tashiro, H. 1987. Turfgrass insects of the United States and Canada. Cornell University

Press, Ithaca, NY.

Wenninger, E.J. and A.L. Averill. 2006. Mating disruption of oriental beetle (Coleoptera:

Scarabaeidae) in cranberry using retrievable, point-source dispensers of sex

pheromone. Environmental Entomology 35: 458-464.

Wise, J.C.,C. Vandervoort and R. Isaacs. 2007. Lethal and sublethal activities of imidacloprid

contribute to control of adult Japanese beetle in blueberries. Journal of Economic

Entomology 100: 1596-1603.

Timms JESO Volume 140, 2009

GROWING PAINS: HOW THE BIRTH OF THE

ENTOMOLOGICAL SOCIETY OF CANADA AFFECTED THE

IDENTITY OF THE ENTOMOLOGICAL SOCIETY OF ONTARIO

L. TIMMS

Faculty of Forestry, University of Toronto

33 Willcocks Street, Toronto, Ontario, Canada MS5S 3B3

email: laura.timms@utoronto.ca

Special Contribution J. ent. Soc. Ont. 140: 49-56

The histories of the Entomological Societies of Ontario and Canada are inextricably

entwined. Both lay claim to the same founding story and early historical narrative', yet

also maintain their identities as distinct organizations. Superficially, the history is simple:

the Entomological Society of Canada (ESC) was founded in 1863, and changed its name

to the Entomological Society of Ontario (ESO) in 1871. The name change reflected the

geopolitical changes of the period (Ontario was one half of the Province of Canada when

the society was formed and part of the rapidly growing Dominion of Canada when the name

changed), as well as a pledge of annual financial support from the government of Ontario

received in that year (Saunders 1883). Despite its new name the ESO continued to operate

as a national body for almost eighty years, with branches across the country. However, in

the period after the Second World War a number of members began to suggest that it was

time for the formation of a truly national society. Thus, in 1950 the ESC was founded and

began to fulfill its chief function: “to serve as a national society and as the parent association

of, or as the link between, the other entomological societies in Canada” (Ozburn 1950).

The foundation of the new ESC effectively resulted in the demotion of the

ESO to a regional society. Although other histories written on the subject indicate that

this was a smooth transition (Spencer 1964; Holland 1966; Connor 1982), in this paper

| argue that the appropriation of their role by the national society caused a great deal of

conflict between the two societies as the members of the ESO were forced to reevaluate the

purpose and identity of the Ontario society. I will show how this anxiety was manifested

in disagreements between the parent society and its offspring over a number of matters in

the period between 1950 and 1963, as well as how the ESO began to redefine its identity in

the years after 1963. Specifically, I will examine the conflicts and issues surrounding the

disposition of shared assets and the organization of annual general meetings, as well as the

societies’ publications. This paper will also provide the first written history of the ESO in

the years after the foundation of the ESC. It is not my intention to stir up old controversies

or animosity, but to provide a written record of this period in the history of both societies — a

record that I hope will prove to be both informative and interesting to its readers.

' See for example the web pages for each society, which contain the same description of their origins:

About the ESO, online, no date, available at: http://www.entsocont.com/ (accessed: 1 May, 2008)

and History of the Entomological Society of Canada, online, no date, available at: http://esc-sec.org/

(accessed: 1 May, 2008)

a a aC A ae Cy

Essay JESO Volume 140, 2009

A Complicated Relationship

The foundation of the ESC created a society whose affairs were so entangled with

those of the ESO that, much like a divorce, a legal agreement was required to sort out

which society was responsible for what. The Instrument of Agreement, developed after

much negotiation between the two societies, was signed on 1 November 1954 and applied

retroactively to the activities of the previous three years (Ozburn and Baker 1954). The

Instrument formalized a number of the arrangements that had been vaguely outlined in the

original motion approving the formation of the ESC (Ozburn 1950). These included the

point that the ESO should retain possession of its library, a collection of some significance

accumulated throughout the life of the Society, as well as the periodicals received in

exchange for the societies’ publications.’ It also defined the understanding that membership

in the ESC was compulsory for members joining the ESO. Its main purpose, however, was

to clarify the responsibilities related to the joint publication of The Canadian Entomologist,

an internationally recognized journal which had been published continuously on a monthly

basis since 1868. Although it seemed clear that a journal with that title should be published

by a national society, after having published it for more than 80 years the ESO was not ready

to give it up completely.

