NATIVE PARASITES OF COLEOPHORA WlRICELLA (LEPIWPTERA

NATIVE PARASITES OF COLEOPHORA WlRICELLA (LEPIWPTERA:

COLEOPHORIDAE) IN BRITISH COLUMBIA

Gordon Edward Miller

BSc.(Hons.), Simon Fraser University, 1972

A THESIS SUBMITTED IN PARTIAL FULFILLMENT OF THE

mQUIWMENTS FOR THE DEGREE OF

MASTER OF SCIENCE

in the Department

of

Biological Sciences

@ Gordon Edward Miller 1976 SIMON FRASER UNIVERSITY

July 1976

All rights reserved. This thesis may not be re- produced in whole or in part, by photocopy or other means, without permission of the author.

APPROVAL

Name: Gordon Edward Miller

Degree: Master of Science

Title of Thesis: Native Parasites of Coleophora l a r i c e l l a (Lepidoptera: Coleophoridae) in British Columbia.

Examining Committee:

Chairperson: Dr. J. S. Barlow

Prof. T. Finlayson 4 Senior Supervisor

--- - ' \. Dr. 2. H. Borden

! - //

Dr. J. P. M. Mackauer

- 6 - r - , * Dr. R. F. ~hepKerd

cific Forest Research Centre Canadian Forestry Service

Victoria, B.C.

-,/- - y . ' - - ~ r r B. P. Beirne 2

External ~xamin< Professor

Simon Fraser University

Date Approved : 2 8 I / 7/76 ' -

I hereby g r a n t t o Simon F r a s e r U n i v e r s i t y t h e r i g h t t o lend

my t h e s i s o r d i s s e r t a t i o n ( t h e t i t l e o f which i s shown be low) t o u s e r s

o f t h e Simon F r a s e r U n i v e r s i t y L i b r a r y , and t o make p a r t i a l o r s i n g l e

c o p i e s o n l y f o r s u c h u s e r s o r i n r e s p o n s e t o a r e q u e s t f rom t h e l i b r a r y

o f a n y o t h e r u n i v e r s i t y , o r o t h e r e d u c a t i o n a l i n s t i t u t i o n , on i t s own

b e h a l f o r f o r one of i t s u s e r s . I f u r t h e r a g r e e t h a t p e r m i s s i o n f o r

m u l t i p l e copy ing o f t h i s t h e s i s f o r s c h o l a r l y p u r p o s e s may be g r a n t e d

b y me o r t h e Dean of Gradua te S t u d i e s . It i s u n d e r s t o o d t h a t c o p y i n g

o r p u b l i c a t i o n of t h i s t h e s i s f o r f i n a n c i a l g a i n s h a l l n o t b e a l l o w e d

w i t h o u t my w r i t t e n p e r m i s s i o n .

T i t l e o f ~ h e s i s I ~ i s s e r t a t i o n :

Native Pa ras i t e s of Coleophora L a r i c e l l a (Lepidoptera: Coleophor ida~)

i n B r i t i s h Columbia

A u t h o r :

( s i g n a t u r e )

Gordon Edward Mil ler

(name )

Ju ly 28hh, 1976

( d a t e )

ABSTRACT

The parasite complex of the larch casebearer, Coleophora l a r i -

c e l l a (Hbn.), was examined in the Kootenay area of ~ r i t i s h Columbia i n

1973 and 1974, pr ior t o the establishment of exotic parasites released

for biological control of the larch casebearer. Forty-two species of

hymenopterous parasites were obtained from mass-rearings of fourth-instar

larval and pupal C. l a r i c e l l a , of which 30 were considered actual para-

s i t e s of C. l a r i c e l l a . Fifteen species were confirmed by individual

rearings. No parasites were obtained from eggs, needle-mining larvae o r

casebearing third-instar larvae. The complex of C. l a r i c e l l a parasites

i n British Columbia was comparable t o tha t in other areas of North

America, .as well as the parasite complex of a native coleophorid in

British Cplumbia. Dicladocerus spp. and S p i l o c h a l c i s a l b i f r o n s (Walsh)

dominated the complex in British Columbia. A l l of the parasite species

obtained were non-specific and non-synchronized with C . l a r i c e l l a . The

incidence of parasitism ranged from O t o 27.7 percent.

Density of C. l a r i c e l l a and percentage parasitism by Dic ladocerus

spp. and S . a l b i f r o n s did not d i f f e r significantly between the five

height classes of t rees sampled. The within-tree distribution of C.

l a r i c e l l a , Dic ladocerus spp. and S . a l b i f r o n s varied, depending on the

t ree class sampled.

The exotic species tha t were recently released against C. l a r i -

c e l l a i n western North America are taxonomically related to the native

iii

s p e c i e s . They a r e non-spec i f ic and non-synchronized w i t h C. l a r i c e l l a

and have a minor role i n de te rmin ing l a r c h ca sebea re r popu la t i ons i n

Europe. A s a r e s u l t , t h e i r e f f e c t i v e n e s s a s b i o l o g i c a l c o n t r o l agen t s

may be l im i t ed .

ACKNOWLE DGMlENTS

I would l i k e t o express my apprecia t ion t o :

Professor T. Finlayson f o r guidance and supervision during

the course of t h e research and prepara t ion of t h i s manu-

s c r i p t ; D r s . J. H. Borden, J. P. M. Mackauer, and R. F.

Shepherd f o r advice on research and i n prepara t ion of t h e

manuscript; D r s . A. L. Turnbull and J. A. McLean f o r

advice on s t a t i s t i c a l ana lys i s ; and t o t h e s t a f f o f t h e

Entomology Research I n s t i t u t e , Agriculture Canada, Ottawa,

p a r t i c u l a r l y D r s . C. M. Yoshimoto, W . R. M. Mason, J. R.

Barron, and L. Masner, and Messrs. H. E. Bisdee and M.

Ivanochko f o r i d e n t i f i c a t i o n of p a r a s i t e specimens.

TABLE OF CONTENTS

Page

APPROVAL . . . . . . . . . . . . . . . . . i i

A B S T R A C T . . . . . . . . . . . . . . iii ACKNOWLEDGMENTS . . . . . . . . . . . . . . . v

TABLE OF CONTENTS . . . . . . . . . . . . . . . . vi

LIST OF TABLES . . . . . . . . . . . . . . . . . viii

. . . . . . . . . . . . . . . . LIST OF FIGURES X

INTRODUCTION . . . . . . . . . . . . a * . . . 1

GENERAL SURVEY FOR PARASITES OF COLEOPHORA LARICELLA IN BRITISH COLUMBIA . . . . . . . . . . . Methods and Materials . . . . . . . . . . . . . . Survey for Parasites of Mature Larvae and Pupae . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . Mass Rearings

Individual Rearings . . . . . . . . . . . . . Survey for Egg Parasites . . . . . . . . . . . . Survey for Parasites of Early-Instar

. . . . . . . . . . . . . . . . . Larvae

~ensities of Coleophora laricella in . . . . . . . . . . . . . . British Columbia

Parasites of Coleophora laricella in . . . . . . . . . . . . . . . British Columbia

Page

Parasites of Mature Larvae and Pupae . . . . . . . 9

Species Obtained . . . . . . . . . . . . . 9

. . . . . . . . . . . . . Introduced Species 17

Relative Importance of Parasite Species . . . . . . . . . . . . . . . . 18

Parasitism at Individual Sites . . . . . . . . . 18

Aspects of Parasite Biology . . . . . . . . . . 22

. . . . . . . . . . . . . . . Brood Size 22

Hyperparasitism . . . . . . . . . . . . . 22

Emergence Patterns . . . . . . . . . . . . 23

Parasites of Egg and Early-Instar Larvae . . . . . . . . . . . . . . . . . 23

DISTRIBUTIONS OF COLEOPHORA LARICELLA AND ITS PARASITES INWESTERNLARCHCROWNS . . . . . . . . 26

Methods and Materials . . . . . . . . . . . . . . 26

Results and Discussion . . . . . . . . . . . . . 27

CONCLUDING DISCUSSION . . . . . . . . . . . . . . 46

APPENDICES

I . SITE DESCRIPTIONS . . . . . . . . . . . . . . 52

I1 . PERCENTAGE PARASITISM BY INDIVIDUAL SPECIES AT SITES IN BRITISH COLUMBIA . . . . . . . 61

REFERENCES . . . . . . . . . . . . . . . . . . 72

CURRICULUM VITAE

LIST OF TABLES

Table Page

I, Numbers reared and densities of Coleophora laricella in samples of mature larvae and pupae taken from eight 1973 and 14 1974 collection sites in British Columbia . . . . . . . 10

11. Parasites obtained from mass-rearings of mature larvae and pupae of Coleophora laricella in 1973 and 1974 collections in British Columbia . . . . . . . . . . . . . . 13

111. Number of taxa and total percentage parasitism at individual sites in 1973 and 1974 collections in British Columbia . . . . . . . . . 2 0

IV. Density of Coleophora laricella and percentage parasitism by Dicladocerus spp. and by Spilochalcis albifrons in five classes of trees on 13 June 1974 at Shoreacres, British Columbia . . . . . . . . . . . . . . . . 28

V. Analysis of variance for the distribution of. Coleophora laricella in five classes of trees on 13 June 1974 at Shoreacres, British Columbia . . . . . . . . . . . . . . . . 29

VI. Analysis of variance for the distribution of Dicladocerus spp. in five classes of trees on 13 June 1974 at Shoreacres, British Columbia . . . . . . . . . . . . . . . . 3 0

VII. Analysis of variance for the distribution of Spilochalcis albifrons in five classes of trees on 13 June 1974 at Shoreacres, British Columbia . . . . . . . . . . . . . . . . 31

VILI. Analysis of variance for the within-tree distributions of Coleophora laricella in one class of tree on 15 May and five classes of trees on 13 June 1974 at Shoreacres, British Columbia . . . . . . . . . . . . . . . . 34

viii

Table Page

IX. Analysis of variance for the within-tree distributions of Dicladocerus spp. in five classes of trees on 13 June 1974 at Shoreacres, British Columbia . . . . . . . . . . 39

X. Analysis of variance for the within-tree distributions of Spilochalcis albifrons in five classes of trees on 13 June 1974 at Shoreacres, British Columbia . . . . . . . . . . 4 3

XI. Percentage parasitism of Coleophora laricella by individual parasite species in samples taken at eight sites in British Columbiaon8-9May1973 . . . . . . . . . . . 6 2

XII. Percentage parasitism of Coleophora laricella by individual parasite species in samples taken at eight sites in British Columbia on 23-25 May 1973 . . . . . . . . . . 63

XIII. Percentage parasitism of Coleophora laricella by individual parasite species in samples taken at 14 sites in British Columbia on 14-15 May 1974 . . . . . . . . . . 6 7

XIV. Percentage parasitism of Coieophora iari- cella by individual parasite species in samples taken at 14 sites in British

. . . . . . . . . . Colwnbiaon12-13June1974 69

LIST OF FIGURES

Figure Page

1. Distribution of Coleophora laricella in British Columbia and location of the eight 1973 and the 14 1974 collection sites. . . . . . . . . . . . . . . . . . 2

2. Emergence patterns af ~icladocePus spp. , Spilochalcis albifrons, and Mesopolobus sp. in relation to that of Coleophora laricella under a 12: 12 hour light-dark cycleat22OC . . . . . . . . . . . . . . . 2 4

3. Schematic representation of within-tree distributions of Coleophora laricella in one class of tree on 15 May 1974 and five classes of trees on 13 June 1974 at Shoreacres, British Columbia . . . . . . . . . . 3 2

4. Schematic representation of within-tree distributions of Dicladocerus spp. in five classes of trees on 13 June 1974 at Shoreacres, British Columbia . . . . . . . . . . 3 7

5. Schematic representation of within-tree distributions of Spilochalcis albifrons in five classes of trees on 13 June 1974 at Shoreacres, British Columbia . . . . . . . . . 41

INTRODUCTION

The l a rch casebearer , Coleophora l a r i c e l l a (Hbn.) (Lepidoptera:

Coleophoridae), is cur ren t ly the most se r ious d e f o l i a t o r of western l a rch ,

Larix occ iden ta l i s Nutt. in western North America. It was f i r s t d is -

covered i n t h e P a c i f i c Northwest near Ste. Maries, Idaho, i n 1957 (Denton

1958), and apparently entered B r i t i s h Columbia some t i m e before 1966 from

i n f e s t a t i o n s i n Washington, Idaho, and Montana (Dawson 1971). Its p resen t

d i s t r i b u t i o n i n B r i t i s h Columbia (Fig. 1) is along the i n t e r n a t i o n a l

border from Anarchist Summit t o Roosvil le and nor th t o the Cranbrook,

Lardeau, and Nelson areas . Most de fo l i a t ion is confined t o s tands below

an e leva t ion of 3,000 f e e t (Morris and Monts 1972) . C. l a r i c e l l a is univol t ine i n western North America (Dawson 1971;

Denton and Tunnock 1972). Moths emerge from l a t e May t o e a r l y Ju ly .

