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A study of some factors influencing the abundance of Adelges cooleyi (Gill.) on Douglas fir Kozak, Antal 1961

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i  A STUDY OF SOME FACTORS INFLUENCING THE ABUNDANCE OF ADELGES COOLEYI (GILL.) ON DOUGLAS FIR.  by  ANTAL KOZAK B.S.F., The University of B r i t i s h Columbia (Sopron D i v i s i o n ) , 1959  A THESIS SUBMITTED IN PARTIAL FULFILMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF FORESTRY  i n the Department of FORESTRY  We accept t h i s thesis as conforming to the required standard  THE UNIVERSITY OF BRITISH COLUMBIA A p r i l , 1961  In the  presenting  requirements  of  British  it  freely  agree for  that  an  advanced  for  available  I  copying  gain  shall  by or  not  his  Apn'1  degree  at  the  reference  and  study.  I  extensive  may  be  granted  representatives.  allowed  of  copying  this  without  by  of  the  It thesis  is  Columbia,  make  further this  Head  of  thesis my  understood  for  my w r i t t e n  of  University shall  of  10,  fulfilment  Library  The U n i v e r s i t y o f B r i t i s h V a n c o u v e r 8, Canada. Date  p a r t i a l  the  publication be  in  that  for  purposes  or  agree  for  permission  that  Department  thesis  Columbia,  scholarly  Department  this  f i n a n c i a l  permission.  ABSTRACT  A study was  carried out to determine factors which influence the abun-  dance of Adelges (Chermes) cooleyi G i l l , on Douglas f i r . through the observation  This was  of populations i n the f i e l d , supported by some labora-  tory work designed to show, that certain influences are important. was  done  The work  done during the summer' of I960 i n a young stand i n Totem Park near Marine  Drive and Agronomy Road on the U.B.C. campus, Vancouver. The following items were investigated: 1.) Intra tree v a r i a t i o n s , 3.)  Inter tree differences,  2.)  Population changes with time, h») M o r t a l i t y of the  insect. Abundance was  affected by e x t r i n s i c influences on the trees, such as  l o c a t i o n and exposure and i n t r i n s i c factors such as time: of bud  opening and'  twig length. Within a tree the abundance of Adelges cooleyi was  affected mostly by  microclimatic f a c t o r s , r e s u l t i n g i n high abundance of the insect i n the  pe-  r i p h e r a l part of the lower crown. The average number of l i v i n g insects decreased with time r e c t i l i n e a r l y i n generation 1 (Sexuparae and Progredientes) and l o g a r i t h m i c a l l y i n generation 2 (Neosistens).  A c r i t i c a l period during establishment of generation 2  caused the logarithmic changes. M o r t a l i t y estimates by d i r e c t counts were subject to a large error because many of the dead insects f e l l o f f .  iii  ACKNOWLEDGEMENTS  Acknowledgement i s made to the U n i v e r s i t y of B r i t i s h Columbia f o r the laboratory and education f a c i l i t i e s , which aided m a t e r i a l l y i n t h i s research. Grateful appreciation i s expressed by the writer to the members of the s t a f f of the Department of Zoology and Faculty of Forestry, expecially to Dr. Kenneth Graham f o r advice and encouragement throughout the work,and to Dr. J . Harry G. Smith f o r counsel on s t a t i s t i c a l  analyses.  Special thanks are due Mr. Jozsef Csizmazia f o r h i s help i n evaluating some of the s t a t i s t i c a l analyses, using the ALwac I I I E e l e c t r o n i c computer.  iv  CONTENTS Page TITLE PAGE  i  ABSTRACT  i i  ACKNOWLEDGEMENT  i i i  CONTENTS  iv  TABLES  vi  FIGURES  ix  INTRODUCTION  1  MATERIALS AND METHODS  3  1. THE INSECT  3  2. THE STAND  Ik  3. DESCRIPTION OF SAMPLE TREES AND SAMPLING METHODS. h. PROCESSING OF DATA EXPERIMENTAL RESULTS  . . .  15" 18 20  1. POPULATION CHANGES BY TIME  21  2. INTER TREE DIFFERENCES  33  3. INTRA TREE VARIATIONS  Ul  it. MORTALITY  51  5. NEED FOR FURTHER STUDIES  54 57  DISCUSSION 1. INTER TREE DIFFERENCES  57  LOCATION OF THE TREES  57  EXPOSURE  58  BUD OPENING TIME  60  TWIG LENGTH  61  2. INTRA TREE VARIATIONS EXPOSURE  62 62  HEIGHT ON LIVING CROWN  Page ~oT  CHANGES IN ABUNDANCE OF INSECT IN HORIZONTAL DIRECTIONS OF THE TREE CROWN  63  TWIG LENGTH  6k  3. MORTALITY  6k  k. NATURAL ENEMIES  66  5. BIOTIC POTENTIAL  67  SUMMARY  68  CONCLUSIONS  69  APPENDICES  70  BIBLIOGRAPHY  93  vi  TABLES  Number 1  Heading  Page  Relative numbers of sexuparae and progredientes on Douglas f i r i n three counts  7  2  Analysis of variance  f o r generation 1  20  3  Analysis of variance  f o r generation 2  21  h  Calculation of the equation and variables by period and average needle Calculation of the equation and variables by period and average needle.  5  6  c o r r e l a t i o n c o e f f i c i e n t of number of l i v i n g insects per 23 c o r r e l a t i o n c o e f f i c i e n t of number of dead insects per 2h  Calculation of the equation and c o r r e l a t i o n c o e f f i c i e n t of the number of l i v i n g insects per needle at low crown l e v e l of the trees on period  26  Calculation of the equation and c o r r e l a t i o n c o e f f i c i e n t of the number of dead insects per needle at low l e v e l of the trees on period  26  Calculation of the equation and c o r r e l a t i o n c o e f f i c i e n t of average number of l i v i n g insects per needle on time. . . .  30  Calculation of the equation and c o r r e l a t i o n c o e f f i c i e n t of average number of dead insects per needle on time. . . .  31  Calculation of the equation and c o r r e l a t i o n c o e f f i c i e n t of average number of l i v i n g insects per needle on time. . .  32  Calculation of the equation and c o r r e l a t i o n c o e f f i c i e n t of average number of dead insects per needle on time. . . .  32  12  Analysis of variance  33  13  Differences  .  3U  lU  Influence of p o s i t i o n of tree i n the stand on abundances of Adelges cooleyi  35  Differences among trees by abundance of Adelges c o o l e y i . The trees were treated as two population, separated by location  36  Average numbers of l i v i n g insects per needle, by exposures and l e v e l s f o r the marginal trees  37  7  8 9 10 11  15  16  among trees, l e v e l s and generations.  among trees by abundance of Adelges cooleyi.  vii  Number  Heading  Page  17  Analysis of variance  18  Differences between d i f f e r e n t l y exposed trees by the abundance of Adelges cooleyi  38  Influence of 8 factors on the average number of l i v i n g insects per needle  kO  Calculation of the equation and c o r r e l a t i o n c o e f f i c i e n t of average number of l i v i n g insects per needle on height from the ground (data of June 5)  kl  Calculation of the equation and c o r r e l a t i o n c o e f f i c i e n t of average number of l i v i n g insects per needle on height from the ground (data of J u l y 25)  k3  Calculation of the equation and c o r r e l a t i o n c o e f f i c i e n t of average number of l i v i n g insects per needle on height from the ground (data of September 26)  kk  Calculation of the equation and c o r r e l a t i o n c o e f f i c i e n t of average twig length on height from the ground  k6  Calculation of the equation and c o r r e l a t i o n c o e f f i c i e n t of average numbers of l i v i n g insects per needle on distances from the stem (data of June 5)  k8  Calculation of the equation and c o r r e l a t i o n c o e f f i c i e n t of average numbers of l i v i n g insects per needle on distances from the stem (data of J u l y 25)  k8  Calculation of the equation and c o r r e l a t i o n c o e f f i c i e n t of average number of l i v i n g insects per needle on distances from the stem (data of September 26)  k9  Calculation of the equation and c o r r e l a t i o n c o e f f i c i e n t of average lengths of twigs on distances from the stem. . .  