By the end of the 1950s, the Instrument of Agreement, whose articles had been

designed to help the ESC get off the ground, was causing a great deal of friction between

the two societies and was in serious need of revision. In 1957, inspired by a conflict over

the distribution of shared membership fees, a special meeting of the ESO Board of Directors

was called to address some of the issues that had been “smouldering for years” (Dustan

1957a) between the two societies. The ESO found itself in a difficult position; they no

longer wanted to be “tied to the Canada Society” (Dustan 1957b), but they also did not

want to lose their share in The Canadian Entomologist. By 1958, however, the ESO was

forced to admit that “for all intents and purposes the Canadian society had assumed full

control” (Peterson 1960) of the journal. A revised Instrument of Agreement was developed

in 1960, in which the ESO relinquished its rights as publisher of the journal but retained

certain residual rights (Peterson et al. 1960). From the ESO’s perspective, chief among

these residual rights was the request that their historical role be acknowledged in perpetuity

on the inside cover of The Canadian Entomologist; the particular wording of this clause was

the subject of much negotiation (Peterson 1960).

It is likely that much of the conflict between the ESC and the ESO was due to

their dual claim to the history of one of the oldest scientific societies in North America; it

wasn’t clear which, if either, had more of a right to it. This ambiguity stems from the fact

that, although the organization had Ontario in its name for seventy-nine years before the

founding of the national society, it spent the first eight years of its life with the designation

of Entomological Society of Canada. Most histories of the subject written by entomologists

take the stance that “though provincial in name, the Society was always national in

outlook and objectives” (Holland 1966) and claim the whole record as that of the ESC.

The only analysis written by a historian puts forward the opposite opinion; that the ESO

was effectively only ever a regional society and that there was no “truly and officially

national, professional scientific society” (Connor 1982) of entomology until 1960 when the

ESC assumed full control of The Canadian Entomologist. Regardless of which viewpoint

Timms JESO Volume 140, 2009

is correct, it seems likely that “the frictions and conflictions of interest” (Dustan 1957a)

between the two societies were not only result of the ESO being upset by the assumption of

its national role by the ESC, but also. by what it saw as the appropriation of its history.

One Hundred Years of Entomology

Shortly after the revision of the agreement between the ESO and the ESC, the

societies were faced with another challenge to their harmonious existence: the celebration

of the 100th Annual Meeting of the Society. In previous years, the annual meeting had been

one of the most important traditions for the ESO. Special exhibits and scrapbooks were put

together for the 25th, 50th, 60th, and 75th Annual Meetings? , large celebrations were thrown

for many of these events, including invitations to and participation by representatives of

societies and institutions across Canada, the United States, and even the United Kingdom

and Europe’. When the ESC was formed, it was decided that its annual meeting would

always be held in conjunction with one of the provincial societies — the former branches

of the ESO (Ozburn 1950). In its first ten years of existence as an independent society,

the ESC held joint annual meetings with the ESO four times. In light of both this and

their shared history, it is not surprising that the ESO and the ESC chose to co-organize the

100th annual meeting in 1963. However, given the already established disagreements and

resentments that were brewing, it seems inevitable that the situation would end badly.

Celebration of the centennial anniversary became a matter of intense debate and

controversy between the two societies, highlighting the underlying tensions between them.

On the surface, much of the debate was about the location of the meeting. At the 1960 Annual

Meeting of the ESO, the membership voted to hold the centennial meeting in Guelph, a place

that many felt was “inseparably linked with the growth and development of the Society”

(McBain Cameron 1962). The ESC centennial committee, however, felt that the meeting

should be held in Ottawa, a “location in keeping with the importance of the event” (Holland

1961). The centennial committee presented and won support for their case at the 1961 ESC

Annual Meeting in Quebec. The matter went back to the ESO membership at their 1961

Annual Meeting, and after “rather extensive discussion” (Holland 1961), ESO voted to keep

the Guelph decision. This resulted in a flurry of angry letters between the board members of

both societies, and a special ballot sent out to the membership asking which decision they

felt should stand — Guelph or Ottawa. The results of the ballot were dramatic; the decision

came in at 80 votes for Guelph and 81 votes for Ottawa, with two votes for Guelph coming

in after the deadline had passed (Wressell 1962b). The centennial committee got its way

and the meeting took place in Ottawa. The closeness of the vote, however, indicated that

beneath the “whole contentious mess” (Wressell 1962a) of the location of the centennial

> Programs of many of the Annual Meetings are available in the Entomological Society of Ontario

Collection, University of Guelph Archival and Special Collections, Boxes 9, 16 and 20.