After mating, the female moth deposi t s about 50 eggs, usual ly s ing ly , on

the undersides of needles. Two weeks t o a month l a t e r , t he l a rva hatches

and bores through t h e eggshell and d i r e c t l y i n t o t h e needle, where it

feeds f o r one t o two months a s a needle miner ( i n s t a r s 1-111). The t h i r d -

i n s t a r l a r v a cons t ructs a case by l i n i n g t h e mined needle sec t ion with

s i l k and chewing t h i s sec t ion f r e e from t h e r e s t o f the needle, leaving

both ends open. Bearing the case , t h e l a r v a feeds ex te rna l ly upon f o l i -

age from mid-August onwards. I n October t h e t h i r d - i n s t a r l a r v a leaves

t h e fo l i age , a t t aches i t s case t o a twig o r branch, and overwinters in-

s i d e t h e case. The l a rva resumes feeding with t h e appearance of new

1

Fig. 1. Dis t r ibu t ion of Coleophora laricella i n B r i t i s h Columbia (1972) and loca t ion of t h e e i g h t 1973 and the 1 4 1974 c o l l e c t i o n sites (adapted f r o m Shepherd and Ross unpubl . * )

* R. F. Shepherd and D. A. Ross, P a c i f i c Fores t Research Centre,

Vic tor ia , B.C.

Scale : miles 0 30 -

Range of Larix occidentalis

Range of Coleoohora

i

COLLECTION SITES I ' I

I I i 1 A N A R c H m s u M M r r r. , J s - w M t A w .

? 2 9WtNSTOME CREEK PARK 9 KOOTENAY BAY

v 3 CASCADE 40 ARROW CREEK

'4 4 ROSSLAND $I1 RYKERTS

/ @ 5 FRUITVALE 4 2 YAHK

b 6 SHEEP'S CREEK /13 .CRANEROOK - J 7 SHOREACRES & $14 -RQQSVCCLE

fo l i age , usually i n mid t o l a t e Apri l . I n May t o ea r ly June t h e fourth-

i n s t a r l a rva a t t aches t h e case t o a needle o r twig and pupates. Length of

the pupal period is about one month.

Defoliat ion of l a r c h r e s u l t s i n growth l o s s and i n p red i spos i t ion

t o a t t a c k by o the r organisms (Tunnock e t a l . 1969).

Introductions of exo t i c p a r a s i t e s have begun i n t h e western

United S t a t e s and B r i t i s h Columbia, i n an attempt t o con t ro l C. l a r i c e l l a

b io log ica l ly (Denton 1972; Morris and Monts 1972; Ryan and Denton 1993;

Ryan e t a l . 1975). Turnbull and Chant (1961) argued t h a t t h e ecology of

a p e s t being considered a s a t a r g e t f o r a b io log ica l con t ro l program

should be s tud ied i n t h e a r e a of proposed r e l e a s e p r i o r t o the introduc-

t i o n of exo t i c na tu ra l enemies. The occurrence of na t ive p a r a s i t e s of

C. l a r i c e l l a i n B r i t i s h Columbia has not been w e l l documented although

Andrews and Geis t l inge r (1969) reared nine species o f p a r a s i t e s from

small rear ings of casebearers from 1966 t o 1968. The ob jec t ive of t h e

present work was t o determine t h e incidence of na t ive p a r a s i t e s o f C.

l a r i c d l a over i t s range i n B r i t i s h Columbia, as a prelude t o the in t ro-

duction and establishment of e x o t i c p a r a s i t e s .

CENERAL SURVEY OF PARASITES OF COLEOPHORA LARICEL-

I N BRITISH COLUMBIA

Methods and Materials

Survey f o r P a r a s i t e s of Mature Larvae and Pupae

Mass Rearings

I n 1973 c o l l e c t i o n s were made on 8-9 May and 23-25 May, t h e for-

m e r c o l l e c t i o n c o n s i s t i n g mainly of C. laricella four th - ins ta r l a rvae

and t h e l a t t e r o f pupae. Samples were taken a t e i g h t s i t e s : Anarchist

Summit, A r r o w Creek, Cascade, F ru i tva le , Rossland, Sheep's Creek cutoff

( 12 m i l e s south o f Salmo) , Shoreacres, and Yahk (Fig. 1).

A t eacl. site 10 trees were sampled a t the crown l e v e l s 4-6 f t .

(1.22-1.83 m) and 10-12 E t . (3.05-3.66 m) above t h e ground during each

co l l ec t ion . Five primary branches were taken from a l l s i d e s of each

tree i n each crown l e v e l and then c u t i n t o 1-1% f t . (30.5-45.8 cm) sec-

t ions . The samples taken on 8-9 May were placed i n p l a s t i c bags and

t r a n s f e r r e d t o r e a r i n g cages 24-48 hours l a t e r , whereas t h e samples taken

on 23-25 May were placed d i r e c t l y i n t o r e a r i n g cages i n t h e f i e l d , before

t r anspor t t o t h e laboratory.

Two types of cages w e r e used for mass-rearing of t h e samples.

One type was a s tandard 1' x 1' x 2 ' (30.5 x 30.5 x 61.0 cm) i n s e c t cage,

cons i s t ing of a wooden frame and bottom with mesh on a l l s ides and top.

he o the r type was a 1' x 1' x 1' (30.5 x 30.5 x 30.5 cm) corrugated

cardboard box with t h e top and one s i d e removed and covered with 0.2 mm

mesh. 5

The i n s e c t s were reared i n a labora tory i n which temperature and

humidity f luc tua t ions were recorded. Emerged moths and p a r a s i t e s were

c o l l e c t e d d a i l y and placed d i r e c t l y i n t o 70 percent ethanol .

The number of casebearers i n each sample was determined a f t e r

emergence was completed by manually removing the cases from t h e rea r ing

cages. Casebearer dens i ty i n each sample was determined by d iv id ing t h e

number of f a s c i c l e s ( i . e . , needle c l u s t e r s ) i n t o t h e number of cases

found. The mean number o f f a s c i c l e s pe r inch o f branch was ca lcu la ted

from 100 branch sec t ions from each co l l ec t ion . When emergence of moths

and p a r a s i t e s had ceased, lengths of a l l of t h e branch sec t ions i n a

sample were measured and mul t ip l ied by t h e previously determined r a t i o

of branch length: number of f a s c i c l e s i n each sample.

Unemerged p a r a s i t e s were detec ted by immersing a l l cases i n warm

10 percent KOH f o r 15 minutes, and then examining under a d i s s e c t i n g

microscope.

The data from the two height i n t e r v a l s were combined i n t o one

group. There were no obvious d i f fe rences i n e i t h e r casebearer dens i ty

o r t o t a l percentage paras i t i sm; however, some p a r a s i t e species occurred

only i n one crown l e v e l (Mil ler and Finlayson 1974).

I n 1974 t h e number of survey s i t e s was increased from 8 t o 14 by

adding t h e Cranbrook, Johnstone Creek Park , Kootenay Bay, ~ o o s v i l l e ,

Rykerts, and Winlaw s i t e s (Fig. 1) . Col lec t ions were made a t t h e sites

on 14-15 May (casebearer la rvae) and 12-13 June (casebearer pupae).

The crown l e v e l s i n 1974 were 5-10 ft. (1.53-3.05 m) and 20-25 f t .

(6.10-7.63 m) above t h e ground. The higher samples were obtained with a

pole pruner. Mass-rearings were done i n I' x 2 ' x 1' (30.5 x 61 x 30.5

cm) cardboard boxes, the t o p of which had been replaced by 0.2 mm mesh.

Otherwise, handling and rea r ing was t h e same a s f o r t h e 23-25 May 1973

co l l ec t ion .

Individual Rearings

A t o t a l of 18,300 l a rvae and pupae of C. l a r i c e l l a were indi -

v idual ly reared from samples taken a t the Rossland and Shoreacres s i t e s

on 30 Apr i l t o 1 May 1974; a t t h e Cascade, Rosslan,d, Sheep's Creek, and

Shoreacres sites on 14-15 May 1974; and a t t h e Rossland, Sheep's Creek,

and Shoreacres s i t e s on 29-30 May 1974.

The sample taken a t each s i t e cons is ted of f i v e branches per

he igh t i n t e r v a l from each of t e n t r e e s , encompassing the f u l l circum-

ference of each t r e e . The samples were taken from 5-10 f t . a t a l l s i t e s

during a l l c o l l e c t i o n s , and from 20-25 f t . a t t h e Sheep's Creek and

Shoreacres sites during t h e 14-15 May c o l l e c t i o n and a t the Shoreacres

s i t e during t h e 29-30 May co l l ec t ion . These samples were t ranspor ted t o

t h e labora tory i n p l a s t i c bags.

The casebearers were removed manually from t h e branches and

placed i n %-dram s h e l l v i a l s , one casebearer per v i a l , with f r e sh l a r c h - needles. The open end of t h e v i a l was then plugged with cot ton bat ten .

Dried l a r c h needles were pe r iod ica l ly replaced with f r e sh ones. Rearing

was done under a 12:12 hour l ight -dark cycle a t a temperature of about

22.OC (21-2S•‹C). Moth and p a r a s i t e emergences were recorded da i ly .

Af te r emergence had ceased, cases were d i s sec ted t o determine

the host stage from which each parasite had emerged. Occurrence o f hyper-

parasitism was determined by examining the remains within 25 cases that

had produced each of Dicladocerus spp., Mesopolobus sp., and S. albifrons.

Survey f or-Eg_s Parasite-%

C. laricella eggs were collected from 10 trees at each of the

eight 1973 and 14 1974 sites. The 1973 samples were taken on 21-22 July

and the 1974 samples on 29-30 July. The terminal 6" (15.3 cm) of 12

secondary or tertiary branches was taken from all sides of each tree from

the same crown levels as the late-instar larval-pupal collections. These

samples were mass-reared in 15 x 35 mrn petri dishes.

Parasite emergence was checked daily for six weeks, after which

the eggs were counted but the egg density was not estimated.

Early larval instars of C. laricella were collected at the Ross-

land and Shoreacres sites in 1973 on 21 August (needle-mining larvae)

and 7 October (casebearing larvae). Collection, handling, and rearing

techniques were the same as in the 23-25 May 1973 collection.