k9  19 20  21  22  23 2k  25  26  27 28  Analysis of variance  among exposures and l e v e l s  of dead insects among trees, l e v e l s  and exposures (generation 29  Analysis of variance Analysis of variance  1) 2)  32  52  of dead insects among trees, l e v e l s  and generations 31  51  of dead insects among trees, l e v e l s  and exposures (generation 30  38  53  Variance r a t i o t e s t between the main factors and f i r s t order interactions  53  Percentage of mortality by l o c a l i t y of the trees and generations  5k  viii  Number 33 3k-5k  Heading Percentage of m o r t a l i t y by l e v e l s and generations. See: L i s t of Appendices  Page . . .  5k 70  ix  FIGURES  Number  Heading  Page  1  The l i f e cycle of Adelges cooleyi  k  2  A, G a l l i c o l a migrans with wings closed laying eggs on f i r needle (X20) B, Egg-mass produced by G a l l i c o l a or progrediens ( X 3 0 ) . .  6  A, Winged sexupara (X60) B, Antenna of winged sexupara (X2lj0) C, Antenna of winged g a l l i c o l a (X210)  9  3  h 5> 6  7 8  9 10 11 12 13  lh 15  A, Sistens nymph, f i r s t B, Progrediens adult  instar 10  Number of l i v i n g and dead insects per needle i n generation 1 f o r d i f f e r e n t periods  22  Number of l i v i n g and dead insects per needle i n generation 1 f o r d i f f e r e n t periods at the low l e v e l of the l i v i n g crown  2$  Number of l i v i n g and dead insects per needle i n generation 2 f o r d i f f e r e n t periods  27  Number of l i v i n g and dead insects per needle i n generation 2 f o r d i f f e r e n t periods at the low l e v e l of the l i v i n g crown  28  Transformed curves of f i g . 7 (B) and f i g . 8 (A).  29  .  .  .  Relationship between twig lengths and number of l i v i n g insects per needle  39  Relationship between number of l i v i n g insects per needle and height on the l i v i n g crown  U2  Relationship between twig length and height on the l i v i n g crown  U5  Relationship between number of l i v i n g insects per needle and distance from the stem at the low l e v e l of the l i v i n g crown.  U7  Relationship between twig length and distance from the stem at the low l e v e l of the l i v i n g crown  50  Relationship between the number of l i v i n g insects per needle and height on the l i v i n g crown  59  A STUDY OF SOME FACTORS INFLUENCING THE ABUNDANCE OF ADELGES COOLEYI (GILL.) ON DOUGLAS FIR.  INTRODUCTION.  The cooleyi spruce g a l l aphid Adelges (Chermes) cooleyi ( G i l l . ) , as a pest of Douglas f i r and S i t k a spruce, varies i n abundance from year to year, place to place, and even between trees i n the same l o c a l i t y .  I t i s so v a r i a -  ble i n numbers as to merit examination of the factors influencing i t s abundance. The insect and i t s secondary host, Douglas f i r , Pseudotsuga Menziesii var.  Menziesii (Mirb.) Franco, o f f e r a unique opportunity f o r the study of  environmental influences on an i n s e c t .  The insect lends i t s e l f well to  observation and sampling because i t i s s e s s i l e during most of i t s l i f e , i s thus suitable f o r quantitative work.  and  Douglas f i r i s so variable i n i t s  phenology that a wide range of phase r e l a t i o n s between i t and iU cooleyi  may  be found. The cooleyi spruce g a l l aphid was named and described by G i l l e t t e i n 1907.  He c a l l e d the generations on spruce: Chermes c o o l e y i , and generations  on Douglas f i r Chermes cooleyi var. cowenii.  In 191U  ChrystaL's (6) experi-  ment proved that, G i l l e t t e ' s Chermes cooleyi var. cowenii i s the c o l o n i c i stage of Chermes c o o l e y i . the species belongs i n 1928,  Annand changed the name of the subfamily to which from Chermisinae to Adelginae, because the former  name had been e a r l i e r used f o r the p s y l l i d s groups (7). No detailed work has been done about the factors influencing the abundance of iU cooleyi on Douglas f i r .  The f a c t has been stated by Chrystal  (7) and Cameron ( k ) , that Adelges cooleyi i s more abundant on young trees i n  2  the nursery and j u s t a f t e r plantation (23).  Chrystal (7)  mentioned that i n  a dense wood of Douglas f i r the cooleyi spruce g a l l aphid was  found c h i e f l y  on the trees near the margin and was not present i n the centre of the plantation.  He has also stated that the insects were more frequent on the lower  and middle branches of the trees.  Both i n England and Germany i t has been  established that, Pseudotsuga Menziesii var. glauca (Beissn) Franco i s more r e s i s t a n t to  cooleyi than the Pseudotsuga Menziesii var. Menziesii  (Mirb.) Franco. The main b i o t i c f a c t o r s - a f f e c t i n g the abundance of A. cooleyi - are the natural enemies.  According  attacked by comparatively  to the present knowledge the insect i s  few enemies.  Two  species of Syrphidae were found by  Cumming (10) which attack the exposed forms on Douglas f i r , and eggs of Syrphidae were also found i n one g a l l c a v i t y on spruce.  The yellow and  Ladybird beetle i s known to feed on Adelges on Douglas f i r ( 7 ) .  Several  authors have noted large numbers of Spiders and Red-Spider-Mites feeding A. c o o l e y i .  Gyorfi (17)  black  on  has mentioned that Hymenopterous parasites of the  family Chalcididae attack c e r t a i n species of European Adelges. Heavily infested Douglas f i r s are not k i l l e d d i r e c t l y by Adelges, but are so reduced i n vigour that they are more susceptible to attack of secondary i n s e c t and disease.  The cumulative e f f e c t of the attack probably reduces  the rate of growth, reducing the active surface of a s s i m i l a t i o n , r e s u l t i n g i n reduced food suply (12, 13),  Although the trees are not k i l l e d by Adelges, i t  i s recommended that Douglas f i r and S i t k a spruce mixture not be planted unsuitable s i t e or l o c a l i t i e s .  on  The most c r i t i c a l stage i n the l i f e both of  Douglas f i r and S i t k a spruce i s j u s t a f t e r the plantation has been made  (7).  MATERIALS AND METHODS 1. The Insect: Two generations of the A. cooleyi were the main material of the experiment, established to examine the factors influencing t h e i r abundance. Adelges cooleyi prefers two d i f f e r e n t species «of host-trees i n i t s complete l i f e cycle.  In the P a c i f i c Coast the primary host i s S i t k a spruce (Picea  sitchensis (Bong.) Carr.), and the secondary host i s Douglas f i r  (Pseudotsuga  *—X  Menziesii var. Menziesii (Mirb.) Franco).  The complete l i f e cycle of  A. cooleyi consists of 5 "generations" ( F i g . l ) s Fundatrix Generation: The f i r s t - s t a g e l a r v a of fundatrix overwinters on a Sitka spruce terminal twig adpressing i t s body to the twig, and sunking i t s s t y l e t deeply i n t o the tissue through bark f i s s u r e s .  