>For example, the scrapbook for the 50th anniversary celebrations included telegrams and letters of

congratulations from 35 groups and institutions, and attendees of the meeting included representatives

from an additional 56 different societies, institutes, departments, etc. Entomological Society of

Ontario Collection, University of Guelph Archival and Special Collections, Box 19.

Essay JESO Volume 140, 2009

meeting, a deeper divergence between the two societies had formed.

Modern Times

In the decades following the centennial celebration, the ESO seems to have become

more resigned to its “now wholly provincial” (Holland 1961) role. As the ESC went on to

address matters of national policy in science’, the ESO became more concerned with keeping

their society solvent and relevant. In 1969, the Society gave away one of its most valuable

assets, the library it had negotiated to keep from the ESC, to the University of Guelph. It

did this despite the original efforts it had gone through to hold on to the library, and despite

the fact that it was “worth at least $50,000” (McBain Cameron 1969), because the space the

library was occupying in the Biology buildings was needed, and the ESO could not afford

to move them elsewhere (Herne 1968). The ESO also became more interested in letting the

ESC take on tasks that it might previously have handled. For example, in 1985 the Public

Education committee decided to stop pursuing the idea of creating a brochure to promote

careers in entomology because of the “feeling” that it should be “developed by the national

society’ (Anonymous 1985). Perhaps most surprising, especially in contrast to the issues of

the centennial celebration, is that the 125th annual meeting of the ESO seems to have passed

with a minimum of fanfare. It was not held jointly with the ESC, and the ESO secretary

remarked in the January newsletter that the meeting had “a smaller turnout than usual”

(Smith 1989) of only seventy attendees. Although there was a speaker at the meeting who

reviewed the contribution of the ESO over the years, the President reported that “financial

support was not found for a proposal to prepare a history of the Society” (Jaques 1989).

Financial problems became more critical for the ESO as it was forced to turn its

secondary publication, The Annual Report, into the primary journal of the Society after

letting go of The Canadian Entomologist for good. The Annual Report had never been

as widely read as The Canadian Entomologist; its continued publication was carried out

in large part to fulfill an obligation to the Ontario government. One of the stipulations of

the 1871 grant from the Ontario Council of Agriculture was that the society must furnish

an annual report on “insects injurious or beneficial to agriculture” (Saunders 1883).

For this reason, the papers published in the Report were often less representative of the

range of papers presented at the annual meeting than they were focused on economic and

applied issues of entomology. In an effort to change this image of the journal and boost

readership, in 1959 the ESO changed the name of the periodical to the Proceedings of the

Entomological Society of Ontario and began actively to solicit papers of all types. It is not

clear how closely the two events are related, but shortly after the name change, the Ontario

Department of Agriculture proposed to withdraw its financial support (Boyce 1968). The

ESO was then faced with the problem of supporting the cost of publication itself, which

* For example, the ESC became heavily involved in the Biological Council of Canada in the 1970s

and provided a number of briefs to the Federal Government on various issues such as the teaching

of Biology in Canadian Universities and the publication of Canadian Science Journals, see Boxes |

and 2 in the Entomological Society of Ontario Collection, University of Guelph Archival and Special

Collections.

Timms JESO Volume 140, 2009

left them wondering if The Proceedings were “worth the struggle”, especially “since most

entomologists publish elsewhere anyway” (Salkeld 1968).