All of the rearing cages were the cardboard-box type. The

samples were checked daily for parasite emergence for four weeks. C.

laricella density was calculated only for the 7 October samples. The

number of casebearers in the 21 August samples was determined by dissec-

tion of all needles which appeared to contain mines and in the 7 October

samples by manually removing the cases.

Results ~ n $ - D ~ s c u s s ion

Densit ies -- of Coleophora l a r i c e l l a i n B r i t i s h Columbia -----

A t o t a l of 134,501 four th - ins ta r la rvae and pupae of C. l a r i c e l l a

were mass-reared i n t h i s s tudy. The numbers reared and t h e casebearer

d e n s i t i e s i n samples from each s i t e a r e l i s t e d i n Table I. I n add i t ion ,

2,427 eggs, 19,279 needle-mining larvae , and 6,890 casebearing th i rd -

i n s t a r l a rvae were reared.

C. l a r i c e l l a d e n s i t i e s decl ined over t h e per iod of t h e study.

The primary cause o f t h e dec l ine may have been the severe drought, o r

high temperatures associa ted with it, t h a t occurred i n t h e summer of 1973.

The d e n s i t i e s o f t h e casebearer a t Rossland and Shoreacres on 7 October

were 13.6 and 46.7 pe r 100 f a s c i c l e s , r e spec t ive ly , compared t o 66.7

and 112.8 per 100 f a s c i c l e s on 25 May. Desiccation o f eggs and espec ia l ly

t h e needle-mining larvae , the s t ages which occurred during the drought,

is known t o be an important mor ta l i ty f a c t o r (Eidmann 1965; Jasch 1973).

Other important mor ta l i ty f a c t o r s known t o reduce C. l a r i c e l l a popula-

t i o n s include unfavorable winter and sp r ing weather, i n t r a s p e c i f i c compe-

t i t i o n of needle-mining larvae , b i r d predation during winter , and poor

synchronization of la rvae emerging from diapause with budding of t h e

l a r c h trees (Eidmann 1965; Jasch 1973; Sloan 1965; Webb 1953).

P a r a s i t e s of Coleophora l a r i c e l l a i n B r i t i s h Columbia

P a r a s i t e s of Mature Larvae and Pupae

Species Obtained

A t o t a l of 42 species of hymenopterous p a r a s i t e s were obtained

m r- . . 0 0

C O O d N

cy t. . . 0 0

0-l d rl m

0 N . . 4 0 CJ f-i

rl m a, d' 0 w . 4

r- '3' . . CJ 03 m rl

d ' o r l c n 0 In . rl

a,

N rl d

from the niass-reari .ngs of fourth-instar larvae and pupae of C. laricella

(Table 11). In 1973 and 1974, 32 and 25 species, respectively, were

taken; 15 being common to both years.

As the samples were mass-reared, some of the parasites could have

in fact emerged from hosts other than C. laricella that were accidentally

included in the rearings. A mass-reared parasite was considered a larch

casebearer parasite if the species emerged in individual rearings done to

verify host relationships in this study or in others (Bousfield and Load

1973; Denton 1972; Sloan 1965; Webb 1953). Thirty of the species in 24

genera were considered parasites of C. laricella (Table 11).

Most of the species represent new host records for British

Columbia. Four of these were previously recorded by Andrews and Geist-

linger (19691, namely: Gelis tenellus (Say), Scambus decorus Wly., Tetra-

stichus ecus [=xanthops (Ratz.)], and Spilochalcis albifrons (Walsh) . They also obtained Bracon sp. which may well have been B. pygmaeus Prov.,

Amblymerus sp. which may have been the same species as lfesoplobus sp.

in this study, and Dicladocerus westwoodii Westw. which may be one of

the species obtained in this study; as well as Scambus transgressus

(Holmg.) and Sceptrothelys deione (Wlkr.) which were not taken in this

study. The average percentage parasitism of 3,245 casebearers sampled

by Andrews and Geistlinger was less than 1 percent in 1966 and 1967, and

4 percent in 1968. The highest percentage parasitism at an individual

site was 14 percent at Creston in 1968.

Several species excluded as C. laricella parasites could in fact

be parasites of C. laricella. Species of Acrolyta and Hypsoter have

Tab

le 11.

Pa

ra

sit

es o

bta

ine

d

fro

m m

ass-r

eari

ng

s

of

matu

re la

rva

e a

nd

p

up

ae

of

Co

leo

ph

ora

la

ric

ell

a i

n 1

97

3 a

nd

1974 c

oll

ec

tio

ns i

n B

rit

ish

C

olu

mbia

Hym

enopte

rous

Pa

ra

sit

es

Ab

un

da

nc

e

19

73

1

97

4

8-9

M

ay

23-2

5

May

14-1

5

May

1

2-1

3 June

- N

o.

No.

N

o.

No.

N

o.

Sp

ecim

en

s N

o.

Sp

ecim

en

s N

o.

No.

S

ite

s

Sit

es

S

pecim

ens

Specim

ens

Ob

tain

ed

O

bta

ined

O

bta

ined

S

ite

s

Ob

tain

ed

S

ite

s

Bra

co

nid

ae

P

Ap

hid

ius

sp

. 1

1

*B

raco

n p

ygm

aeu

s P

rov

. a

,b,c

3

2 5

2 73

8

9

5

6

4

Ich

neu

mo

nid

ae

Ac

roly

ta

sp

.

*C

am

po

ple

x r

ufi

pe

s P

rov

.

*Dia

degm

a sp

. a

, c

"G

eli

s t

en

ell

us

(S

ay

) a,b

rc

*G

eli

s sp

. a

,b,c

Hy

ps

ote

r s

p.

a

*It

op

l ec

tis v

esc

a T

ow

nes

*P

rist

om

eru

s sp

. a

,b

*S

cam

bu

s d

ec

oru

s w

all

ey

a

Mym

arid

ae

An

ap

hes

sp

.

Tab

le

11

--C

on

tin

ued

Hym

enopte

rous

Pa

rasit

es

Ab

un

da

nc

e

19

73

1

97

4

8-9

M

ay

23-2

5

May

14-1

5

May

12-1

3 June

No.

N

o.

No.

N

o.

No.

N

o.

No.

N

o.

Sp

ecim

en

s S

pecim

en

s S

pecim

en

s S

pecim

ens

Sit

es

O

bta

ined

O

bta

ined

S

ite

s

Sit

es

S

ite

s

Ob

tain

ed

O

bta

ined

Eu

lop

hid

ae

Ap

ros to

cet u

s sp

p .

( 2 )

*C

hry

soch

ari

s (K

ra to

ch

vi 1

ian

a)

lar

icin

ell

ae

(R

atz

. ) a.c

C

"C

irro

spil

us

pic

tus

(Nees

*D

icla

do

ceru

s sp

p.

(3) a

,b,c

~ig

lyp

hu

s sp

. C

"E

lac

he

rtu

s p

rote

ote

rati

s (H

ow. )

C

*

Eu

der

us

cush

ma

ni

(Cra

w fo

rd)

*E

ulo

ph

us

sp.

c,d

Me

litt

ob

ia sp

.

*T

etr

ast

ich

us

do

losu

s (G

ahan) b

*T

etr

ast

ich

us

ec

us

W

alk

er a

,b

*Zag

ram

mos

oma

am

eric

an

um

Gir

. b

Th

ysa

nid

ae

Th

ysa

nu

s sp

.

Tab

le

11

--C

on

tin

ued

Ab

un

da

nc

e

19

73

1

97

4

8-9

M

ay

23

-25

M

ay

14

-15

M

ay

12

-13

Ju

ne

Hy

me

no

pte

rou

s P

ara

sit

es

NO

. N

o.

No

. N

o.

No

. S

pe

cim

en

s

No

. S

pe

cim

en

s

No

. N

O.

Sit

es

Sp

ec

ime

ns

S

ite

s

Sp

ec

ime

ns

O

bta

ine

d

Ob

tain

ed

O

bta

ine

d

Sit

es

Ob

tain

ed

S

ite

s

En

cy

rti

da

e

C

*C

op

i dos

oma

sp

. P

tero

ma

l id

ae

*C

ato

lac

cu

s a

en

eo

vir

idis

(G

ir.

1 b,c

Cy

rto

ga

ste

r v

ulg

ar

is W

alk

er

*H

ab

rocy

t us

ph

yc

i d

is A

s hm

. b

~c

*M

eso

po

l ob

us

sp

. arb

Ch

alc

idid

ae

i

*S

pil

oc

ha

lcis

alb

i fro

ns

(Wa

lsh

) a

&,c

Eu

ryto

mid

ae

"Eu

ryto

ma

sp

. c ,d

Ce

rap

hro

nid

ae

Aph

anog

mus

sp

.

been reared from another coleophorid, Coleophora rna l ivore l la m y . (Beacher

1947). Diglyphus sp. was obtained i n s u f f i c i e n t l y l a r g e numbers t o indi -

c a t e t h a t it d id emerge from C . l a r i c e l l a .

Introduced Species

Two of t h e species , Chrysochar i s l a r i c i n e l l a e ( ~ a t z . ) and Cirro-

s p i l u s p i c t u s ( ~ e e s ) , a r e European species t h a t were re leased aga ins t C.

l a r i c e l l a i n eas te rn North ~ m e r i c a i n the l a t e 1930's (Dowden 1962;

McGugan and Coppel 1962). C . l a r i c i n e l l a e was re leased agains t C. l a r i -

c e l l a i n t h e western United S t a t e s (Idaho and Washington) i n 1972 (Ryan

and Denton 1973) bu t was probably present i n western North America before

1972 as specimens were reared from C . l a r i c e l l a long d is tances from t h e

re l ease po in t s i n the same year (Ryan e t a l . 1974). The c l o s e s t r e l e a s e

point was over 200 miles from the s i t e s i n B r i t i s h Columbia where speci-

mens were c o l l e c t e d , a long d i s t ance t o d i spe r se i n one season.

Poss ib le explanations f o r the presence of C. L a r i c i n e l l a e i n t h e

P a c i f i c Northwest a r e : (1) it has spread from eas te rn North America on

a l t e r n a t i v e hos ts ; ( 2 ) it was introduced with C . l a r i c e l l a and spread

with it; o r ( 3 ) it was introduced with e a r l y in t roduct ions of A g a t h i s

pumila (Ratz . ) , a European species previously re leased aga ins t C. l a r i -

c e l l a i n western North America (Ryan e t a l . 1974). These explanations

could a l s o apply t o C. p i c t u s .

A. pumila is conspicuous by i t s absence i n t h i s study. This

species was re leased i n B r i t i s h Columbia i n 1969 a t two s i t e s , F ru i tva le

and Arrow Creek (not a t s i t e s of the same names i n t h i s s tudy, although

the Arrow Creek s i t e s were l e s s than one mile a p a r t ) and became

es tab l i shed a t both (Morris and Monts 1972). The absence of A. pumila

a t t h e F ru i tva le and Arrow Creek s i t e s of t h i s study ind ica tes t h e slow-

ness with which t h i s p a r a s i t e i s dispers ing . I n t h e western United

S t a t e s t h e most d i s t a n t recovery from a re l ease po in t was only s i x miles

10 years a f t e r r e l ease , possibly due t o the excessively l a r g e numbers of

C. l a r i c e l l a ava i l ab le i n t h e immediate area and consequent slow dis-

p e r s a l r a t e (Denton 1972).

