native parasites of coleophora wlricella (lepiwptera
TRANSCRIPT
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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.
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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 ' -
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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 )
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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
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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 .
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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.
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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
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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
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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 lari- cella in 1973 and 1974 collections in British Columbia . . . . . . . . . . . . . . 13
111. Number of taxa and total percentage para- sitism at individual sites in 1973 and 1974 collections in British Columbia . . . . . . . . . 2 0
IV. Density of Coleophora laricella and percen- tage 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 dis- tributions 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
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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 lari- cella by individual parasite species in samples taken at eight sites in British Columbiaon8-9May1973 . . . . . . . . . . . 6 2
XII. Percentage parasitism of Coleophora lari- cella by individual parasite species in samples taken at eight sites in British Columbia on 23-25 May 1973 . . . . . . . . . . 63
XIII. Percentage parasitism of Coleophora lari- cella 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
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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
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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
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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.
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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
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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 .
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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
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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 .
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(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
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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.
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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
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m r- . . 0 0
C O O d N
cy t. . . 0 0
0-l d rl m
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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
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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
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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
.
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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
.
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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
.
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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
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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
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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).
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m r - w
o r - m . . (U d ' r -
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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
.
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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).
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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.
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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
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IL- C. laricella 11-11 Dicladocerus spp. -I- S. albifrons mle..-.-l.... Mesopolobus sp.
0 4 8 12 16 20 24
DAYS
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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.
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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
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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
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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 signifi- cance)
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
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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 signifi- cance)
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 16.890 0.81 - 3 7.499 0.53 -
Within Classes
Total 39 3 4
- not significant
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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 sig- nificance)
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 4 108.958 5.39 - 3 51.452 1.31 -
Within Classes
Total 39 3 4
- not significant
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Fig. 3. Schematic representation of within-tree distri- butions 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 .)
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1 May 15, 1974 collection
-
1 June 13, 1974
~ u n e 1 3, 1974 collection
I I I
2 collection
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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
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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
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Fig. 4. Schematic representation of within-tree distribu- tions of Dicladocerus spp. in five classes of trees on 13 June 1974 at Shoreacres, British Columbia. (Numbers repre- sent percentage parasitism, with outer being those of outer branch half and inner those of inner branch half)
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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
-
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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
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Fig. 5. Schematic representation of within-tree distribu- tions 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)
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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 -
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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
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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).
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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 ,
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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
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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
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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
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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)