Although the ESO had difficulty maintaining the relevance of and interest in its

publication over the past thirty years, it persevered, in large part due to the fact that the Board

has been “reluctant to break a series” (Ellis 1984). Once the society was responsible for

the cost of the publication, it was forced to institute-a page charge policy, requiring authors

to pay for each page of their articles. This meant fewer manuscripts were put forward, and

that one of the tasks of the editor was to constantly badger the membership for submissions

(eg. Ellis 1982, Prévost 2002, Richards 2006). Fewer submissions made it harder to stick

to the annual publication schedule; a variety of creative means to catch up were employed,

including publishing in one volume all of the papers presented at annual meeting symposia

(Anonymous 1985b; Kevan 1987; Bolter 1990), as well as dedicating volumes to particular

entomologists (Anonymous 1985a; Richards 2007). In 1989, severe financial troubles

obligated the ESO to solicit and accept donations from a variety of sources, including a

large pesticide company, in order to continue publication of the journal (Kinoshita 1989).

The most recent efforts to increase the profile of the publication included renaming it again

in 2003, this time as the Journal of the Entomological Society of Ontario (Prévost 2003),

as well as making it available online (Richards 2006). During all this time, “discontinuing

The Proceedings” was “out of the question” (Marshall 1989), an attitude which emphasizes

the ESO’s particular commitment to its history.

While much of the evident tension between the two societies appears to have

dissipated after the 1963 meeting, the ESC continued to experience problems with the

ESO that it did not encounter with other provincial societies, specifically related to the

organization of joint annual meetings. One past president of the ESC did not hesitate to

point out Ontario as an example of the “problems [that] do exist in some regions” (Cooper

1976). It was perhaps for this reason that the ESC executive voted in 1977 to hold a joint

meeting with the Entomological Society of America in 1982 in Toronto without the support

of the ESO, a decision that the Ontario society felt left them “out in the cold” (Smith 1977).

Finances and annual meetings were another issue. Beginning with the 1963 meeting, the

ESC attempted to establish a procedure for the sharing of profits and losses related to joint

annual meetings (Munroe 1962). As of 2005, the ESC was still trying to formalize this

process. Although most of the other provincial societies have abided by the “rather loose

arrangement” (Shore 2005) of sharing half the profits of joint annual meetings with the

national society, the ESO has not always been cooperative; after the 2001 joint annual

meeting in Niagara Falls the ESO declined to give any of the profits to the ESC (Hunt 2001),

causing much consternation in the national society. However, after the 2008 joint annual

meeting in Ottawa the ESO gave 51% of the profits to the ESC (C. Scott-Dupree, personal

communication), a sign of the generally amiable relationship that currently exists between

the two societies.

> Interestingly, volume 116 (1985) as well as volumes 137 and 138 (2006 & 2007) were all dedicated

to D.H. Pengelly, former secretary and treasurer of the ESO. As far as I am aware, no other person

has had volumes of the Proceedings / Journal dedicated to them.

> aaemeieelieieed

Essay JESO Volume 140, 2009

Conclusion

All previous histories of the ESO and ESC have ended their narratives at or shortly

after the celebration of the centennial; in reading them one gets the impression that the

creation of the national society had been the ultimate goal of the ESO. This paper has

shown that this was not the case, that it took at least a decade for the members of the ESO

to adjust to their altered role as a provincial society and that occasional remnants of this

strain remain in evidence to this day. Furthermore, the shared history and subsequently

tumultuous division of the two societies created a distinctive connection between them

that deserves to be celebrated and explored. This investigation should be of interest to

those wishing to produce more complete histories as the 150th anniversary of organized

entomology in Canada approaches.

Acknowledgments

The assistance of the staff at the archives, McLaughlin Library, University of

Guelph, in providing access and photocopies of archival material is greatly appreciated.

Information and suggestions provided by Kevin Barber (Treasurer, ESO) and Sandy Smith

(Former Secretary and President of the ESO) were also helpful. Finally, I thank Steve

Walker for useful comments on an earlier version of this manuscript.

Notes and References

Anonymous. 1985a. Dedication. Proceedings of the Entomological Society of Ontario

116: iv.

Anonymous. 1985b. White pine symposium: foreword. Proceedings of the Entomological

Society of Ontario — Supplement to volume 116: v.

Bolter, C.J. 1990. Sustainable agriculture and forestry: an entomological perspective.

Proceedings of the Entomological Society of Ontario 121: 1-3.

Boyce, H.R. 1968. Letter to D.H. Pengelly, dated 16 January 1968. Entomological Society

of Ontario Collection, University of Guelph Archival and Special Collections, Box

2h.