Relat ive Importance of P a r a s i t e Species

Although many p a r a s i t e species were reared, only a few predomi-

nated. Of t h e seven species present i n t h e c o l l e c t i o n of 8-9 May 1973,

Dicladocerus spp. represented 86.7 percent and B. pygmaeus 8.5 percent

of t h e t o t a l number of p a r a s i t e s . Of the 26 species reared from the col-

l e c t i o n o f 23-25 May 1973, seven species cons t i tu ted 96.6 percent of t h e

t o t a l , namely: Dicladocerus spp. (46.0%), S . a l b i f r o n s (32.8%), B.

pygmaeus (8.5%) , Mesopolobus sp. (4.9%) , and T e t r a s t i c h u s e c u s

(4.4%). These species were a l s o t h e most widespread (Table 11). I n

1974, 11 species were taken i n t h e c o l l e c t i o n of 14-15 May with Dicla-

docerus spp. (83.2%), and Mesopolobus sp. (13.3%) c o n s t i t u t i n g 96.5 per-

cen t of the t o t a l number of p a r a s i t e s . I n t h e c o l l e c t i o n of 12-13 June,

17 species were reared, with Dic ladocerus spp. (63.8%), S . a l b i f r o n s

(23.5%), and Mesopolobus sp. (9.9%) c o n s t i t u t i n g 97.2 percent of the

t o t a l . These species were a l s o t h e most widespread i n 1974 (Table 11).

Paras i t i sm a t Individual S i t e s

The number of taxa and t h e t o t a l percentage paras i t i sm f o r each

s i t e and c o l l e c t i o n a r e summarized i n Table 111. Percentage pa ras i t i sm

by individual species a t each s i t e i n each c o l l e c t i o n a r e given i n

Appendix 11.

The percentage pa ras i t i sm and t h e number of taxa were g rea te r i n

t h e second c o l l e c t i o n than t h e f i r s t i n both years (Table 111). Bousfield

and Lood (1971) and Beacher (1947) found s i m i l a r increases i n pa ras i t i sm

of C. l a r i c e l l a and C. m a l i v o r e l l a , respect ive ly , during t h e sp r ing feed-

i n g per iod of t h e hos ts . Evidently, a c t i v e p a r a s i t i z a t i o n o f C. l a r i c e l l a

occurs during t h i s period, suggest ing t h a t p a r a s i t e a d u l t s had j u s t become

a c t i v e o r t h a t C. l a r i c e l l a reached a suscep t ib le s tage . Sweep-net col-

l e c t i o n s o f a d u l t s o f B. p y p a e u s , I t o p l e c t i s vesca Townes, Dicladocerus

spp., and Hesopolobus sp. during t h e f i r s t 1974 c o l l e c t i o n confirmed t h e

presence o f a d u l t p a r a s i t e s a t t h a t t i m e . I n Europe, G e l i s spp., D.

westwoodi i , and ,Vesopolobus subfumatus Ratz. a r e known t o a t t a c k C, l a r i -

c e l l a j u s t a f t e r t h e hos t la rvae have resumed feeding i n the spr ing , a s

probably do most o t h e r European p a r a s i t e species (Jasch 1973). B.

pygmaeus a t t a c k s Coleophora p run i e l l a C1. i n Wisconsin during t h e same

per iod (Doner 1934). The increase i n pa ras i t i sm by S. a l b i f r o n s between

c o l l e c t i o n s was probably cor re la t ed with t h e increase i n hos t pupal popu-

l a t i o n s which i s thought t o be t h e s t a g e a t tacked by t h i s species (Bous-

f i e l d and Lood 1971).

B. pygnaeus i s known t o have two generat ions pe r year on C.

p r u n i e l l a during t h e sp r ing (Doner 1934). I f t h i s species a l s o has two

on,C. l a r i c e l l a i n B r i t i s h Columbia, t h e a c t u a l percentage paras i t i sm

may d i f f e r from t h a t ind ica ted i n e i t h e r c o l l e c t i o n of each year , depend-

i n g on t h e amount of generat ion overlap (which was unknown).

m r - w

o r - m . . (U d ' r -

Ta

ble

1

11

--C

on

tin

ue

d

19

73

1

97

4

Cro

wn

8-9

M

ay

23

-25

M

ay

14

-15

M

ay

12

-13

Ju

ne

L

ev

el

No.

of

To

tal

%

No.

o

f T

ota

l %

N

o.

of

To

tal

%

No.

of

To

tal

%

Sit

e

(ft.

)

Tax

a*

Pa

ras

itis

m

Tax

a*

t P

ara

sit

ism

T

axa*

P

ara

sit

ism

T

axa*

P

ara

sit

ism

Ro

ss la

nd

5-

10

3

6.7

1

5

16

.9

4

9.3

3

27

.7

Sh

ee

p's

C

reek

5-1

0

3 P

7

2

.5

3

9.2

5

16

.8

Sh

ore

ac

res

5-1

0

3

1.4

1

6

6.3

6

1

1.9

8

18

.3

20

-25

3

1

0.2

3

1

3.8

Yah

k 5-1

0

1

5.6

2

2.1

1

1.9

1

6.9

20

-25

-

2.0

1

7.7

* On

ly

emer

ged

s

pe

cie

s c

ou

nte

d;

Dic

lad

oc

er

us

sp

p.

we

re t

re

ate

d a

s o

ne

sp

ec

ies

as

th

ey

co

uld

no

t be

sep

ara

ted

.

'Te

len

om

us

sp

p.

we

re c

ou

nte

d a

s o

ne

sp

ec

ies

.

-Un

emer

ged

o

nly

.

Aspects of Parasite Biology

Brood Size:

A c h r y s o c h a r e l l a sp. was the only gregarious parasite species indi-

cated by the individual rearings. The mean number produced from four

cases was 3.25 (range L - 5 ) . A l l individuals that emerged from one case I

were of the same sex. Bousfield and Lood (1973) also found a very low

incidence of gregarious parasites. However, these researchers found

three species, A c h r y s o c h a r e l l a s i l v i a G i r . , T . e c u s , and Mesopolobus sp.,

tha t occasionally produced more than one adult per case.

Hyperparasitism:

No evidence of hyperparasitism by D i c l a d o c e r u s spp. , Mesopolobus

sp., o r S. a l b i f r o n s was found in the dissected cases of C. l a r i c e l l a .

They have been recorded as hyperparasites, as well as primary parasites,

of other insects (Muesebeck e t a l . 1951; Peck 1963; Telford 1961). It

could not be determined i f any of the other species obtained were hyper-

parasites because of the small numbers present in the individual rearings.

Species taken during th i s study tha t have been recorded both as hyper-

parasites and primary parasites (Muesebeck e t a l . 1951; Peck 1963;

Telford 1961) include: G. t e n e l l u s , G e l i s sp., T . e c u s , Zagramnosom

a m r i c a n u m Wlkr., C a t o l a c c u s a e n e o v i r i d i s ( G i r . ) , and Habrocytus p h y c i d i s

Ashm, The l a t t e r species was found t o be both a primary and secondary

parasite on the same host, A c r o b a s i s r u b r i f a s c i e l l a Pack., by Finlayson

(1967).

Emergence Pat terns :

Only Dicladocerus spp. , S . a l b i f r o n s , and Mesopolobus sp. were

taken i n s u f f i c i e n t l y l a rge numbers i n individual rear ings t o permit t h e

p l o t t i n g of emergence curves (Fig. 2 ) . No c o r r e l a t i o n e x i s t e d between

p a r a s i t e emergence and temperature over t h e range observed. Dic ladocerus

spp. emergence began t h r e e days a f t e r t h a t o f C. l a r i c e l l a and l a s t e d 14

days, peaking on t h e s i x t h day. Mesopolobus sp. emergence began t h r e e

days a f t e r t h a t of Dicladocerus spp. and l a s t e d 12 days, peaking on the

f i f t h day. S. a l b i f r o n s emergence began 11 days a f t e r t h a t of Diclado-

c e r u s spp. and l a s t e d 10 days, peaking on the four th day.

The emergence p a t t e r n s o f Dic ladocerus spp. and S . a l b i f r o n s

d i f f e r from t h e p a t t e r n s reported by Bousfield and Lood (1971) . The

emergence began, peaked, and ended sooner i n t h e p resen t study. These

d i f ferences may have been the r e s u l t of d i f ferences i n r ea r ing condit ions,

which were no t reported by Bousfield and Lood (1971).

P a r a s i t e s of Eggs and Ear ly- Ins tar Larvae

No p a r a s i t e s were taken from mass-rearings of C. l a r i c e l l a eggs,

needle-mining l a rvae o r casebearing t h i r d - i n s t a r larvae. Sloan and

Coppel (1965) d i d not f ind any egg p a r a s i t e s i n Wisconsin and none have

been reported elsewhere. Whether t h e l ack of r epor t s of ea r ly - ins ta r

l a r v a l p a r a s i t e s i n o t h e r areas of North America is due t o the lack of

p a r a s i t e s o r d e f i n i t i v e research i s not known.

Fig. 2. Emergence p a t t e r n s of Dicladocerus spp. , S p i l o - c h a l c i s a l b i f r o n s , and Mesopolobus sp. i n r e l a t i o n t o t h a t of Coleophora l a r i c e l l a under a 12:12 hour l ight-dark cycle a t 22OC. The po in t s represent t h e da i ly ( i n r e l a - t i o n t o casebearer emergence which began 11-12 June) aver- ages of t h e samples co l l ec ted on 29-30 May 1974 and reared individual ly . Day 1 was t h e f i r s t day of casebearer emergence i n the sample from a s i t e o r he ight

IL- C. laricella 11-11 Dicladocerus spp. -I- S. albifrons mle..-.-l.... Mesopolobus sp.

0 4 8 12 16 20 24

DAYS

DISTRIBUTIONS OF COLEOPHORA LARICELLA AND ITS PARASITES

I N WESTERN LARCH CROWNS

Methods and Materials

The d i s t r i b u t i o n s of C. l a r i c e l l a and t h e p a r a s i t e s Dic ladocerus

spp. and S . a l b i f r o n s i n crowns of western l a r c h were examined on 15 May

and 13 June 1974 a t Shoreacres. A t o t a l of 40 t r e e s were sampled from

f i v e c l a s s e s of t r e e s during t h e 1 3 June co l l ec t ion . The f i v e c l a s s e s

and t h e number o f t r e e s sampled i n each c l a s s were:

Class Description No. of Trees

1 Open-grown t r e e s a t l e a s t 100 yd. from road and over 40' high

10

2 Same a s c l a s s 1 except 25-35' high 10

3 >

Same a s c l a s s 1 except 10-15' high 5

4 Same a s c l a s s 1 except t r e e s were road- s i d e

Same a s c l a s s 1 except t r e e s formed

5 closed canopy. Trees sampled were a t l e a s t twice t h e he ight of the t r e e s

10

from t h e edge o f the stand.

During t h e 15 May c o l l e c t i o n only c l a s s 1 t r e e s were sampled.

Samples were taken a t two crown l e v e l s , 5-10 f t . and 20-25 f t .

above t h e ground. Two primary branches were taken from both t h e sunny

and shaded s i d e s of each t r e e from each crown l e v e l and c u t i n h a l f .

The branch halves were mass-reared i n p a i r s according t o t r e e , crown

l e v e l , s i d e of t r e e , and branch h a l f . Otherwise these samples were

handled and reared i n t h e same manner a s t h e mass-reared 1974 general

survey samples.

2 7

For s t a t i s t i c a l a n a l y s i s l o g X t ransformat ions were c a r r i e d 10

o u t on l a r c h casebearer d e n s i t i e s (no./100 f a s c i c l e s ) and a r c s i n e t r ans -

formations were done on percentage p a r a s i t i s m da ta because var iances were

no t independent of means i n t h e untransformed da t a , var iances i nc reas ing

wi th means. The t ransformat ions made t h e var iances independent. I n

a n a l y s i s of var iance (Dixon 1973) of t h e i n t r a - t r e e d i s t r i b u t i o n s of each

c l a s s , t r e e s were allowed t o go random, r e s u l t i n g i n conserva t ive F

va lues and genera l ized s ta tements o f da t a . The d a t a a r e given i n t h e

unt rans formed form.