Passing through the winter i n t h i s stage, they  become active i n the f i r s t part of A p r i l , and begin to feed. are reported to have three l a r v a l i n s t a r s (9, 10).  The fundatrices  These l a r v a l stages are  present f o r a r e l a t i v e l y short time i n the spring. The f i r s t - i n s t a r larvae are dark brown with s l i g h t l y greenish colour. The general appearance the second-instar larvae resemble the adults, having thinner c u t i c l e .  The adults are dark brown with a green tinge (10).  The  body of the adult or stem-mother i s covered with a white wax-mass, under which she lays a large number of eggs (150-350) (10) at the end of A p r i l . G a l l i c o l a Migrans: The g a l l i c o l a e migrantes are the gall-forming generation on spruce. This generation i s the progeny of fundatrices.  This summer generation hatches  about 10 - 15* days a f t e r the eggs are deposited by the stem-mothers, and migrates to the inner bases of young needles, which are j u s t breaking from the bud at t h i s time.  As soon as they have s e t t l e d , they begin to feed, and  g a l l s begin t o develop, caused by some stimulation, induced by the feeding  Fig-1 The life cycle of Adelges cooleyi  insect (LU).  This stimulant i s supposed to be an auxin (1, 26).  The g a l l v a r i e s i n length from 1 - 5 the twig attacked (18).  inches, varying with the vigour of  The number of chambers within the g a l l ranges from  kO to 200, the number of young i n each chamber varying from 1 to 15, with an average of 5 ( 6 ) . The larvae go through four i n s t a r s within the g a l l s (9, 10). reddish-brown coloured larvae are covered with a fine powdery wax. wing pads can be seen at the t h i r d - i n s t a r .  The The  They emerge from the g a l l s i n  the fourth-instar and s e t t l e on the needles, and moult to the adult stage. The parthenogenetic adults are reddish-brown with a well s c l e r o t i z e d thorax. They f l y to the Douglas f i r , where being s e t t l e d f o r a few days, they secrete wax from the anterior margin of the head and the abdomen. abdomen and the wings protects the eggs which they l a y .  The wax from the  The average number  of eggs l a i d was found to be 65 of 10 counts (9, 10) ( F i g . 2). Sistens (Fundatrix spuria!); The sistens are the overwintering stage on Douglas f i r .  They are  progeny of the g a l l i c o l a migrans or progeny of the second generation of wingless females, s t a i d on Douglas f i r on the previous summer. says that the sistens hatch 6-7  days a f t e r the o v i p o s i t i o n .  has been stated by Cumming (10) as about three weeks. the time required i n I960 was 2 or 2\ weeks.  Chrystal (6) This period  In the present study  The youngs s e t t l e on the lower  surface of Douglas-fir needles, and pass the winter i n t h i s stage.  This  dormant form of sistens generation used to be called neosistens ( F i g . h ) . This generation was one subject i n the experiment, and was 8 times during the summer of I960.  sampled  This generation w i l l be c a l l e d generation  2 i n the further part of the t h e s i s . The neosistens are light-brown with a darker prothorax and head.  It i s  u n l i k e l y that t h i s stage feeds before going into the dormant condition.  Fig- 2  A, Gallicola migrans with wings closed laying eggs on fir needle (X 20) B, Egg-mass produced by Gallicola or progrediens (X 30 )  7  However, they move from the needle a f t e r being hatched, looking f o r suitable needles t o overwinter, where they press t h e i r bodies to the underside o f the needles, i n s e r t i n g t h e i r s t y l e t s i n t o the raesophyll t i s s u e .  This i n s e r t i o n  i s believed to afford a d d i t i o n a l anchorage, rather than providing food. When s e t t l e d , they secrete a fringe of white wax around the edge of the body. The f i r s t moult occurs i n the spring. larvae i s covered with wax. larvae but with more wax.  The body of the second-instar  The adults are s i m i l a r to the second-instar  The adults are parthenogenetic females, laying  from 30 to hO eggs (6). There are two kinds of progeny of the sistens: a. )The progredientes, being wingless parthenogenetic females. They remain on Douglas f i r . b. )The sexuparae, being winged parthenogenetic females.  They  migrate t o spruce. The proportion of these two forms varies widely.  Ghrystal (6) found  that about $0% o f the brood was sexupara i n Stanley Park i n 1915.  Three  counts are given by Cumming (10) from the Kananaskis Forest Experiment Station (Table l ) . Table 1.  Relative numbers of sexuparae and progredientes on Douglas f i r i n three counts.  Date  , Sexuparae  Number counted Progredientes Doubtful J-  Per cent of Sexuparae  6 - 711 - 55  63  5  16  75  15 - 711 - 55  72  U  0  95  3 - 711 - 57  8U  76  11  U9  The Sexuparae and Progredientes; They hatch about a week a f t e r o v i p o s i t i o n . time when the buds s t a r t t o open.  Hatching begins about the  After hatching they move to the needles  -•-Larva too small to d i s t i n g u i s h form.  8  and develop i n t o winged adults (Sexuparae) or wingless adults (Progredientes). The larvae and the seasonal development resemble generation 2, being l i g h t yellow-brown, but become darker and darker as they pass through the moults. The larvae resemble those of the g a l l i c o l a e , but the g a l l i c o l a e have more glands.  Wax  can be found on the d o r s a l and v e n t r a l surfaces of the head  during the l a r v a l stage. There are four l a r v a l i n s t a r s , three of them s i m i l a r i n morphology of the two forms.  Wing pads are present on the fourth i n s t a r larvae of  sexuparae. The generation of sexuparae and progredientes was also material of the experiment, being sampled 5 times during the summer of I960.  The progre-  dientes and sexuparae were not observed separately, because there was reason to t r e a t them separately f o r the subject of the experiment.  no  This  generation w i l l be c a l l e d as generation 1 i n the further part of the t h e s i s . Sexuparae t The adults of sexuparae are winged parthenogenetic females ( F i g . 3). Wax  i s present on a l l parts of the body, except the v e n t r a l surface of the  abdomen, the adults are h e a v i l y s c l e r o t i z e d .  The eyes, head and metathorax  are dark grey-black, and the remainder of the body i s brown.  The wings are  grey with green bases and greenish-gray costa. They migrate back to spruce, s e t t l e on the needles, then l a y eggs (10).  The eggs are protected by wax  5-20  and the folded wings of the female.  Progredientes: The adults of progredientes are wingless parthenogenetic females (Fig.U), resembling the s i s t e n s .  They stay on Douglas f i r .  by Progredientes i s v a r i a b l e . eggs.  The number of eggs l a i d  Chrystal (6) has stated that they l a y 30 - UO  The average number of eggs i s given as 15 with a range from 3 to 25  i n 11 counts by Cumming (10).  F i f t e e n egg-clusters were counted i n Totem  A, Winged sexupara ( X 60 ) B, Antenna of winged sexupara ( X 240) C, Antenna of winged gallicola (X 210) (After Chrystal)  Fig- 4  A, Sistens nymph,first instar B, Progrediens adult  Park on June 25,  The average of these was 27,  I960.  with a range from 5 to  k3. Progredientes hatch 2 - 3  weeks a f t e r oviposition, and overwinter i n  t h i s undeveloped stage (Neosistens).  