Connor, J.T.H. 1982. Of Butterfly Nets and Beetle Bottles: The Entomological Society

of Canada, 1863-1960. History of Science and Technology in Canada Bulletin 6:

154-1716

Cooper, G.S. 1976. Letter to M.E. MacGillivray, dated 6 December 1976. Entomological

Society of Ontario Collection, University of Guelph Archival and Special

Collections, Box 1.

Dustan, G.G. 1957a. Letter to B.M. McGugan, dated 22 November 1957. Entomological

Society of Ontario Collection, University of Guelph Archival and Special

Collections, Box 10.

Tine JESO Volume 140, 2009

Dustan, G.G. 1957b. Letter to A.S. West, dated 27 November 1957. Entomological Society

of Ontario Collection, University of Guelph Archival and Special Collections, Box

10. ;

Ellis, C.R. 1982. Editor’s Report, dated 26 November 1982. Entomological Society of

Ontario Collection, University of Guelph Archival and Special Collections, Box

Ellis, C.R. 1984. Editorial Committee Report. Entomological Society of Ontario Collection,

University of Guelph Archival and Special Collections, Box 17.

Herne, D.C. 1968. Letter to W.C. Allan, no date 1968. Entomological Society of Ontario

Collection, University of Guelph Archival and Special Collections, Box 10.

Holland, G.P. 1961. Report number two of the Centennial Committee to the Board of the

ESC, dated 15 December 1961. Entomological Society of Ontario Collection,

University of Guelph Archival and Special Collections, Box 21.

Holland, G.P. 1966. Entomology in Canada, pp. 7-13 /n Centennial of Entomology in

Canada 1863-1963. G.B. Wiggins (ed.), Toronto.

Hunt, D. 2001. Minutes of the Interim Board Meeting, dated 20 April 2001. Entomological

Society of Ontario Collection, University of Guelph Archival and Special

Collections, Box 31.

Jaques, R. 1989. Report of the Entomological Society of Ontario 1988-89. Entomological

Society of Ontario Collection, University of Guelph Archival and Special

Collections, Box 30.

Kevan, P.G. 1987. Alternative pollinators for Ontario’s crops: prefatory remarks to

papers presented at a workshop held at the University of Guelph, 12 April, 1986.

Proceedings of the Entomological Society of Ontario 118: 109-110.

Kinoshita, G.B. 1989. Letter to H. Goulet, dated 5 June 1989. Entomological Society of

Ontario Collection, University of Guelph Archival and Special Collections, Box

30.

Marshall, S. 1989. Treasurer’s Report. ESO Newsletter, January 1989, Entomological

Society of Ontario Collection, University of Guelph Archival and Special

Collections, Box 30.

McBain Cameron, J. 1962. Letter to H. B. Wressell, dated 14 March 1962. Entomological

Society of Ontario Collection, University of Guelph Archival and Special

Collections, Box 21.

MacBain Cameron, J. 1969. Letter to W.C. Allan, dated 10 March 1969. Entomological

Society of Ontario Collection, University of Guelph Archival and Special

Collections, Box 10.

Munroe, E.G. 1962. Letter to H.B. Wressell, dated 9 November 1962. Entomological

Society of Ontario Collection, University of Guelph Archival and Special

Collections, Box 21.

Ozburn, R.H. 1950. Report of Council, 1950. Annual Report of the Entomological Society

of Ontario 81: 6.

Ozburn, R.H. and Baker, A.W. 1954. Instrument of Agreement between the Entomological

Society of Canada and the Entomological Society of Ontario, Entomological Society

of Ontario Collection, University of Guelph Archival and Special Collections, Box

18.

Essay JESO Volume 140, 2009

Peterson, D.G. 1960. Letter to B. Hocking, dated 18 January 1960. Entomological Society

of Ontario Collection, University of Guelph Archival and Special Collections, Box

30.

Peterson, D.G, Allan, W.C., Hocking, B., and Reed, L.L. 1960. Revision to Instrument of

Agreement Between the Entomological Society of Canada and the Entomological

Society of Ontario. Entomological Society of Ontario Collection, University of

Guelph Archival and Special Collections, Box 30.