Resul t s and Discussion

The mean dens i ty of C. l a r i c e l l a and t h e mean percentage para-

s i t i s m by Dicladocerus spp. and by S. a l b i f r o n s i n t h e crown l e v e l s o f

each t r e e c l a s s a r e l i s t e d i n Table I V . There were no s i g n i f i c a n t d i f -

fe rences i n h o s t d e n s i t y o r i n p a r a s i t i s m by e i t h e r spec i e s between t h e

f i v e c l a s s e s (Tables V - V I I ) .

The mean d e n s i t i e s of C. l a r i c e l l a i n t h e var ious po r t ions of

t h e t r e e crown a r e i l l u s t r a t e d i n Fig. 3. Analysis of var iance showed

a h ighly s i g n i f i c a n t v a r i a t i o n i n C. l a r i c e l l a dens i ty between crown

l e v e l s , between s i d e s o f t h e t r e e , and between branch ha lves i n c l a s s 1

c o l l e c t i o n s (Table V I I I ) . Density was s i g n i f i c a n t l y higher a t t h e lower

crown l e v e l than a t t h e h igher l e v e l ; on t h e sunny s i d e of t r e e s than on

t h e shaded s i d e ; and on t h e o u t e r h a l f of t h e branch than on t h e i n n e r

h a l f . S i m i l a r d i s t r i b u t i o n s i n c l a s s e s 2 and 4 a l s o proved t o be s i g n i f -

i c a n t (Fig. 3; Table V I I I ) . I n c l a s s 3 , s i g n i f i c a n t v a r i a t i o n occurred

Table I V . Density of Coleophora l a r i c e l l a and percentage parasitism by Dicladocerbls spp. , and by S p i l o c h a l c i s a l b i f r o n s i n five classes of t rees on 13 June 1974 a t Shoreacres, Bri t ish Columbia. (X=mean, SD=standard devi- ation)

- -- - - ---

Crown C. l a r i c e l l a density % Parasitism Level (no. /lo0 fascicles) Dicladocerus spp. S . a l b i f r o n s - - -

Class (ft. ) X SD X S D X S D

Table V. Analysis of variance for the distribution of Coleophora Laricella in five classes of trees on 13 June 1974 at Shoreacres, British columdia. (DF=degrees of freedom, MS=mean square, F=F ratio, LS=level of significance)

C r o w n L e v e l 5-10 ft. 20-25 ft.

Source of Variation DF MS F LS DF MS F LS

Between Classes 4 .0627 5.22 - 3 -0255 2.57 -

Within Classes

Total 3 9 3 4

- not significant

Table VI. Analysis of variance for the distribution of Dicladocerus spp. in five classes of trees on 13 June 1974 at Shoreacres, British Columbia. (DF=degrees of freedom, MS=mean square, F=F ratio, LSzlevel of significance)

C r o w n L e v e l 5-10 ft. 20-25 ft.


Between Classes 4 16.890 0.81 - 3 7.499 0.53 -

Within Classes

Total 39 3 4

- not significant

Table VII. Analysis of variance for the distribution of Spilochalcis albifrons in five classes of trees on 13 June 1974 at Shoreacres, British Columbia. (DF=degrees of freedom, MS=mean square, F=F ratio, LS=level of significance)

C r o w n L e v e l 5-10 ft. 20-25 ft.


Between Classes 4 108.958 5.39 - 3 51.452 1.31 -

Within Classes

Total 39 3 4

- not significant

Fig. 3. Schematic representation of within-tree distributions of Coleophora laricella in one class of tree on 15 May 1974 and five classes of trees on 13 June 1974 at Shoreacres, British Columbia. (Numbers represent number of casebearers per 100 fascicles, with outer being those of outer branch half and inner those of inner branch half .)

1 May 15, 1974 collection

-

1 June 13, 1974

~ u n e 1 3, 1974 collection

I I I

2 collection

Ta

ble

VI

II

. A

na

lys

is o

f v

ari

an

ce

fo

r t

he

wit

hin

-tre

e

dis

trib

uti

on

s o

f C

ole

o-

phor

a la

ric

ell

a i

n o

ne

cla

ss

of

tre

e o

n

15

May

a

nd

f

ive

cla

sses o

f tr

ee

s o

n

13

Ju

ne

1

97

4 a

t S

ho

rea

cre

s,

Brit

ish

Co

lum

bia

. (D

F=

deg

rees

of

fre

ed

om

, M

S=

mea

n

sq

ua

re,

F=

F

ra

tio

, L

S=

lev

el

of

sig

nif

ica

nc

e)

Ma

y

15

J

un

e

13

-

So

urc

e o

f C

la

ss

1

Cl

as

s

1

Cl

as

s

2

Va

ria

tio

n

Err

orT

erm

D

F

MS

F

LS

DF

M

S F

LS

DF

MS

F

LS

Mai

n Effects

He

igh

t H

t-T

r 1

2.1

95

1

14

.46

**

* 1

1.7

86

2

17

.69

**

* 1

2.3

28

1

68

.03

**

* S

ide

S

d-T

r 1

0.1

53

2

2.6

6

***

1

0.1

41

1

7.3

6

***

1

0.2

07

2

6.9

9

***

Ha

lf

Hf -

Tr

1

0.2

81

7

8.5

8

***

1

0.2

55

3

8.6

5

***

1

0.6

60

5

9.2

8

***

Inte

rac

tio

ns

Ht-

Sd

H

t-S

d-T

r 1

0.0

05

0

.62

-

1

0.0

11

1

.44

-

1

0.0

17

1

.87

-

Ht-

Hf

Ht-

Hf-

Tr

1

0.0

15

2

.88

-

1

0.0

04

0

.59

-

1

0.0

64

1

0.9

0

x

Sd-

Hf

Sd-H

f -T

r 1

0.0

31

2

.46

-

1

0.0

44

4

.13

-

1

0.0

03

0

.50

-

Ht-

Tr

- 9

0

.01

9

9

0.0

08

9

0

.01

4

Sd

-Tr

- 9

0

.00

7

9

0.0

08

9

0

.00

8

Hf -

Tr

- 9

0

.00

4

9

0.0

07

9

0.0

11

Ht-

Sd-H

f H

t-S

d-H

f-T

r 1

0.0

14

2

.11

-

1

0.0

08

1

.06

-

1

0.0

10

6

.59

*

Ht-

Sd

-Tr

- H

t-H

f -T

r -

Sd-H

f -T

r -

Ht-

Sd-H

f -T

r -

9

0.0

07

9

0.0

08

9

0

.00

1

between branch halves but not between sides of trees (Fig. 3; Table VIII).

Density was higher on the outer branch half than on the inner half. In

class 5, significant variation occurred between branch halves but not be-

tween crown levels or sides of trees (Fig. 3; Table VIII). The density

was higher on the outer branch half than on the inner half.

Webb (1953) found a similar distribution of C. l a r i c e l l a with

respect to crown level and branch portion. The density at the top of

the crown was about 2/3 that at the base, and more casebearers were on

the terminal portion than the base of the branch. The abundance of C.

l a r i c e l l a larvae and pupae on the sunny side of the tree and the outer

half of the branch may reflect the oviposition behavior of the female

moth (Sloan and Coppel 1965; Webb 1953).

Parasitism by Dicladocerus spp. in class 1 was significantly

higher on the inner than on the outer branch half whereas there were no

significant variations between crown levels or sides of trees (Fig. 4;

Table IX). Similar significant and non-significant variations occurred

in class 2 (Fig. 4; Table IX). In classes 3, 4, and 5, no significant

variations occurred between crown levels, sides of trees or branch halves

(Table IX) . Parasitism by S. a l b i f r o n s was significantly greater at the lower

than at the higher crown level and on the outer than on the inner branch

half, whereas no significant variation occurred between tree sides in

class 1 (Fig. 5; Table X). A similar distribution occurred in class 2

(Fig. 5; Table X). No significant variations occurred between tree sides

or branch halves in classes 3 or 4, as well as between crown levels in

Fig. 4. Schematic representation of within-tree distributions of Dicladocerus spp. in five classes of trees on 13 June 1974 at Shoreacres, British Columbia. (Numbers represent percentage parasitism, with outer being those of outer branch half and inner those of inner branch half)

Tak d

e IX. Analysis of variance for the within-tree distributions of Dicladocerus

spp. in five classes of trees on 13 June 1974 at Shoreacres, British Columbia.

(DF=degrees of freedom, MS=mean square, F

=F

ratio, LS=level of significance)

Source of

Class 1

Class 2

Class

3

Variation

Error Term

DF

MS

F

LS

DF

MS

F

LS

DF

MS

F

LS

Main Effects

Height

Side

Half

Interactions

Ht-Sd

Ht-Hf

Sd-Hf

Ht-Tr

Sd-Tr

Hf -Tr

Ht-Sd-Hf

Ht-Sd-Tr

Ht-Hf -Tr

Sd-Hf-Tr

Ht-Tr

Sd-Tr

Hf -Tr

Ht-Sd-Tr

Ht-Hf -w

Sd-Hf-Tr

- - - Ht-Sd-Hf-Tr

- - - Ht-Sd-Hf-Tr

-

Ta

ble

IX

--C

on

tin

ued

So

urc

e o

f V

ari

ati

on

E

rro

r T

erm

C

la

ss

4

Cl

as

s

5

DF

MS

F

LS

DF

MS

F

LS

Mai

n E

ffe

cts

Heig

ht

Ht-

Tr

Sid

e

Sd-T

r

Half

H

f -T

r

Inte

rac

tio

ns

Ht-

Sd

Ht-

Sd-T

r

Ht-

Hf

Ht-

Hf-

Tr

Sd-

Hf

Sd-

Hf

-Tr

Ht-

Tr

- S

d- T

r -

Hf-

Tr

- H

t-S

d-H

f H

t-S

d-H

f-T

r

Ht-

Sd-T

r -

Ht-

Hf

-Tr

- S

d-H

f -T

r -

Ht-

Sd-H

f -T

r -

- n

ot

sig

nif

ica

nt;

*

P<

0.0

5;

***

P<

0.0

01

; W

0.0

1

Fig. 5. Schematic representation of within-tree distributions of Spilochalcis albifrons in five classes of trees on 13 June 1974 at Shoreacres, British Columbia. (Numbers represent percentage parasitism, with outer being those of outer branch half and inner those of inner branch half)

Tab

le

X.