Consequently  a subcycle of the l i f e  cycle can be found on Douglas f i r ( F i g . 1), consisting of two generations; Sistens (Neosistens) and Progredientes (Sexuparae) (28).  This means that the  Adelges cooleyi can develop without i t s primary host. Sexualis Generation; This i s the one generation of the Adelges cooleyi where sexual reproduct i o n occurs. Chrystal (6)  This generation i s not described w e l l i n most of the l i t e r a t u r e . did not observe t h i s generation i n B. C ,  the generation i n B r i t a i n .  but l a t e r (7)  he found  S t i U l a t e r Annand i n C a l i f o r n i a , Cameron (k) i n  B r i t a i n , Francke-Grosmann ( l k ) i n Germany and Cumming (10) i n Alberta recorded the sexual generation. The most detailed report on t h i s generation was given by Cumming (9, 10). According to him they hatch a week or more a f t e r the o v i p o s i t i o n of the sexuparae.  Chrystal (6),  distinguished 3 only.  and Cameron (k) found $ l a r v a l i n s t a r s .  Cumming (10)  I t i s possible that the f i r s t two i n s t a r s can be  so s i m i l a r , that they were not distinguished (10). The colour of the larvae i s reddish-brown.  The adults move back from  the present year growth to the old growth, as f a r as k - 5 years-old twigs. This habit of moving back helps them i n the mating, helping the male f i n d the female.  They l a y eggs between the old bud scales and twigs.  The  female secretes wax, which surrounds but does not cover completely the egg. I t i s believed that the female lays only one egg (10, l k ) . Morphology of Adelges  cooleyi;  The description of the generations presented here w i l l not be complete, only the differences between the generations w i l l be pointed out.  The most  remarkable differences can be seen i n the gland facets, i n t h e i r shapes and sizes.  There are also differences i n the arrangement of the plates on which  the gland facets occur.  The v e n t r a l plates are s l i g h t l y s c l e r o t i z e d , and  always have small round gland f a c e t s .  The dorsal plates are heavily  s c l e r o t i z e d , being i n three p a i r s , showing a single transverse row on each segment.  This row i s arranged from the mesothorax to the s i x t h or seventh  segment of the abdomen.  Two rows can be found on the head and prothorax.  The c e n t r a l p a i r of glands on the dorsal surface i s c a l l e d "mesial", the one beside them " p l e u r a l " , and the l a t e r a l one i s "marginal" (10). Fundatrix; A l l the dorsal gland facets of the f i r s t - i n s t a r larvae are angular i n form, and everyone i s w e l l outlined except those on the sixth and seventh segments of abdomen.  The posterior plates of the adults have a higher propor-  t i o n of round gland f a c e t s .  They also have angular gland facets which are  not so large as those of the sistens and progredientes (10). G a l l i c o l a Migrans; The f i r s t - i n s t a r larvae have only setae on the thoracic and abdominal segments.  These larvae are the smallest of a l l generation of Adelges  The number of gland areas increases i n s t a r after i n s t a r .  cooleyi.  The fourth-instar  resembles the adult, but the gland areas are smaller on the abdomen. The gland facets of the dorsal surface o f the adult are round. of the head and thorax are constant.  The gland areas  These areas vary on the abdomen. The  mesial areas of the abdomen are fused at the centre l i n e , and the p l e u r a l areas are broken up into small groups of gland f a c e t s .  The hamuli on the  hind wing v a r i e s from 2 to 5. Sexupara: The f i r s t - i n s t a r l a r v a has setae surrounded by small p l a t e s .  The adults  have large v a r i a t i o n i n the number and arrangement of gland areas on the  13  abdomen.  The antennae are shorter than those of the g a l l i c o l a ( F i g . 3).  The hamuli on the hind wing i s varying from 1 to h (10). Sistens and Progrediens: The f i r s t - i n s t a r sistens l a r v a can be distinguished by the appearance of the p l a t e s . the centre l i n e . laterally.  Two s c l e r o t i z e d areas cover the head which are separated at Gland facets can be found on these areas a n t e r i o r l y and  On each side of the prothorax a large sclerotized area can be  found, each of them i s fused from four p l a t e s .  The mesial, p l e u r a l and  marginal areas of the mesothorax, metathorax and the abdominal segments^ from 1 to U have rectangular p l a t e s . and seventh abdominal p l a t e s .  Gland facets can be seen on the f i f t h , s i x t h The gland facets resemble those of the fun-  datrix l a r v a e . The larvae of the progrediens resemble the. sistens, but they have more glands at the l a t e r a l margins and the r e c t i c u l a t i o n s are more conspicuous. The separation of the sistens and progredientes can be done by the gland facets on the a n t e r i o r mesial p l a t e of the prothorax and by the p l e u r a l plate of the f i r s t abdominal segment.  The sistens has smaller number of  gland facets i n both areas than the progrediens, and the whip of the antenna i s shorter than i n that of the progrediens.  More sistens have gland  area on the hind coxa than progredientes (10). Sexualls: The male and female cannot be distinguished i n the f i r s t - i n s t a r . l a s t - i n s t a r female has a wax gland at the p o s t e r i o r end. smaller than any of the generations.  The  The adults are  Setae are present on the head thorax  and abdomen of the male, where gland areas can be seen on the other forms.  2. The Stand; The place of the experiment was Totem Park and i t s surrounding planted stand.  The main component of the stand i s Douglas f i r .  Three age classes  can be found; most o f the trees are 2k - 26 years o l d , but some 20 and 28 - 29 years o l d trees are also present. natural regeneration,  The oldest class seems to be  and the two younger classes are planted.  The main  material of the plantation was Douglas f i r , with some S i t k a spruce. other tree species are also present as a r e s u l t of natural The vegetations  Some  regeneration.  of three layers of the f o r e s t , as tree crown l a y e r (A),  shrub l a y e r (B) and ground vegetation  (C) are the follows:  The Vegetation of the Forest; A l e v e l ; crown closure; 95$ Pseudotsuga Menziesii Thuja p l i c a t a Tsuga heterophylla Picea sitchensis Taxus b r e v i f o l i a ALnus rubra Acer macrophyllum Prunus sp. B l e v e l : closure 20 - 2$% Gaultheria shallon Sarabucus pubens Rubus s p e c t a b i l i s Rubus v i t e f o l i u s Rosa gymnocarpa Rubus p a r v i f l o r u s Symphoricarpus r i v u l a r i s  Vaccinium parviflorum Physocarpus capitatus C l e v e l : closure 25 - 30% Polystichum nunitum Pteridium  aquilinum  Sorbus sitchensis Dryopteris  austiaca  Sambucus pubens Claytonia s i l i r i c a Trientalis l a t i f o l i a Galium t r i f l o r u m Dicentra formosa Pseudotsuga Menziesii V Tsuga heterophylla Acer macrophyllum In the southern part of the forest most of the B and C l e v e l vegetation has been removed.  The s o i l of t h i s part i s tramped down, having a thick  l e v e l of raw humus. Generally the s o i l of the f o r e s t has been developed on sandy deposits of the Fraser River, to a podzolic s o i l . Ao, 1 inch, A l , 2 inches, and A2, I2 The  The surface layers of the s o i l are:  inches.  "B or depositional l a y e r i s also present but i t was too deep to dig n  down to "C or basic l a y e r to measure "B". M  I t appears that the f o r e s t type  - indicated by the s i t e index, s o i l and vegetation - i s a disturbed Douglas f i r - Moss association (11). 3. Description of Sample Trees and Sampling Methods: Twenty-five trees were sampled during the period of study, from June to 'the end of September.  These trees were marked with numbered tags from 1 to  25.  The trees were chosen with regard to two 1.  things:  Location of the treej three classes were established:  a. ) Trees growing at the center of the f o r e s t . b. ) Trees growing at the d i f f e r e n t margins of the f o r e s t . c. ) Open-growth t r e e s . Eight of the 25 trees were chosen at the center of the f o r e s t , 13  of  them at the margins and k of them were open growing. 2.  Time of bud-opening: considerable differences occurred i n the  time of opening of buds of Douglas f i r (16).  Three classes were established  to d i f f e r e n c i a t e the trees: a. ) E a r l y (E): the buds were open before May b. ) Medium (M): the buds opened from May  15.  15 to May  c. ) Late ( L ) : the buds opened l a t e r than May  21.  21.  Ten of the 25 trees were early-opening, k of them were medium, and  11  were late-opening. The trees were chosen, as much as possible, with regard to height, D.B.H., crown class and age.  The opportunity f o r selecting f o r these v a r i -  ables was l i m i t e d , because the f o r e s t consists of planted trees of l i m i t e d v a r i a t i o n i n these f a c t o r s . Both of the progrediens and sistens (Neosistens)  s e t t l e d on the current  year's growth, consequently the new growth of foliage was sample u n i t was  established as a needle.  of the l i v i n g crown of each t r e e . v i s u a l l y separated  sampled.  The  basic  Samples were drawn from three l e v e l s  The l i v i n g crowns of the trees were  into three l e v e l s (Low, Medium, High), and the samples  were drawn from the middle of each l e v e l (Appendix E ) .  North, East, South  and West sides of the trees were sampled i n each l e v e l . drawn from the outer part of the crown. cut from a normal t r e e .  Consequently 12  The sample© were sample twigs were  Some of the trees are not normal i n crown shape  (Appendix L ) ,  lacking some part of the crown.  margins are of t h i s kind. from one tree.  Most of the trees on the  In these cases the 12 samples could not be drawn  Some of the sample trees were very high, and d i f f i c u l t to  sample with safety i n the high l e v e l and sometimes even i n the medium l e v e l . These l e v e l s were sampled only three times during the summer. The frequency of sampling was varied according to circumstances.  The  trees were sampled once per week from June U, to August 6, and after t h i s time only once per two weeks or once per three weeks.  The change of the  length of sampling period was suggested by the low a c t i v i t y of insects a f t e r August 6, because by t h i s time most of the neosistens were established on the needle preparatory to overwintering. Prunners were used to cut the sample twigs from the lower trees, and i n the lower l e v e l s of the higher trees.  Two prunners were used, one of  the two was 8 feet i n length, and the other one consisted of three 1 5 - feet - long pieces, working t e l e s c o p i c a l l y .  This second prunner was  not u s e f u l over 3 0 - 3 5 f e e t i n length, because i t was very f l e x i b l e . For sampling the higher trees a ladder was used, which could be extended to 3h f e e t from which pole prunners could be operated.  Beyond the reach of  prunners, samples were obtained by climbing the t r e e . The cut twigs were placed i n nylon bags, i n which the samples were taken to the laboratory, where the l i v i n g and dead insect were counted on 5  ten randomly chosen needles of each twig.  D i f f e r e n t i a t i o n between dead and  l i v i n g larvae was made with the aid of a microscope.  The information was  kept separately f o r l i v i n g and dead insects (Appendix A, B). The d i f f e r e n t stages and i n s t a r s of the insects were observed at the same time, when they were counted.  The progrediens and sexupara were the  material of the sampling from June k to J u l y 2 , and the neosistens generation was the material from J u l y 9 to September 2k (Appendix F ) .  18  The trees, No. U, 10 and  25jWere  specially sampled to establish the  distribution of the Adelges cooleyi horizontally and vertically within a The samples by height were drawn from every 5 feet of the outer part  tree.  of the living crown, from four sides of the tree (North, West, South and East).  The samples radially out from the stem to the outer part of the crown  were drawn from every 3 feet, to four directions from the stem (North, West, South and East) in the low level of the crown. This special sampling was repeated 3 times during the summer, with sampling of the progrediens and sexupara generation on June 5, and the neosistens generation on July 25 and on September 26 (Appendix G).  Five twig lengths were also measured in each  sampling place on September 26 (Appendix G). The current year growths of the 25 sample trees were measured 2 times during the summer, on June k and September 26.  Ten twigs were measured i n  each sampling location on the 25 trees (Appendix J ) . The growth of foliage was also recorded once per week on two trees, on tree No. lb, (early opened) and on tree No. 15 (late opened) (Appendix H ) . The crown shapes of the 25 trees were studied and three typical shapes were drawn from two different views (Appendix L ) . Owing to lack of equipment, the meteorological data were not measured. The data presented as Appendix K are from the U.B.C. Meteorological Station. The place of observation of these data was within 300-1*00 yards of Totem Park. The natural enemies of Adelges cooleyi were observed, but lack of time did not permit measurement of mortality attributable to them. k. Processing of data; To determine the relationships between the abundance and factors, most of the data were subjected to statistical analyses.  The detailed processing  of data w i l l be discussed in the next chapter. Some of the measured data were not analysed, but they w i l l be used to  explain the e f f e c t of several f a c t o r s .  20  EXPERIMENTAL RESULTS: Six trees - No. 1 , 2 , 1 0 , 1 1 , l h and 2 0 - o f the 2 5 could be sampled i n each l e v e l and at every c a r d i n a l points.  The abundance of the l i v i n g i n s e c t s  on these 6 trees was analyzed by three factors: l e v e l , exposure (cardinal points) and bud-opening time (Appendix M), separately f o r the two generations (Table 2 and 3 ) . Table 2 . Analysis of variance f o r generation  1.  Degrees of freedom  Net sum squares  Mean sura squares  Variance ratio (F)  Significance at 0 . 0 5 level  Levels (L)  2  0.74  0.37  7.40  H.S.*  Exposures (E)  3  0.03  0.01  - .  Bud opening (B)  2  1.21  0.61  12.20  H.S.  L x E  6  0.05  0.01  -  N.S.  L x B  h  0.14  O.Oij  -  N.S.  E x B  6  0.08  0.01  L x E x B  12  0.10  0.01  Residual  36  1.94  0.05  Total  71  4.29  Source  N.S.  N.S.  -  N.S.  * H.S.=highly s i g n i f i c a n t , S i g n i f i c a n t , N.S.