Prévost, Y. 2002. Proceedings of the Entomological Society of Ontario. Newsletter of

the Entomological Society of Ontario 7(2): 7. Available online at: http://www.

entsocont.com/newsletters.htm.

Prévost, Y. 2003. Journal of the Entomological Society of Ontario Renamed! Newsletter

of the Entomological Society of Ontario 8(1): 1. Available online at: http://www.

entsocont.com/newsletters.htm.

Richards, M.H. 2006. JESO News. Newsletter of the Entomological Society of Ontario

11(2): 3. Available online at: http://www.entsocont.com/newsletters.htm.

Richards, M.H. 2007. From the Editor. Journal of the Entomological Society of Ontario

138: 1.

Salkeld, E.H. 1968. Letter to D.H. Pengelly, no date 1968. Entomological Society of

Ontario Collection, University of Guelph Archival and Special Collections, Box

21,

Saunders, W. 1883. Annual Address of the President. Annual Report of the Entomological

Society of Ontario, 14: 8-13.

Shore, T. 2005. Survey for refinements to the arrangements between the Entomological

Society of Canada and the regional societies for organization and operation of joint

annual meetings, dated 21 November 2005. Received by e-mail to author from

David Hunt on November 22, 2005.

Spencer, G.J. 1964. A century of entomology in Canada. Canadian Entomologist 96: 33-

59.

Smith, M.V. 1977. Letter to E.A.C. Hagley, dated 18 October 1977. Entomological Society

of Ontario Collection, University of Guelph Archival and Special Collections, Box

Smith, S. 1989. 1988 Annual Meeting (Guelph). ESO Newsletter, January 1989,

Entomological Society of Ontario Collection, University of Guelph Archival and

Special Collections, Box 30.

Wressell, H.B. 1962a. Letter to A.W.A. Brown, dated 14 March 1962. Entomological

Society of Ontario Collection, University of Guelph Archival and Special

Collections, Box 21.

Wressell, H.B. 1962b. Letter to A.W.A. Brown, dated 24 August 1962. Entomological

Society of Ontario Collection, University of Guelph Archival and Special

Collections, Box 21.

Marshall JESO Volume 140, 2009

BOOK REVIEW

Field Guide to the Dragonflies and Damselflies of Algonquin Provincial Park and the

Surrounding Area. By Colin D. Jones, Andrea Kingsley, Peter Burke and Matt Holder.

Algonquin Field Guide Series, published by The Friends of Algonquin Park.

263 pp. ISBN 978-1-894993-29-6 (Soft cover: October 2008). $28.95 CAN.

This book is billed as a comprehensive field guide to the dragonflies and damselflies

found in Algonquin Provincial Park and surrounding area, but it is much, much more than

that. The coverage actually extends across south-central Ontario and into southwestern

Quebec, and includes 135 out of the provincial total of 172 Odonata species. The detailed,

full-colour illustrations set a new standard for the illustration of field guides with stunning

watercolours showing the intricate colours of males, females and variants in brilliant

detail.

The introductory text is excellent, with clear and nicely illustrated treatments of

Odonata morphology, behaviour, and life cycle, but the real strength of this book is in

the profusely illustrated and carefully organized identification tools that lead the reader to

informative treatments of each species. There are no keys, but instead the authors have

used tables and charts illustrated by line drawings, watercolors and some photographs. The

nine families involved are easily separated using three colour pages devoted to diagnosing

the families. Within each family there are pages combining line-drawings of the male

and female genitalic characters that define the species. These are of tremendous value in

confirming the identification of difficult species, but it is unlikely that the average user will

refer to them very often; most users will instead identify their odonate finds by thumbing

through the profusely illustrated species accounts in search of a “match”. Useful diagnostic

characters are highlighted or indicated with arrows and captions, and actual size is indicated

with a silhouette. Once a match is located, the reader is provided with a “description” (a

one-paragraph diagnosis), a very useful discussion of similar species, and information about

habitat, behaviour, abundance, and distribution. Flight period is given both in the text and

in graphical form along the heading for each species.