Anal

.ysis

of

va

ria

nc

e fo

r t

he

wit

hin

-tre

e

dis

trib

uti

on

s o

f Spilochalcis

albifrons in

fiv

e c

las

se

s o

f tr

ee

s o

n

13

Ju

ne 1

97

4 a

t S

ho

rea

cre

s,

Brit

ish

C

olu

mbia

. (D

F=

deg

rees

of

freed

om

, M

S=m

ean

sq

ua

re,

F=

F ra

tio

, L

S=

lev

el

of

sig

nif

- ic

an

ce

)

So

urc

e o

f C

la

ss

1

Cl

as

s

2 C

la

ss

3

V

ari

ati

on

E

rro

r T

erm

DF

MS

F

LS

DF

MS

F

LS

DF

MS

F

LS

Main Effects

Heig

ht

Ht-

Tr

Sid

e

Sd-T

r

Half

H

f -T

r

Interactions

Ht-

Sd

Ht-

Sd-T

r

Ht-

Hf

Ht-H

f -T

r

Sd-H

f S

d-H

f -T

r

Ht-

Tr

- S

d-T

r -

Hf-

Tr

- H

t-S

d-H

f H

t-S

d-H

f-T

r

Ht-

Sd-T

r -

Ht-

Hf-

Tr

- S

d-H

f-T

r -

Ht-

Sd-H

f -T

r -

Ta

ble

X--

Co

nti

nu

ed

So

urc

e o

f V

ar

iati

on

E

rr

or

Te

rm

Cl

as

s

4 C

la

ss

5

DF

MS

F

LS

D

F M

S F

L

S

Ma

in E

ffects

He

igh

t H

t-T

r

Sid

e

Sd

-Tr

Ha

lf

Hf -

Tr

Inte

rac

tio

ns

Ht-

Sd

H

t-S

d-T

r

Ht-

Hf

Ht-

Hf-

Tr

Sd-

Hf

Sd

-Hf-

Tr

Ht-

Tr

- S

d-T

r -

Hf-

Tr

- H

t-S

d-H

f H

t-S

d-H

f-T

r

Ht-

Sd

-Tr

- H

t-H

f-T

r -

Sd-

Hf

-Tr

- H

t-S

d-H

f -T

r -

- n

ot

sig

nif

ica

nt;

*

PC

0.0

5;

** P

<0

.01

; **

* P

<0

.00

1;

N<

0.0

1

class 4. In class 5, significantly greater parasitism occurred at the

lower than at the higher crown level but no significant variations

occurred between tree sides or branch halves (Fig. 5; Table X).

The distribution of Dicladocerus spp. could be affected by move-

ments of C. laricella larvae during the period of parasite attack and

development. Since S. albifrons attacks the sessile pupae, host move-

ments would not affect the distribution of this species. The amount of

spring movement by casebearer larvae is influenced by casebearer density,

greater movements occurring at higher densities (Webb 1953). At the

casebearer densities that occurred during this study the movements of

casebearer larvae would not appear to have affected the distribution of

Dicladocerus since the distributions of C. laricella were the same in

the 15 May and 13 June collections (Fig. 3).

The within-tree distributions of Dicladocerus spp. and S. albi-

frons in classes 1 and 2 are similar to those in 30-40 ft. trees in the

western United States (Tunnock et al. 1972). The distributions of

Dicladocerus spp. and S. albifrons within trees probably reduces compe-

tition for hosts between these species (Tunnock et al. 1972).

CONCLUDING DISCUSSION

Although 30 parasites of C. l a r i c e l l a were taken in British

Columbia, the aggregate percentage parasitism was not large. However,

the native parasite complex and the incidence of parasitism in British

Columbia were comparable to those in other areas of North America

(Bousfield and Lood 1971, 1973; Denton 1972; Sloan 1965; Webb 1953).

The parasite complex and incidence of parasitism in British Columbia

also resembled those in the Alps region of Europe, the area of origin

of C. l a r i c e l l a on European larch, L a r i x decidua Mill. (Jasch 1973),

although more major species (in terms of relative abundance and con-

stancy) occurred in the Alps. The parasite complexes in other areas of

western Europe were not as rich in species as was the Alps (Jasch 1973).

Zwijlfer and Pschorn-Walcher (1968) reviewed the parasite com-

plexes of several introduced pests and came to the following conclusions

about possibilities for parasitism in areas of introduction: (1) the

introduced pest may not be attacked by native parasites because the pest

is not similar taxonomically to native hosts; (2) the introduced pests

may be only occasionally and to a minor extent attacked by native parasites

similar to the parasites in the original area of the host or by polyphagous

parasite species; or (3) the introduced pest may be attacked to a large ext-

ent by very adaptable parasite species. C. l a r i c e l l a in British Columbia (and

North America generally) fits into the second category. The polyphagous

species taken include G. t e n e l l u s , S. d e c o r u s , E. cushmani, 2'. d o l o s u s ,

4 7

C . a e n e o v i r i d i s , and S . a l b i f r o n s while species somewhat restricted to

casebearers and needle miners as hosts include B. pygmaeus, C. r u f i p e s ,

I . v e s c a , Z . americanum, and H . p h y c i d i s (Krombein 1958; Krombein and

Burks 1967; Muesebeck et al. 1951; Peck 1963). The other specimens were

only identified to genus so it was not possible to determine the recorded

host spectrum of these. However, some of the species of the genera

Diadegma, G e l i s , P r i s t o m e x u s , E u l o p h u s , Copidosoma, Mesopolobus , Eury toma,

and Telenomus are polyphagous, while some species of the genera P r i s -

t o m e r u s , D i c l a d o c e r u s , and Mesopolobus are somewhat restricted to case-

bearers and needle miners (Krombein 1958; Krombein and Burks 1967;

Muesebeck et al. 1951; Peck 1963). Species of the genera Diadegma, G e l i s ,

I t o p l e c t i s , D i c l a d o c e r u s , E l a c h e r t u s , T e t r a s t i c h u s , Habrocy tus , and

Mesopolobus have been taken in both British Columbia and Europe (Jasch

19731, as have the two introduced species taken in British Columbia.

There has been an apparent increase in both number of species

and total percentage parasitism in British Columbia between 1966 (Andrews

and Geistlinger 1969) and 1973 to 1974. A similar increase occurred in

the western United States (Denton 1972). This increase raises the ques-

tion of what the ultimate role of native parasites might be. As the

native parasite complex and incidence of parasitism of C . l a r i c e l l a are

comparable to those of the native cherry casebearer, C. p r u n i e l l a , in

British Columbia (Waddell 1952), at least during outbreaks, the ultimate

role may already be reached (under the present circumstances).

Mortality of C . l a r i c e l l a caused by the native parasites may be

limited by the number of alternate hosts available to the parasites in

the absence of suitable stages of C. l a r i c e l l a since these or related

species are known to have more than one generation per year (Clausen

1962; Dowden 1941; Jasch 1973) and not all of these can be spent on C.

l a r i c e l l a . For example, in Europe Dicladocerus wes twoodi i Westw. that

emerged from C. l a r i c e l l a , attacked the larch grey roller moth, z e i r a p h e r a

d i n i a n a Guen. (Aeschlimann 1969). S. a l b i f r o n s is more dependent on

alternate hosts than other species as very few females of this species,

2.5 percent of the species total in 1973 and 0 percent in 1974, were

taken from C. l a r i c e l l a . The predominance of males of S. a l b i f r o n s from

C. l a r i c e l l a has been recorded in other areas (Bousfield and Lood 1973;

Webb 1953). The lack of alternate hosts has been suggested as a limiting

factor of parasitism of C. m a l i v o r e l l a (Beacher 1947) and C. p r u n i e l l a

(Doner 1934, 1936). Similarly, the minor influence of parasitism on

populations of the leafminer P h y l l o c n i s t i s l a b y r i n t h e l l a Bjerk. was the

result of the lack of suitable host stages for the parasites emerging

from this host in the spring (Sundby 1957). A positive trend was noted

between total percentage parasitism and the total number of lepidopteran

and sawfly larvae (which may or may not be alternate hosts of the para-

sites taken) at five sites during this study.

Alternate hosts are known to be important in other systems. For

example, the mymarid Anargus epos Gir., an important egg parasite of the

grape leafhopper Erythroneura e l e g a n t u l a Osborn, is known to overwinter

in the eggs of D i k r e l l a c r u e n t a t a (Gillette), a leafhopper on wild Rubus

spp. and not on the grape leafhopper. The ability of A. epos to reduce

the level of grape leafhopper infestations is a function of the distance

between vineyard and the endemic ecosystem where wild Rubus spp. occur

(mutt and Nakata 1973). Other examples of the role of alternate hosts

in parasitism are given by DeBach (1964).

Exotic parasites were released against C. l a r i c e l l a in eastern

North America (Dowden 1962; McGugan and Coppel 1962) because of the inef-

fectiveness of the native parasites in controlling the outbreak. They

have been credited with control of C. l a r i c e l l a (Munroe 1971; Turnbull

and Chant 1961) and of the non-target C. mal i vore l l a (LeRoux et al. 1963).

One of the two species credited with control of C. l a r i c e l l a , A. pumila,

is specific and synchronized with this host (Jasch 1973) while the other

species, C. l a r i c i n e l l a e , is synchronized with C. l a r i c e l l a in the pres-

ence of A. pumila or in the presence of alternate hosts (Quednau 1970).

In western North America, the native parasites did not prevent

C. l a r i c e l l a from reaching damaging population levels. A. pumila, the

first exotic parasite released in the western United States, did not

become established at all release sites nor has it dispersed well where

establishment did occur (Denton 1972). As a result, C. l a r i c i n e l l a e ,

Dicladocerus spp., Necremnus metalarus (Wlkr.), Diadegma l a r i c i n e l l a

(Strobl), and Elacher tus a rg i s sa Wlkr. were released (Ryan and Denton

1973; Ryan et al. 1975).

The chances of success for biological control with these species

seem limited. They are similar taxonomically and biologically to the

parasite species already present. Since they cannot spend all their gen-

erations on C. l a r i c e l l a (Jasch 1973), lack of alternate hosts or other

factors that may be limiting the native parasites may also limit these

exotic species. C. laricinellae cannot control C. laricella in the

absence of A. pumila or sufficient alternate hosts (Quednau 1970). Con-

trol of C. malivosella by C. laricinellae was apparently facilitated by

necessary alternate hosts (LeRoux et al. 1963). Population studies of

C. laricella in Europe showed parasitism to be a minor factor influencing

host populations (Eidmann 1965; Jasch 1973). Climatically, the areas of

larch casebearer infestation in British Columbia and the western United

States resemble the Alps region more than eastern North America.

It is possible that the exotic species may have different alter-

nate host spectra than the native species or other adaptations that might

allow for greater parasitism of C. laricella than that observed for

native species. Similarly, further releases of C. laricinellae from dif-

ferent source areas may result in the introduction of a better-adapted

strain. Strains are known to have different biological characteristics

affecting the success of biological control attempts (Messenger and van

den Bosch 1971). Further releases would also aid in the dispersal of

this species.

Studies of parasitism of C. laricella referred to above were

carried out during outbreaks. Percentage parasitism was apparently

inversely host-density dependent (although the significance of the rela-

tionship was not stated) in Europe (Jasch 1973). A similar tendency

occurred in British Columbia (increases in percentage parasitism coin-

cided with decreases in C. laricella density at the eight sites sampled

in both years); however, the number of samples was too small to determine

the significance of the relationship. Varley and Gradwell (1970)

considered factors that operate inversely density-dependently over the

whole range of an insect's densities are relatively rare but cite the

non-specific, non-synchronized parasites of the winter moth, Operophtera

brumata L., as an example. If a significant inverse host-density re-

sponse does exist, the parasites may be relatively more effective and

may be capable of controlling C. laricella at endemic (non-damaging)

population levels.