=Not s i g n i f i c a n t . The l e v e l s and the bud-opening times show s i g n i f i c a n t d i f f e r e n c e s . In table 3 the bud-opening times show also s i g n i f i c a n t d i f f e r e n c e , the l e v e l s do not, but the "F" value of l e v e l s i s very close to the s i g n i f i c a n c e level.  Examining the factors i n table 2 and 3 , the average numbers of l i v i n g  insects per needle were greater at the low l e v e l ( 0 . 3 1 i n generation 1 and 0.39  i n generation 2) than at the medium l e v e l (O.2I4 i n generation 1 and 0 . 2 8  i n generation 2 ) , and i t was greater at the medium l e v e l than at the high l e v e l (0.07 i n both generations) of the l i v i n g crown.  The average number of  l i v i n g insects was also greater on the e a r l y bud-opening trees ( O . 3 8 i n  21  Table 3.  Analysis of variance f o r generation  2.  Degrees of freedom  Net sum squares  Mean sum squares  Variance ratio (F)  Significance at 0.05 l e v e l  Levels (L)  2  1.26  0.63  3.15  N.S.  Exposures (E)  3  0.19  0.06  -  N.S.  Bud opening (B)  2  3.56  1.78  8.90  H.S.  L x E  6  0.07  0.01  -  N.S.  L x B  h  0.75  0.19  1.00  N.S.  E x B  6  0.3k  0.06  -  N.S.  L x E x B  12  0.12  0.01  -  N.S.  Residual  36  7.31  0.20  Total  71  13.60  Source  generation 1 and 0.56 1 and 0.09 and 0.09  i n generation 2)  i n generation 2)  i n generation  It  than on the medium (0.17  and l a t e bud opening trees (0.07  i n generation  i n generation 1  2).  Both of the analyses of variance indicate that there i s no s i g n i f i c a n t difference between exposures, consequently the data of the four cardinal points w i l l not be treated separately i n the further analyses. 1.  Population changes with time;  The f i r s t f i v e sampling time of the 13 remainder covered the generation  covered the generation 1,  and the  2.  a. Population changes with time i n generation  1:  F i r s t , the c o r r e l a t i o n between number of insects per needle and time was tested f o r the 6 trees mentioned. plotted by time ( F i g . 5). separately.  The average number of insects per needle  was  The numbers of dead and l i v i n g insects were plotted  The points suggested  the c a l c u l a t i o n of linear-regression equa-  tions f o r both numbers of l i v i n g and numbers of dead insects (Table U,  5).  22  June 4  II  18 Sampling Period  25  July 2  Fig 5 Number of living and dead insects per needle in generation I for different periods  23  Table l u Calculation  o f the equation and correlation c o e f f i c i e n t of variables  by period and average number of l i v i n g insects per needle.  t  i  XY  0  0.31  0 .  0.0961  0.00  1  0.25  1  0.0625  0.25  2  0.25  u  0.0L8U  O.hh  3  0.16  9  0.0256  0.1*8  U  0.11  16  0.0121  O.kh  10  1.05  30  0.2UU7  1.61  2.0  0.21  X = period, Y = average number of insects per needle, b • -0.QL9, a * 0.31, r - -0.998, t - -26.97, Sb = 0.00257, -0.057 - i  £  -O.Okl.  Where: b = sample regression c o e f f i c i e n t , a = intercept. r = correlation  coefficient,  t = " t " t e s t f o r correlation  coefficient.  Sb • standard deviation f o r regression, i  = confidence i n t e r v a l s .  The equation o f the regression l i n e i s : Y = -O.OU9X+0.31 The periods.  correlation i s s i g n i f i c a n t between the average number of insects and The values o f " r " and " t " are h i g h l y s i g n i f i c a n t .  The average  number o f insects per needle decreased l i n e a r l y with time. There i s no s i g n i f i c a n t correlation between the average number o f dead insects per needle and time (Table 5 ) , but points f i t well t o the l i n e . The unsignificant  r e s u l t probably was caused by the small sample s i z e .  L o g i c a l l y , the slope of the m o r t a l i t y l i n e should be the reverse o f the l i n e o f l i v i n g insects, but t h i s i s not so, f o r two reasons: i.  about 50$ o f the insects o f t h i s generation develop into winged  Table 5.  Calculation of the equation and c o r r e l a t i o n c o e f f i c i e n t of variables  by period and average number of dead insects per needle.  t  Y  —yT~  Y*—  XY  0  0.21  0  o.ouui  1  0.17  1  0.0289 0.17  2  0.20  a  O.OiiOO  3  0.20  9  o.ouoo 0.60  k  0.23  16  0.0529  0.92  10  1.01  30  0.2059  2.09  2.0  0.00  0.1*0  0.20  X = periods, Y = number of dead insects per needle, b " 0 . 0 0 7 , a r - 0.51,  t - 1.02,  Sb • 0.00686, -0.015  6  m  0.19,  i *+0.029.  The equation of the regression l i n e i s = Y = 0 . 0 0 7 X + 0 . 1 9 adults, and f l y to spruce.  The number of winged adults could not be observed  because of t h e i r f l i g h t . i i . Some of the dead insects could not be observed, because many of them f e l l o f f , and also many of them were eaten by predators. The c o r r e l a t i o n between number of insects per needle and period, was extended f o r every tree sampled.  The average numbers of insects at the lower  crown l e v e l of the trees were used, because they were most abundant there ( F i g . 6) (Appendix C) (Table 6, 7 ) . The " t " and " r " show that (Table 6) the c o r r e l a t i o n i s s i g n i f i c a n t between the number of l i v i n g insects per needle and time. The lack of c o r r e l a t i o n (Table 7) between the number of dead insects per needle and time probably i s caused by the small sample s i z e . b. Population changes with time i n generation 2: Eight samples out of 13 were drawn from generation 2 . samples were treated i n the same way as i n generation 1 .  The data of these F i r s t the average  June 4  II  18  25  July 2  Sampling Period  Fig  6 Number of living and dead insects per needle in generation I for different periods at the low level of the living crown  26  Table 6. Calculation of the equation and correlation c o e f f i c i e n t of the number of l i v i n g insects per needle at low crown l e v e l of the trees on period.  Total Ave.  X  Y  0  0.28  0  0.078U  0.00  1  0.23  1  0.0529  0.23  2  0.23  k  0.0529  0.U6  3  O.lij  9  0.0196  0.1*2  U  0.12  16  O.OLUU  0.U8  10  1.00  30  0.2182  1.59  2.0  XY  0.20  X = periods, Y - numbers of insects per needle, b •  r - -0.960, t • -5.89, Sb - 0.00775, -0.063  6  -0.0U1,  a •= 0.28,  1 -0.019 £  Equation: Y = -O.OiaX+0.28 Table 7.  Calculation of the equation and c o r r e l a t i o n c o e f f i c i e n t of the number  of dead insects per needle at low l e v e l of the trees on period. X  Total Ave.  Y  —  X " 2  y2  XY  0  0.15  0  0.0225  0.00  1  0.12  1  O.OU4U  0.12  2  0.16  a  0.0256  0.32  3  0.15  9  0.0226  o.a5  U  0.16  16  0.0256  0.6a  10  0.7U  30  0.1106  1.53  2.0  0.15  X = periods, Y = numbers of insects per needle, b -  0.005,  a = O.lh, r =  0.U8,  t - 0.95, Sb - 0.00606, -0.012 * i * 0.022 Equation: Y -  0.005X+0.ia  numbers '-of insects per needle of the 6 trees mentioned were plotted  against  2?  \ 0-3  \  \  \  \  \  a> a.  \  in  8 0 2 c  Number of living insects  a> E  Z a> cr> o > <  0-  Number  July 9  Fig  7  16  Number  23  30  Aug 6  J  L  13 20 27 Sampling Period  Sept 3  of living a n d d e a d insects p e r n e e d l e  for different  periods  10  17  in generation  24  2  28  jul/§  ife  23  30  A u g ^ 13 20 Sampling Period  27  Septl  iO  17  24~  Fig-8 Number of living and dead insects per needle in generation 2 for different periods at the low level of the living crown  Log Sampling Period  Log Sampling Period 01  Fig 9  02  03  0-4  0-5  0-6  07  Transformed curves of fig 7 (B) and fig-8(A)  08  0-9  10  30  time (Fig. 7 ) .  