This is a wonderful book that I think belongs on the shelf of every entomologist and

naturalist in Ontario. Not only is it well organized, beautifully illustrated and informative,

it is also well-packaged. At 14 x 21 cm, it is just the right size to fit into a jacket pocket or

the outside pouch of a day pack, and it looks water-resistant and sturdy enough to hold up

to a bit of bashing. My main criticism of the book centers on what might be perceived by

some as its main strength, which is the degree to which it is focused on Algonquin Park.

i fo

Book review JESO Volume 140, 2009

As far as I know this book can only be purchased from bookstores or from Algonquin’s

website (www.algonquinpark.on.ca), which is likely to limit its readership despite the

current popularity of Odonata among naturalists. The popular online bookseller Amazon.ca

currently lists 36 books on dragonflies and damselflies (it is for good reason that dragonflies

have been described as the “new butterflies”), but Dragonflies and Damselflies of Algonquin

Provincial Park is not among them. This combination of limited availability and a local-

sounding title is likely to limit the number of readers with interests outside the Algonquin

area, which is unfortunate since this is a tremendously useful guide for most of Ontario, and

indeed much of northeastern North America. I’m looking forward to a later edition, or a

follow-up version including all 172 Ontario Odonata species!

STEVE MARSHALL

Department of Environmental Biology

University of Guelph,

Guelph, Ontario, Canada NIG 2W1

samarsha@uoguelph.ca

JESO Volume 140, 2009

THE ENTOMOLOGICAL SOCIETY OF ONTARIO

OFFICERS AND GOVERNORS

2009-2010

President: G. UMPHREY

Department of Mathematics and Statistics

University of Guelph, Guelph, ON N1G 2W1

umphrey@uoguelph.ca

President-Elect: H. FRASER

Ontario Ministry of Agriculture, Food and Rural Affairs

4890 Victoria Ave. North, P.O. Box 8000

Vineland, ON LOR 2E0

hannah. fraser@ontario.ca

Past President: C. SCOTT-DUPREE

Dept. of Environmental Biology

University of Guelph, Guelph, Ontario NIG 2W1

cscottdu@uoguelph.ca

Secretary: N. MCKENZIE

Vista Centre, 1830 Bank Street, P.O. Box 83025

Ottawa, ON K1V 1A3

nicole_mckenzie@hc-sc.gc.ca

Treasurer: K. BARBER (until 31 Dec. 2009)

Natural Resources Canada, Canadian Forest Service

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

kbarber@nrcan.ge.ca

S. LI (beginning | Jan. 2010)

Pest Management Centre, Building 57

Agriculture and Agri-Food Canada

960 Carling Ave., Ottawa, ON K1A 0C6

Dr.Shiyou.Li@nrcan.ge.ca

Directors:

H. DOUGLAS

Canadian Food Inspection Agency

960 Carling Ave., Ottawa ON K1A 06C

douglash@inspection.ge.ca

S. LACHANCE (2010-2011)

Université de Guelph - Campus d’ Alfred

31 St. Paul Street, Alfred, ON KOB 1A0

Slachance@alfredc.uoguelph.ca

S. KULLIK

Department of Environmental Biology

University of Guelph, Guelph, ON NIG 2W1

sigrun.kullik@sympatico.ca

K. RYALL (2009-2011)

Canadian Forest Service, Great Lakes Forestry Centre,

Natural Resources Canada, 1219 Queen Street East,

Sault Ste. Marie, ON P6A 2E5

Krista.Ryall@NRCan-RNCan.gc.ca

K. RYAN (2008-2010)

Faculty of Forestry, University of Toronto

Toronto, ON MSS 3B3

kathleen.ryan@utoronto.ca

I. SCOTT

Agriculture and Agri-Food Canada

1391 Sandford Street, London, ON NSV 4T3

Ian.Scott@agr.gc.ca

(2008-2010)

(2009-2011)

(2010-2011)