APPENDIX I

SITE DESCRIPTIONS

Locat ion : Anarchist Summi t

Elevation: 3,300'

Aspect: South

Stand History: Undisturbed

Stand Composition:

1. Western Larch:

Proportion of stand: 33%

Height: 65-75'

Diameter at Breast Height: 19-27"

2. Other Trees (proportion of Stand): Douglas-fir (33%), Ponderosa

pine (33%)

3. Tree Density: Open grown

4. Understory: undergrowth was sparse; mostly grasses, some others

such as Ribes sp., kinaikinnik

Location: Arrow Creek

Elevation: 2,100'

Aspect: East


Stand Composition:

1. Western Larch:

Proportion of Stand: 75%

Height: 50-60'


2. Other Trees (Proportion of Stand): Lodgepole pine (20%), Western

red cedar and Douglas-fir (5%)

3. Tree Density: Closed canopy

4. Undergrowth: sparse; mostly grasses, some others such as waxberry,

Rubus sp. present

Location: Cascade

Elevation: 2,100'

Aspect: North


Stand Composition:

1. Western Larch:


Height: 60-65'


2. Other Trees (Proportion of Stand): Lodgepole pine (50%), Douglas-

fir (15%)


4. Undergrowth: very sparse; some grass, few waxberry bushes

Location: Cranbrook

Elevation: 3,000'

Aspect: Northwest


Stand Composition:

1. Western Larch:


Height: 50-60'


2. Other Trees (Proportion of Stand): Lodgepole pine (60%)


4. Undergrowth: very sparse; grasses

Location: Fruitvale

Elevation: 1,400'

Aspect: North


Stand Composition:

1. Western Larch:


Height: 60-65'


2. Other Trees (Proportion of Stand): Douglas-fir (50%)


4. Undergrowth: very sparse; western red cedar, bracken and grasses

present

Location: Johnstone Creek Park

Elevation: 2,700'

Aspect: East South East


Stand Composition:

1. Western Larch:


Height: 60-70 '


2. Other Trees (Proportion of Stand): ~ouglas-fir (45%), Ponderosa

pine (25%)


4. Undergrowth: blanket of grass

Location: Kootenay Bay

Elevation: 1,850'

Aspect: South


Stand Composition:

1. Western Larch:


Height: 50-60'


2. Other Trees (Proportion of Stand): Western white pine (40%),

Douglas-f ir (30%)


4. Undergrowth: very sparse; few plants such as bracken, tall mahonia,

princess pine, waxberry present

Location: Roosville

Elevation: 2,600'

Aspect: East


Stand Composition:

1. Western Larch:


Height: 45-50'


2. Other Trees (Proportion of Stand): Douglas-fir (50%), Lodgepole

pine (15%)


4. Undergrowth: blanket of grasses

Location: Rossland

Elevation: 2,600'

Aspect: Southwest

Stand History: Partial cut

Stand Composition:

1. Western Larch:


Height: 60-70'


2. Other Trees (Proportion of Stand): Poplar (25%)


4. Undergrowth: blanket of bracken; many other plants such as tall

mahonia, waxberry, and elderberry present

Location: Rykerts

Elevation: 2,000'

Aspect: Southwest


Stand Composition:

1. Western Larch:


Height: 70-75'


2. Other Trees (Proportion of Stand): Douglas-fir (20%)


4. Undergrowth: very sparse; mosses, Clintonia sp. present

Location: Sheep's Creek

Elevation: 1,300'

Aspect: South


Stand Composition:

1. Western Larch:


Height: 30-35'


2. Other Trees (Proportion of Stand): Poplar, Douglas-fir (10%)


4. Undergrowth: even covering of grasses; bracken and a few other

species present

Location: Shoreacres

Elevation: 1,550'

Aspect: Southeast


Stand Composition:

1. Western Larch:


Height: 45-55'


2. Other Trees (Proportion of Stand): Douglas-fir (45%), Miscellaneous

(10%)


4. Undergrowth: blanket of grasses; many species such as bracken, tall

mahonia, waxberry present

Location : Winlaw

Elevation: 1,850'

Aspect: Northwest


Stand Composition:

1. Western Larch:

Proportion of stand: 40%

Height: 50-55'


2. Other Trees (Proportion of Stand): Douglas-fir (30%), Western white

pine (30%)


4. Undergrowth: even cover of grasses; bracken and tall mahonia

present

Location : Yahk

Elevation: 2,100'

Aspect: South


Stand Composition:

1. Western Larch:


Height: 65-70'


2. Other Trees (Proportion of Stand): Douglas-fir (35%), Lodgepole

pine (15%)


4. Undergrowth: sparse; grasses with a few waxberry bushes

APPENDIX I1

PERCENTAGE PARASITISM BY INDIVIDUAL PARASITE SPECIES

AT SITES IN BRITISH COLUMBIA ON FOUR

COLLECTING DATES IN 1973 AND 1974

Table XI.

Percentage parasitism of Coleophora laricella by

individual parasite

species in samples taken at eight sites in British Columbia on 8-9 May 1973

Site

Bracon

Dicladocerus

Eulophus

Habrocytus

Mesopolobus

Unemerged

Total

PY W

eus

SPP

SP

phycidis

SP -

Anarchist Summit

z

* *

1.3

Arrow Creek

* *

* *

Cascade

2.3

~t

3.0

Fruitvale

Rossland

Sheep's Creek

* x

*

Shoreacres

* *

1.4

Yahk

3.7

1.9

5.6

dP f-i

v +J 5 a +J d Q) rn a, & a

*

Table XIII. Percentage parasitism of C

ole

op

ho

ra l

ar

ice

lla

by individual parasite

species in samples taken at 14 sites in British Columbia on 14-15 May 1974

Site

Crown

Bra

co

n

Dia

degm

a P

ris

tom

eru

s Sc

ambu

s A

chry

soch

are

l la

Level

(ft.1

PY g

mae

us

SP .

SP .

decor

us

SP

Anarchist Summit .

. .

5-1

0

Arrow Creek

. . .

. .

Cascade

. .

. . .

. .

Cranbrook

. . . .

. .

Fruitvale

. . . .

. .

Johnstone Creek Park .

Kootenay Bay . . . . .

Roosville

. .

. . . .

Rossland .

. . .

. .

.

Rykerts

. . , . .

. .

Sheep's Creek

. .

. .

Shoreacres . .

. . .

. Winlaw . .

. . . .

. .

Yahk .

. . .

. . . .

.

20

-25

0

* 21

present but

< 1%

Table XIII--Continued

--- -

Site

Crown

Di cl adocerus

Copi dosoma

Mesopol obus

E ur y tom

Level

Unemerged

Total

(ft. 1

SPP.

(3)

SP .

SP

SP .

Anarchist Summit .

..

.....

Arrow Creek ..

..

..

.

Cascade

..

..

..

Cranbrook

Fruitvale

..

..

..

Johnstone Creek Park .

..

..

.

Kootenay Bay

Roosville

..

..

..

Rossland .

..

..

..

Rykerts

..

..

..

.

Sheep's Creek

....

Shoreacres .

..

..

.

Winlaw .

..

..

..

.

Yahk .

..

..

..

..

Table XIV.

Percentage parasitism of C

ole

op

ho

ra l

ar

ice

lla

by individual parasite

species in samples taken at 14 sites in British Columbia on 12-13 June 1974

Site

Crown

Bra

con

C

ampo

pl e

x

Dia

degm

a G

el

is

G

eli

s

Ito

ple

cti

s

Pri

sto

me

rus

Level

PY g

mae

us

ru fi

pe

s

SP .

ten

ell

us

sp.

ve

sca

SP

(ft.1

Anarchist Summit .

. 5-10

1.7

Arrow Creek

. .

. .

Cascade

. .

. . . .

Cranbrook

. .

. . .

Fruitvale

. . .

. .

Johnstone Creek Park

Kootenay Bay

Roosville

. . .

. .

Rossland .

. .

. . .

Rykerts

. .

. . .

. Sheep's Creek

. . .

Shoreacres . . .

. .

Winlaw . .

. .

. . .

Yahk .

..

..

. .. 2

0-25

cn

* U

)

present but

< 1%

Ta

ble

XIV--Continued

Cro

wn

Scambus

Achrysocharel la

Chrysocharis

Dicladocerus

Tetrastichus

Sit

e

Lev

el

decor us

SP

laricinellae

sp

p.

(3)

ecus

(f

t. 1

An

arc

his

t S

um

mit

. .

5-1

0

5.0

20

-25

6

.0

5-1

0

6.4

....

Arr

ow

C

reek

Casc

ad

e

..

..

..

.

..

..

C

ran

bro

ok

..

..

.

Fru

itv

ale

Joh

nst

on

e

Cre

ek

Park

....

Ko

ote

nay

B

ay .....

Ro

osv

ille

......

Ro

ssla

nd

..

..

.

Ry

ke

rts :

..

.

Sh

ee

p's

C

reek

.....

Sh

ore

ac

res

Win

law

.......

Ya

hk

. .......

Table

XIV

--C

on

tin

ued

--

Site

Crown

Zag

ram

mos

oma

Mes

op

ol o

bu

s S

pil

oc

ha

lcis

Level

Unemerged

Total '

am

eri c

anum

SP

alb

i f ro

ns

(ft.)

Anarchist Summit . .

5-10

6.7

....

Arrow Creek

Cascade

..

..

..

Cranbrook

.....

..

..

.

Fruitvale

Johnstone Creek Park

Kootenay Bay ..

..

.

Roosville

Rossland .

..

..

.

Rykerts

..

..

..

...

Sheep's Creek ..

..

.

Shoreacres

..

..

..

.

Winlaw

Yahk ........

REFERENCES

Aeschlimann, J. P. 1969. Contribution a ll&ude de trois gspeces d' Eulophides (Hymenoptera: Chalcidoidea) parasites de la Tordeuse grise du ~el&ce, Zeiraphera d i n i a n a Guen. (Lepid.: Tortricidae) en Haute- Engadine. Entomophaga 14:261-319.

Andrews, R. J. and N. J. Geistlinger. 1969. Parasites of the larch casebearer, Coleophora l a r i c e l l a (Hbn.) , in British Columbia (Lepidoptera: Coleophoridae). J. entomol. Soc. Brit. Columb. 66:50-51.

Beacher, J. H. 1947. Studies of pistol case-bearer parasites. Ann. ent. SOC. Am. 40:530-544.

Bousfield, W. E. and R. C. Lood. 1971. Impact of parasites on the larch casebearer in the Northern Region 1970. U.S.D.A. For. Serv. Rept. 71-4, Northern Region, Missoula, Mont.

Bousfield, W. E. and R. C. Lood. 1973, Parasites of the larch casebearer in Montana, Idaho, and Washington. Environ. Ent. 2:212-213.

Clausen, C. P. 1962. Entomophagous Insects. Hafner Publ. Co., New York.

Dawson, A. F. 1971. Larch casebearer in British Columbia. Can. For. Serv., Pac. For. Res. Cent. Pest Leafl. No. 34.

DeBach, P. (Ed. 1964. B i o l o g i c a l C o n t r o l o f I n s e c t P e s t s and Weeds. Reinhold Publ. Corp., New York.

Denton, R. E. 1958. The larch casebearer in Idaho--a new defoliator record for western forests. U.S.D.A. For. Serv. Res. Note 51, Intermt. For. and Range Exp. Stn.

Denton, R. of lax Idaho.

E. 1972. Establishment of A g a t h i s pumila (Ratz.) for control .ch casebearer, and notes on native parasitism and predation in U.S.D.A. For. Serv. Res. Note INT-164, Intermt. For. and

Range Exp. Stn.

Denton, R. E. and S. Tunnock. 1972. Larch casebearer in western larch forests. U.S.D.A. For. Serv., For. Pest Leafl. No. 96.

Dixon, W. J. 1973. BMD Biomedical Computer Programs. Univ. of Calif. ,Press, Los Angeles.

Doner, M. H. 1934. Observations on the biology of Microbracon pygmaeus

(Prov.), an important parasite of Coleophora p r u n i e l l a C1. Ann. ent. SOC. Am. 27:435-442.

Doner, M. H. 1936. Hymenopterous parasites of Coleophora p r u n i e l l a Cl., and parasites recorded from other species of Coleophora. Ann. ent. SOC. Am. 29:224-244.

mutt, R. L. and J. Nakata. 1973. The Rubus leafhopper and its egg para- sitoid: an endemic biotic system useful in grape-pest management. Environ. Ent. 2:381-386.

Dowden, P. B. 1941. Parasites of the birch leaf-mining sawfly (Phyl- lotoma nenwrata) . U.S.D.A. Tech. Bull. 757.

Dowden, P. B. 1962. Parasites and predators of forest insects liberated in the United States through 1960. U.S.D.A. Handbk. 226.