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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.
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APPENDIX I
SITE DESCRIPTIONS
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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 History: Undisturbed
Stand Composition:
1. Western Larch:
Proportion of Stand: 75%
Height: 50-60'
Diameter at Breast Height: 23-27"
2. Other Trees (Proportion of Stand): Lodgepole pine (20%), Western
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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 History: Undisturbed
Stand Composition:
1. Western Larch:
Proportion of Stand: 35%
Height: 60-65'
Diameter at Breast Height: 20-26"
2. Other Trees (Proportion of Stand): Lodgepole pine (50%), Douglas-
fir (15%)
3. Tree Density: Closed canopy
4. Undergrowth: very sparse; some grass, few waxberry bushes
Location: Cranbrook
Elevation: 3,000'
Aspect: Northwest
Stand History: Undisturbed
Stand Composition:
1. Western Larch:
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Proportion of Stand: 40%
Height: 50-60'
Diameter at Breast Height: 17-20"
2. Other Trees (Proportion of Stand): Lodgepole pine (60%)
3. Tree Density: Closed canopy
4. Undergrowth: very sparse; grasses
Location: Fruitvale
Elevation: 1,400'
Aspect: North
Stand History: Undisturbed
Stand Composition:
1. Western Larch:
Proportion of Stand: 50%
Height: 60-65'
Diameter at Breast Height: 16-24"
2. Other Trees (Proportion of Stand): Douglas-fir (50%)
3. Tree Density: Closed canopy
4. Undergrowth: very sparse; western red cedar, bracken and grasses
present
Location: Johnstone Creek Park
Elevation: 2,700'
Aspect: East South East
Stand History: Undisturbed
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Stand Composition:
1. Western Larch:
Proportion of Stand: 30%
Height: 60-70 '
Diameter at Breast Height: 28-30"
2. Other Trees (Proportion of Stand): ~ouglas-fir (45%), Ponderosa
pine (25%)
3. Tree Density: Closed canopy
4. Undergrowth: blanket of grass
Location: Kootenay Bay
Elevation: 1,850'
Aspect: South
Stand History: Undisturbed
Stand Composition:
1. Western Larch:
Proportion of Stand: 30%
Height: 50-60'
Diameter at Breast Height: 19-23"
2. Other Trees (Proportion of Stand): Western white pine (40%),
Douglas-f ir (30%)
3. Tree Density: Closed canopy
4. Undergrowth: very sparse; few plants such as bracken, tall mahonia,
princess pine, waxberry present
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Location: Roosville
Elevation: 2,600'
Aspect: East
Stand History: Undisturbed
Stand Composition:
1. Western Larch:
Proportion of Stand: 35%
Height: 45-50'
Diameter at Breast Height: 13-17"
2. Other Trees (Proportion of Stand): Douglas-fir (50%), Lodgepole
pine (15%)
3. Tree Density: Open grown
4. Undergrowth: blanket of grasses
Location: Rossland
Elevation: 2,600'
Aspect: Southwest
Stand History: Partial cut
Stand Composition:
1. Western Larch:
Proportion of Stand: 75%
Height: 60-70'
Diameter at Breast Height: 27-33"
2. Other Trees (Proportion of Stand): Poplar (25%)
3. Tree Density: Open grown
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4. Undergrowth: blanket of bracken; many other plants such as tall
mahonia, waxberry, and elderberry present
Location: Rykerts
Elevation: 2,000'
Aspect: Southwest
Stand History: Undisturbed
Stand Composition:
1. Western Larch:
Proportion of Stand: 80%
Height: 70-75'
Diameter at Breast Height: 20-27"
2. Other Trees (Proportion of Stand): Douglas-fir (20%)
3. Tree Density: Closed canopy
4. Undergrowth: very sparse; mosses, Clintonia sp. present
Location: Sheep's Creek
Elevation: 1,300'
Aspect: South
Stand History: Undisturbed
Stand Composition:
1. Western Larch:
Proportion of Stand: 90%
Height: 30-35'
Diameter at Breast Height: 10-12"
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2. Other Trees (Proportion of Stand): Poplar, Douglas-fir (10%)
3. Tree Density: Open grown
4. Undergrowth: even covering of grasses; bracken and a few other
species present
Location: Shoreacres
Elevation: 1,550'
Aspect: Southeast
Stand History: Partial cut
Stand Composition:
1. Western Larch:
Proportion of Stand: 45%
Height: 45-55'
Diameter at Breast Height: 20-23"
2. Other Trees (Proportion of Stand): Douglas-fir (45%), Miscellaneous
(10%)
3. Tree Density: Open grown
4. Undergrowth: blanket of grasses; many species such as bracken, tall
mahonia, waxberry present
Location : Winlaw
Elevation: 1,850'
Aspect: Northwest
Stand History: Partial cut
Stand Composition:
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1. Western Larch:
Proportion of stand: 40%
Height: 50-55'
Diameter at Breast Height: 17-23"
2. Other Trees (Proportion of Stand): Douglas-fir (30%), Western white
pine (30%)
3. Tree Density: Open grown
4. Undergrowth: even cover of grasses; bracken and tall mahonia
present
Location : Yahk
Elevation: 2,100'
Aspect: South
Stand History: Undisturbed
Stand Composition:
1. Western Larch:
Proportion of Stand: 50%
Height: 65-70'
Diameter at Breast Height: 12-17"
2. Other Trees (Proportion of Stand): Douglas-fir (35%), Lodgepole
pine (15%)
3. Tree Density: Closed canopy
4. Undergrowth: sparse; grasses with a few waxberry bushes
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APPENDIX I1
PERCENTAGE PARASITISM BY INDIVIDUAL PARASITE SPECIES
AT SITES IN BRITISH COLUMBIA ON FOUR
COLLECTING DATES IN 1973 AND 1974
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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
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dP f-i
v +J 5 a +J d Q) rn a, & a
*
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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%
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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 .
..
..
..
..
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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%
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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
. .......
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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 ........
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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 case- bearer, 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
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(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 in- sects, 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 intro- duced biological control agents. In C. B. Huffaker (Ed.), Biolog ica l Con t ro l , pp. 68-92. Plenum Press, New York.
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Miller, G. E. and T. Finlayson. 1974. Native parasites of the larch case- bearer, 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 case- bearer, 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 preda- tion 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.
. ent. Tidsskr. Suppl. 2.
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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 bio- logical control of insects in Canada. Can. J. Zool. 39:697-753.
Varley, G. C. and G. R. Gradwell. 1970. Recent advances in insect popu- lation 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.
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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.
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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; col- lection, 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.
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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
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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