ESO Regional Rep to ESC: H. DOUGLAS

Canadian Food Inspection Agency

960 Carling Ave., Ottawa ON K1A 06C

douglash@inspection.gc.ca

- Librarian: J. BRETT

Library, University of Guelph

Guelph, ON NIG 2W1

jimbrett@uoguelph.ca

Newsletter Editor: J. ALLEN

Ontario Ministry of Agriculture, Food and Rural Affairs

1 Stone Road West, Guelph, ON NIG 4Y2

jennifer.allen@ontario.ca

Student Representative: J. GIBSON

Department of Biology, Carleton University

1125 Colonel By Drive, Ottawa, ON K1S 5B6

jgibson5@connect.carleton.ca

Website: B. LYONS

Canadian Forest Service, Great Lakes Forestry Centre,

Natural Resources Canada, 1219 Queen St East,

Sault Ste. Marie, ON P6A 2E5

Barry.Lyons@NRCan-RNCan.ge.ca

JESO Editor: M. RICHARDS

Dept. of Biological Sciences, Brock University

St. Catharines, ON L2S 3A1

miriam.richards@brocku.ca

Technical Editor: S. REHAN

Dept. of Biological Sciences, Brock University

St. Catharines, ON L2S 3A1

sandra.rehan@brocku.ca

Associate Editors:

A. BENNETT

Agriculture and Agri-Food Canada

960 Carling Ave., Ottawa ON K1A 06C

N. CARTER

154 Riverview Blvd.

St. Catharines, ON L2T 3M7

J. SKEVINGTON

Agriculture and Agri-Food Canada

Eastern Cereal and Oilseed Research Centre

960 Carling Ave., Ottawa, ON K1A 0C6

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, and the right to publish in the

Journal, and receive the Newsletter and the Journal. Students, amateurs and retired

entomologists within Canada 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.ca

APPLICATION FOR MEMBERSHIP

Please send your name, address (including postal code) and email address to:

Nicole McKenzie, Secretary, Entomological Society of Ontario

c/o Vista Centre, 1830 Bank Street, P.O. Box 83025 Ottawa, ON K1V 1A3

email: nicole mckenzie@hc-sc.gc.ca

NOTICE TO CONTRIBUTORS

Please refer to the Society web site (http://www.entsocont.ca) for current instructions to

authors. Please submit manuscripts electronically to the Scientific Editor

(miriam.richards@brocku.ca).

ie © 8) See

CONTENTS

L FROM THE EDIT OR cvccscseasesshoccstscnassesontrsiestamaaenaae sesseeessssensesnencnsnsneneneneseenees siintooreipusaiietdl ra

> J

II. ARTICLES vie q

B. J. COOKE, F. LORENZETTI and J. ROLAND — On the duration and distill of

forest tent caterpillar outbreaks in east-central Camada.......siccscssosscsnssesssescessossnvenannssencessoonsesne dite a :

J.H.SKEVINGTON and J.A. GOOLSBY — New records of Pipunculidae attacking proconiine | ’ !

sharpshooters (Auchenorrhyncha: Cicadellidae: oe t

Cc. A. BAHLAI and M. K. SEARS — Population dynamics of Harmonia axyridis and /

glycines in Niagara peninsula soybean fields and vineyards. sinuses

Ill. NOTE

G. C. CUTLER and R. E. L. ROGERS — New record of the Asiatic garden beetle, Mel adera

castanea (Arrow), i In Atlantic CamBGBcsscccadecacdcsbaninneseu SECS SSE SET EEEETESSE SHES EE ESTE EES ri ee 45 ' i

IV. ESSAY

L. TIMMS — Growing pains: How the birth of the Entomological Society of Canada affect

the identity of the Entomological Society of Ontario.......... sinviinensiotemiaseii soo ciniataioaaa ™

V. BOOK REVIEW

S. MARSHALL — Field Guide to the Dragonflies and Damselflies of Algonquin Provincial ‘

Park and the Surrounding Area. 2008. by Colin D. Jones, Andrea Kingsley, Peter Burke and

Matt HON er snssnsessnnoseseentserennstctatnoescoeonteendionnansctasss leap |

VI. ESO OFFICERS AND GOVERNORS 2009-2010 .svscsesesesersctnttuememenenerenernmnrntee 56

) a

VII. ESO OFFICERS AND GOVERNORS 2008-2009 ..osssssssseseseesuseseneeeusinside front cover —

VIN. FELLOWS OF THE FSQ.3..1425.0/5)5 2c. .-suseeninside front cover

IX. APPLICATION FOR MEMBERSHIP.......scccsssccosecccsesccosseccoveeccssseconsseesvees inside back cov er i

X. NOTICE TO CONTRIBUTORS x....<.:enssncstsssenisoxciercscieuieeeseeaaanenan .inside back cover

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