Eidmann, H. H. 1965. Okologische und physiologische Studien iiber die IArchenminiermotte, Coleophora l a r i c e l l a Hbn. Stud. For. Suec, 32.

Finlayson, T. 1967. Taxonomy of final-instar larvae of hymenopterous and dipterous parasites of Acrobasis spp. (Lepidoptera: Phycitidae) in the Ottawa region. Can. Ent. 99~1233-1271.

!$ ~asch. A. 1973. Populationsdynamik und Parasitenkomplex der Mrchen- miniermottr , Coleophora l a r i c e l l a Hbn., im natiirlichen Verbreitungsge- biet der Europaischen Larche, L a r i x decidua Mill. '2. angew. Ent. 73: 1-42.

Krombein, K. V. and others. 1958. Hymenoptera of America North of Mexico. Synoptic Catalog. U.S. Dep. Agric., agric. Monogr. 2, Suppl. 1.

Krombein, K. V. and B. D. Burks. 1967. Hymenoptera of America North of Mexico. Synoptic Catalog. U.S. Dep. Agric., agric. Monogr. 2, Suppl. 2.

LeRoux, E. J., R. 0. Paradis and M. Hudon. 1963. Major mortality factors in the population dynamics of the eye-spotted bud moth, the pistol casebearer, the fruit-tree leafroller, and the European corn borer in Quebec. Mem. ent. Soc. Can. 32:67-82.

McCugan, B. M. and H. C. Coppel. 1962. Biological control of forest insects, 1910-1958. I n A review of the biological control attempts against insects and weeds in Canada. Commonw. Inst. biol. Contr. Tech. Comm. No. 2.

Messenger, P. S. and R. van den Bosch. 1971. The adaptability of introduced biological control agents. In C. B. Huffaker (Ed.), Biolog ica l Con t ro l , pp. 68-92. Plenum Press, New York.

Miller, G. E. and T. Finlayson. 1974. Native parasites of the larch casebearer, Coleophora laricella (Lepidoptera: coleophoridae), in the west Kootenay area of British Columbia. J. entomol. SOC. Brit. C o l d . 71:14-21.

Morris, E. and J. Monts. 1972. Larch casebearer infestations in Nelson Forest District, 1972. Can. For. Serv., Forest Insect & Disease SU~V. Pest Rept., Pacific Forest Research Centre, Victoria, B.C.

Muesebeck, C. F. W., K. V. Krombien and H. K. Townes. 1951. Hymenoptera of America North of Mexico. Synoptic catalog. U.S. Dep. Agric., agric. Monogr. 2.

Munroe, E. 1971. Status and potential of biological control in Canada. In Biological control programmes against insects and weeds in Canada, 1959-1968. Commonw. Inst. biol. Contr. Tech. Comm. 4:213-255.

Peck, 0. 1963. A catalogue of the nearctic Chalcidoidea (lnsecta: Hymenoptera). Can. Ent. Suppl. No. 30.

Quednau, F. W. 1970. Competition and cooperation between C. laricinellae and A . pumila on larch casebearer in Quebec. Can. Ent. 102:602-612.

Ryan, R. B. and R. E. Denton. 1973. Initial releases of Chrysocharis laricinellae and Dicladocerus westwoodii for biological control of the larch casebearer in the western United States. U.S.D.A. For. Serv. Res. Note PNW-200.

Ryan, R. B., W. E. Bousfield, G. E. Miller and T. Finlayson. 1974. Presence of Chrysocharis laricinellae, a parasite of the larch casebearer, in the Pacific Northwest. J. econ. Ent. 67:805.

Ryan, R. B., W. E. Bousfield, R. E. Denton, R. L. Johnsey, L. F. Pettinger, and R. F. Schmitz. 1975. Additional releases of larch casebearer parasites for biological control in the western United States. U.S.D.A. For. Serv. Res. Note PNW-242.

Sloan, N. F. 1965. Biotic factors affecting populations of the larch casebearer Coleophora laricella Hbn. in Wisconsin. Ph.D. Thesis, Univ. of Wisconsin, Madison, Wisc.

Sloan, N. F. and H . , C . Coppel. 1965. Oviposition patterns and egg predation of the larch casebearer, Coleophora laricella Hbn. in Wisconsin. Univ. of Wisconsin For. Res. Notes No. 124.

Sundby, R. 1957. The parasites of Phyllocnistis labyrinthella Bjerk. and their relation to the population dynamics of the leaf-miner. Norsk.

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Telford, A. D. 1961. Lodgepole needle miner parasites: biological control and insecticides. J. econ. Ent. 54:347-355.

Tunnock, S., M. McGregor and W. E. Bousfield. 1972. Distribution of larch casebearer parasites in the crowns of western larch trees in the Northern Region. U.S.D.A. For. Serv. Rept. 72-4, Northern Region, Missoula, Mont.

Tunnock, S., R. E. Denton, C. E. Carlson and W. W. Janssen. 1969. Larch casebearer and other factors involved in the deterioration of western larch stands in northern Idaho. U.S.D.A. For. Serv. Res. Pap. INT-68.

Turnbull, A. L. and D. A. Chant. 1961. The practice and theory of biological control of insects in Canada. Can. J. Zool. 39:697-753.

Varley, G. C. and G. R. Gradwell. 1970. Recent advances in insect population dynamics. Ann. Rev. Ent. 15:l-24.

Waddell, D. B. 1952. Biology and control of the cherry casebearer, Coleo- phora pruniella Clemens, in British Columbia. Proc. entomol. Soc. Brit. Colurnb. 48:85-89.

Webb, F. E. 1953. An ecological study of the larch casebearer, Coleophora laricella Hbn. (Lepidoptera: Coleophoridae). Ph.D. Thesis, Univ. of Michigan, Ann Arbor, Mich.

Zwijlfer, H. and H. Pschorn-Walcher. 1968. Wie verhalten sich Insekten- parasiten gegentiber eingeschleppten, faunenfremden Wirten? Anz. Schaklingsk. 41:51-55.

CURRICULUM VITAE

Gordon Edward MILLER

HOME ADDRESS:

OFFICE ADDRESS:

PERSONAL:

Apartment 428 - 281 Holdom Avenue Burnaby, British Columbia, Canada V5B 3T9 Telephone: (604) 291-1014

April, 1976

Department of Biological Sciences, Simon Fraser University, Burnaby, British Columbia, Canada V5A 1S6 Telephone: (604) 291-4285

Born August 3, 1950; Vancouver, British Columbia; 6'3"; 195 lbs.; married; health excellent

SPECIAL INTERESTS :

My area of professional interest and specialization is pest management. Specific interests include forest and agricultural entomology, insect population dynamics, pheromones, pest damage, pest resistance of plants, integrated control, economic thresholds, and damage by diseases and weeds to crops

EDUCATION:

1974-present

Honors

1966-1968

Honors

Master of Pest Management program at Simon Fraser Uni- versity taken concurrently with the M.Sc. program. Graduation is expected in 1976.

Master of Science program in Biological Sciences at Simon Fraser University.

Simon Fraser University, Burnaby, B.C. B.Sc.(Hons.) in Biological Sciences 1/72. Seventy-two semester hours of course work in bialogical sciences, sixty of which are in upper levels courses.

B.C. Government Scholarships in three semesters. Simon Fraser University Open Scholarship in one semester.

Burnaby Central Senior Secondary School, Burnaby, B.C. Completed grade 12. Diploma.

1962 Roll of Honor for Scholastic Achievement; 1963 Minor Award for Athletics; 1964 Major Award for Athletics.

EDUCATION (Cont Id. 1

1962-1966 Moscrop Junior Secondary School, Burnaby, B.C.

1956-1962 Cascade Heights Elementary School, Burnaby, B.C.

CERTIFICATES HELD:

British Columbia Pesticide Applicator Certificate (No. 4292) in: 1. Agricultural Crop Pest Abatement 2. Forest Pest Abatement 3. Nonagricultural and Nonforestry Vegetation Control 4. Landscape and Garden Pest Abatement 5. Structural Pest Abatement 6. Mosquito Abatement

WORK EXPERIENCE:

1972-present Graduate S t u d e n t and Research A s s i s t a n t . Pestology Centre, Department of Biological Sciences, Simon Fraser University. Survey of insect parasites of the larch casebearer in B.C. for the Department of the Environment; with the degree of parasitism in relation to location within a plot, casebearer density, tree height, and density of alternate hosts.

Responsibilities: Planning and leading field work; collection, preparation, and species identification of insect parasites; supervision of six technicians, five at once.

Research A s s i s t a n t . Pestology Centre, Department of Biological Sciences, Simon Fraser University. Bark and ambrosia beetle biology, rearing, and sexing. Pheromone bioassays. Insect histology. Antenna1 morphology of parasitic Hymenoptera.

Research A s s i s t a n t . Pestology Centre, Department of Biological Sciences, Simon Fraser University. Insect identification.

F i e l d Researcher . Labatt ' s Fraser River Project . Vancouver, B.C. Interviewed companies about pollution problems.

S t r a i g h t e n e r . Western Canada Steel, Vancouver, B.C. Steel processing.

S a n i t a r y Eng ineer . H . R. MacMillan Planetarium, Van- couver, B.C. Responsible for building's appearance.

RESEARCH PUBLICATIONS:

J. H. Borden, G. E. Miller and J. V. Richerson. 1973. A possible new sensillum on the antennae of Itoplectis conquisitor (Hymenoptera: Ichneurnonidae). Can. Ent. 105: 1363-1367.

G. E. Miller and T. Pinlayson. 1974. Native parasites of the larch casebearer, Coleophora laricella (Lepidoptera: Coleophoridae) in the West Kootenay area of British Columbia. J. ent. Soc. Brit. Columb. 71:14-21.

R. B. Ryan, W. E. Bousfield, G. E. Miller and T. Finlayson. 1974. Presence of Chrysocharis laricinellae, a parasite of the larch casebearer in the Pacific North- west. J. econ. Ent. 67:805.

TEACHING EXPERIENCE :

Teaching Assistant in the following courses: 1. Insect Biology 73-3 2. Plant Ecology 72-3 3. Population Dynamics 74-1 4. Introductory Biology 73-1, 75-1

MEMBERSHIPS :

Entomological Society of Canada Entomological Society of America Entomological Society of British Columbia

PROFESSIONAL CONFERENCES ATTENDED:

1975 Entomological Society of British Columbia

1973 Western Forest Insect Workshop, Tucson, Arizona

1972 Joint Meeting of the Entomological Society of America, Pacific Branch and the Entomological Society of British Columbia, Victoria, B.C.

NON-ACADEMIC ACTIVITIES:

University: 1973 Graduate Student Representative on the Department Under-

graduate Curriculum Committee

Community: 1975 Coach of boy's football team 1974 Coach of boy's baseball and football teams. 1973 Coach of boy's soccer team

Hobbies : Volleyball, Floor Hockey, Tennis, Other Sports, Photo- graphy

REFERENCES :

Pro fe s so r T. F in layson , Professor . Pestology Cent re , Department of ~ i o l o g i c a l Sc iences , Simon F r a s e r Univers i ty , Burnaby, B.C., Canada V5A 1S6

D r . J. H. Borden, Professor . Pestology Cent re , Department o f B io log ica l Sc iences , Simon F r a s e r Univer- s i t y , Burnaby, B.C. , Canada V5A 1S6

D r . B. P. Beirne, P ro fe s so r and Director, Pestology Cent re , Department of B io log ica l Sc iences , Simon F r a s e r Univers i ty , Burnaby, B.C., Canada V5A 1S6

NATIVE PARASITES OF COLEOPHORA WlRICELLA (LEPIWPTERA

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