Between the f i r s t and second and between the second and third  sampling periods the line of the living insects i s increasing, because these periods were the hatching time. decrease.  From the third point, the line starts to  The manner of the points suggested to leave out the f i r s t two  points.  The remaining points show a curve, which was transformed to loga-  rithms.  Both the X and Y were changed to logarithms (Table 8 ) (Fig. 9).  Table 8 . Calculation of the equation and correlation coefficient of average number of living insects per needle on time. X  Y  1  logY  0.33  0.000  -O.U 81  0.0000  0.2314  2  0.31  0.301  -0.509  0.0906  0.2591  -0.1532  3  0.22  0.477  -0.658  0.2275  0.4330  -0.3139  5  0.19  0.699  -0.721  O.U 886  0.5198  -0.5040  8  0.16  0.903  -0.796  0.8154  0.6336  -0.7188  10  0.17  1.000  -0.770  1.0000  0.5929  -0.7700  T# 29  1.38  3.380  -3.935  2.6221  2.6698  -2.4599  A*  0.23  0.563  -0.656  U.83  (logX)  (logY)  ' (logX)(logY)  logX  2  2  0.0000  T* = Total A*  m  Average  X = periods, Y = numbers of living insects per needle, b = - 0 . 3 3 9 , a = O . U 6 5 , r - - 0 . 9 6 3 , t = - 6 . 8 0 , Sb - 0 . 0 U 9 , - 0 . 2 0 3 * i  18  -O.U65  Equation: logY = -0.3391ogX-0.U65 Both the values of r " and " t " give significant correlation between the M  transformed "Y" and "X , meaning that the correlation i s logarithmic between n  the average number of living insects per needle and time. The equation of the regression line was also calculated for number of dead insects per needle i n the generation 2 (Fig. 7 ) (Table 9 ) . Significant correlation was found between the average.number of dead  31  insects per needle and time. The  c o r r e l a t i o n analysis between number of insects per needle and period  was extended to include a l l of the 25 trees sampled.  The average number of  insects at the low l e v e l of each tree was used, because insects were most abundant there (Appendix C, D). Table 9« Calculation of the equation and c o r r e l a t i o n c o e f f i c i e n t of average number of dead insects per needle on time. X  Y  0  0.03  1  p -1 8  XY  0  0.0009  0.00  o.ou  1  0.0016  o.ou  2  o.o5  a  0.0025  0.10  3  0.05  9  0.0025  o.i5  U  0.06  16  0.0036  0.2U  6  0.06'  36  0.0036  0.36  9  0.07  81  0.00U9  0.63  11  0.10  121  0.0100  1.10  36  0.U6  268  0.0296  2.62  U.5  0.06  X = periods, Y = numbers of dead insects per needle, b • 0 . 0 0 5 , a • O.OU, r - 0 . 9 U 8 , t - 7 . 2 9 , Sb - 0 . 0 0 0 6 9 0 . 0 0 3 * i  &  0.007  Equation: Y - 0 . 0 0 5 X + 0 . 0 U The  average numbers of l i v i n g insects had to be tested the same way as  i n table 8.  The f i r s t two points were omitted and both X and Y were trans-  formed to logarithms ( F i g . 8, 9 ) (Table 1 0 ) . A s i g n i f i c a n t c o r r e l a t i o n was found between X and Y, meaning that the numbers of l i v i n g insects decreased l o g a r i t h m i c a l l y with time (Table 1 0 ) .  32  Table 10. Calculation of the equation and c o r r e l a t i o n c o e f f i c i e n t of average number of l i v i n g insects per needle on time.  Total Ave.  X  Y  logX  logY  1  0.30  0.000  -0.523  0.0000  0.2735  -0.0000  2  0.23  0.301  -0.638  0.0906  0.1*070  -0.1920  3  0.18  0.1*77  -0.7U5  0.2275  0.5550  -0.355k  5  0.17  0.699  -0.770  0.1*886  0.5929  -0.5382  8  0.15  0.903  -0.82k  0.815U  0.6790  -0.71*1*1  10  0.15  1.000  -0.821*  1.0000  0.6790  -0.821*0  29  1.18  3.380  -4.321*  2.6221  3.1861*  -2.6537  0.20  0.563  -0.721  lw 83  (logX)  (logY)  2  2  (logX)(logY)  X = periods, Y • numbers of l i v i n g insects per needle, b = -0.303, a = -0.551, r = -0.9681*, t =-8.07, Sb = 0.013, -0.267 - i --0.339 Equation: logY - -0.3031ogX-0.551 Table 11. Calculation of the equation and c o r r e l a t i o n c o e f f i c i e n t of average number of dead insects per needle on time. X  Total Ave.  Y  •'X '  XY  2  0  0.011*  0  0.0002  1  0.019  1  0.0001* 0.019  2  0.050  I*  0.0025 0.100  3  0.01*1*  9  0.0019  0.132  1*  0.01*8  16  0.0023  0.192  6  0.067  36  0.001*5 0.1*02  9  0.071*  81  0.0055 0.666  11  0.071*  121  0.0055 0.811*  36  0.390 268  4.5  0.01*9  0.0228  0.000  2.325  33  X = periods, Y ° numbers of dead insects per needle, b r = 0.898, t *  e  0.00$, a  a  0.0026,  5.0U5, Sb - 0.00126, 0.002 * i & .008 0  Equation: Y • 0.005X+0.026  The r s u l t s (Table 11) show s i g n i f i c a n t c o r r e l a t i o n between X and Y, meaning that the numbers of dead insects per needle increased  with time.  2. Inter Tree Differences: The numbers o f l i v i n g and dead insects were summarized by tree, generat i o n and l e v e l i n tree (Appendix N).  Analysis of variance was carried out  f o r the l i v i n g insects (Table 12). Table 12. Analysis of variance among trees, l e v e l s and generations.  Source  Degrees of freedom  Net sum squares  Mean sum squares  CompoVariance S i g n i f i ratio cance at nents of (F) 0.05 l e v e l Variance  Tree (T)  2U  3.770  0.1571  39.27  H.S.  0.0236U  Level (L)  2  0.600  0.3000  75.00  H.S.  0.00575  Generation (G)  1  0.001  0.0010  -  N.S.  -0.00008  T x L  1*8  0.960  0.0200  5.oo  S.  0.00800  T x G  2h  0.330  0.0138  3.U5  s.  0.00326  L x G  2  0.010  0.0050  1.25  N.S.  o.oooou  U8  0.190  o.oouo  T x L x G  Hi9  Total  O.OOliOO  5.861  Table 12, shows a highly s i g n i f i c a n t difference among trees, and among levels. variance.  These r e s u l t s indicate the p o s s i b i l i t y o f s i g n i f i c a n t components of The s i g n i f i c a n t interactions indicate that the factors are not  independent and there i s no good b i o l o g i c a l e x p l a n a t i o n of the i n t e r a c t i o n s . The most economical sample numbers could be calculated from the components of variance f o r the f a c t o r s .  Minimum variance would be obtained i f we  take a large sample i n a large stratum, a large sample when the stratum  Table 13. Differences among trees by abundance of Adelges cooleyi. Generation 1 . I 1U II 0.62 III 19.hit 17 H.S. V M  2 0.21 3.56 s. M  15 0.18 1.98 N.S. M  8 0.17 1.59 N.S. M  13 0.16 1.19 N.S. M  12 0.15 0.79 N.S. M  21 O.OU -3.57 S. I  20 O.OU -3.57 s. I  9 o.ou -3.57 S. M  7 o.ou -3.57 S. M  2U 0.03 -3.97 S. I  19 0.03 -3.97 S. I  18 0.03 -3.97 S. I  16 0.03 -3.97 S. I  17 0.02 -U.37 S. I  5 0.22 2.23 s. M  22 0.22 2.23 s. 0  3 O.17 0.99 N.S. M  23 0.16 0.7U N.S. 0  2 o.U* 0.25 N.S. M  1 0.11* 0.25 N.S. M  8 0.13 0.00 N.S. M  6 0.12 -0.25 N.S. M  12 0.10 -0.7U N.S. M  15 0.10 -0.7U N.S. M  11 7 0 . 0 9 o.oi* -0.99 -2.23 s. N.S. M 0  9 o.ou -2.23 S. M  10 o.ou -2.23 s. 0  19 o.ou -2.23 s. I  20 0.03 -2.U9 s. I  21 0.03 -2.U9 S. I  2U 0.03 -2.U9 S. I  16 0.02 -2.72 s. I  18 0.02 -2.72 S. I  13 0.01 -2.97 S. M  17 0.01 -2.97 S. I  feneration 1 + 2 . I 11* U II 1 . 5 6 O.U 8 III 21.55 3.1a 17 H.S. s. M V M  22 0.U6 3.10 S. 0  5 o.U3 2.6U s. M  2 0.35 1.U0 N.S. M  3 0.32 0.93 N.S. M  23 0.30 0.62 N.S. 0  8 0.30 0.62 N.S. M  15 0.28 0.31 N.S. M  12 0.25 -0.16 N.S. M  1 0.25 -0.16 N.S. M  11 0.2U -0.31 N.S. 0  10 6 0.23 0 . 1 8 -0.1*7 -1.21* N.S. N.S. M 0  13 0.17 -1.U0 N.S. M  7 0.08 -2.79 S. M  9 0.08 -2.79 S. M  19 0.07 -2.95 s. I  20 0.07 -2.95 S. I  21 0.07 -2.95 S. I  2U 0.06 -3.10 s. I  18 0.05 -3.26 S. I  16 0.05 -3.26 S. I  17 0.03 -3.57 S. I  I II III 17 7  22 0.21* ii.37 S. 0  1* 0.23 3.97 s. M  5 0.21 3.56 S.  10 6 O.lit 0 . 1 1 0.U0 - 0 . 7 9 N.S. N.S. M 0  1 0.11 -0.79 N.S. M  I* 0.25 2.97 S. M  generation 2 . I lU II 1.03 III 22.28 H.S. 17 7 M I II III 17 7  I II III 17 .7  M  3 0.15 0.79 N.S. • M  11 23 O.U* 0.15 o.uo 0.79 N.S. N.S. 0 0 25 0.01 -U.76 s. I  25 0.00 -3.22 S. I  25 0.01 -3.88 S. I  35  Key f o r table 13: I " tree No. I I = average number of l i v i n g insects per needle. I l l - value of »t". IV = significance at 0 . 0 5 l e v e l . V  85  l o c a t i o n of the trees.  I = inside growth tree M 0 variance i s Mgh (level).  B  e  margin growth tree open-growth, tree  (trees) and a small sample when the stratum cost i s high  Because of the high cost of sampling at the several locations above  reach from the ground and adequate estimates of variance were obtained from the ground, further sampling probably should be concentrated on providing estimates of tree-to-tree v a r i a t i o n at the ground l e v e l of sampling. The differences between trees were examined by the " t " t e s t .  Table  13  shows that most of the h i g h l y populated trees are edge trees, and the lower populated trees are inside the stand.  By these r e s u l t s , the trees were  separated into three populations, i n t e r i o r , marginal and open trees.  The  average numbers of l i v i n g insects per needle of each tree population were tested against each other by the " t " t e s t (Table l h ) . Table 1U.  Influence of p o s i t i o n of tree i n the stand on abundance of Adelges  cooleyi. Location of trees.  Interior  Interior Marginal Open  Values of t Marginal  —  —  21.1»8 H.S. 5.19  Open  s.  —  —  2.13  N.S.  —  Table i i * shows that the abundance of the Adelges cooleyi on growing withi n a stand i s s i g n i f i c a n t l y d i f f e r e n t from those on the stand edge, or grown  Table 15. Differences among trees by abundance of separated by l o c a t i o n . Generation 1 . Margin growth trees. I 1 7 9 6 3 12 II o.olt 0.0k 0.11 0.11 0.15 0.15 III - 3 . 6 5 -3.65 - 1 . 8 2 -1.82 -0.78 -0.78 S. N.S. N.S. S. N.S. N.S. 17 E L L M L E 7 Inside growth trees. I 25 17 16 18 19 2lt II 0 . 0 1 0.02 0.03 0.03 0.03 0.03 III 0.00 0.00 0.00 0.00 -5.71 -2.86 N.S. N.S. N.S. N.S. S. 17 s. L E E L M L 7 Generation 2 . Margin growth trees. I 13 9 7 15 12 O.Oit II 0 . 0 1 O.Oit 0.10 0.10 III -2.lj8 -2.06 -2.06 -1.2U -1.2it s. N.S. N.S. N.S. N.S. 17 E L L L M 7 Inside growth trees. 18 16 I 25 17 2it II 0.00 0.01 0.02 0.02 0.03 III -2.15 -1.07 0.00 0.00 1.07 N.S. N.S. N.S. N.S. N.S. 17 E M L L L 7 Generation 1 + 2 . Margin growth trees. I 7 13 6 1 9 II 0.08 0.08 0.17 0.23 0.25 III -2.21 -2.21 -1.53 -1.07 -0.£2 N.S. N.S. S. N.S. S. 17 E M M L L 7 Inside growth trees. I 16 18 2it 25 17 II 0 . 0 1 0.03 0.05 0.05 0.06 III -It.60 - 2 . 3 0 0.00 0.00 1.15 S. N.S. N.S. N.S. N.S. 17 7 L L E M L  6  0.12 -0.97 N.S. E  Adelges cooleyi.  13 0.16  8  •0.52 N.S. M  0.17 -0.26 N.S. E  20 O.Oit 2.86  21 0.0U 2.86  S. M  S. E  8 0.13 •0.83 N.S. E 20  1  O.llt  -0.69 N.S. M  15  0.18 0.00 N.S. L.  2  O.llt  -0.69 N.S. L  5  0.21  2 0.21  0.78 N.S. E  0.78 N.S. L  3 0.17 -0.27 N.S. L  5 0.22  N.S. E  0.25 0.83 N.S. E  2 0.35 -0.15 N.S. L  5 0.U3 0.U6 N.S. E  0.U8 0.8lt N.S. E  O.ltl  It  0.23  1.30 N.S. E  it  lit 0.62  11.Ul H.S. E  lit  1.03 11.58 H.S. E  19  21 0.03 1.07 N.S. E  1.07 N.S. M  2.15 N.S. E  12 0.25 -0.92 N.S. E  15 0.28 •0.69 N.S. L  8 0.30 -0.53 N.S. E  21 0.07 2.30 N.S. E  20 0.07 2.30 N.S. M  19 0.07 2.30 N.S. E  0.03  The trees were treated as two  O.Oit  3  0.32  -0.38 N.S. L  U  U4 1.65 9.77 H.S. E  Key: I, I I , I I I , 17 = as i n tal 7,=.Bud opening time E = Early M = Medium L = Late  population,  i n the open.  No s i g n i f i c a n t difference was found between open and margin  growth trees. Then the trees were separated into two groups, i n t e r i o r and edge trees, omitting the open trees and differences between trees were sought, differences were indicated by the " t " t e s t (Table  1$).  S l i g h t differences can be seen between trees within the separated tree populations (Table 15).  Most of the trees, which were s i g n i f i c a n t l y more  populated than the average were e a r l y bud-opening trees, and most of those which were s i g n i f i c a n t l y l e s s populated were l a t e bud-opening trees.  Slight  differences were shown by " t t e s t between the l a t e and early-opening trees n  ( s i g n i f i c a n t at 0.3  l e v e l ) i n respect to the abundance of Adelges c o o l e y i .  The t e s t of differences between late-opening  and medium-opening trees, and  between medium-and early-opening trees could not be carried out, because of the small number of medium-opening trees. The 13 marginal trees had been chosen so 3 of them faced North, 3 East, 3 South and U West (Appendix 0).  These trees were analysed i n r e l a t i o n to  exposures and l e v e l s , leaving out one (No. 5) of the West trees (Table  16,  17 and 18). Table 16.  Average numbers of l i v i n g insects per needle, by exposures and l e v -  e l s f o r the marginal trees. Level  North  South  West  . East  Low  0.56  O.UU  0.51  0.91  Medium  0.33  o.i5  0.23  0.87  High  O.OU  o.ou  o.ou  0.32  Totals  0.93  0.63  0.78  2.10  Ave.  0.31  0.21  0.26  0.70  Both the l e v e l s and exposures show s i g n i f i c a n t differences (Table  17).  The relationship between number of l i v i n g insects per needle and height l e v e l  38  Table 1 7 . Analysis of variance among exposures and levels. Degrees . of freedom  Net sum squares  Mean sum squares  Variance ratio • (F)  Exposure (E)  3  0.U506  0.1502  15.98  H.S.  Level (L)  2  0.1*938"  0.21*69  26.27  H.S.  L xE  6  0.0566  0.009k  Total  11  1.0010  Source  on the living crown w i l l be examined later.  Significance at 0 . 0 5 level  The differences between exposures  were shown out by the least-significant-range test (Table 18). Table 18. Differences between differently exposed trees by the abundance of Adelges cooleyi. Exposures  Significantly different from  North  0.69-1.17  S, E  South  0.39-0.87  N, E  West  0.5U-1.02  E  1.86-2.3k  N,  East The  Ranges  S, W  abundance of Adelges cooleyi was examined in relation to length of  twigs, which showed a high degree of variation between trees.  The. .average  numbers of living insects per needle from the low level of the living crown of the trees were plotted over the average length of twigs from the low level of  the living crown (Appendix j ) (Fig. 1 0 ) .  The data measured on September 26  were used for calculating the average twig length.  It is given by the calcu-  lation of the equation of regression line that(Appendix I): b = 0 . 0 0 2 3 , a - 0 . 2 2 5 , r - 0.228, t = 0 . 1 1 2 , Sb = 0 . 0 2 0 1 3 , - 0 . 0 1 9  6  i - 0.065  As shown by the values of "r" and "t", no significant correlation was found between number of living insects per needle and twig length.  One of the  trees, No. 11*, shows a very high insect population on relatively short twigs,  and another of the trees, No. 10, has very long twigs with a r e l a t i v e l y small number of insects on i t .  Another c a l c u l a t i o n was carried out without these  two trees (Appendix I ) , where:  b = 0.058, a - - 0 . 0 8 , r - 0.680, t - k.25, Sb = 0.01375, 0.30 =" i * 0.086 Equation: Y - OJ0'58X-O.O8 This second c a l c u l a t i o n shows s i g n i f i c a n t c o r r e l a t i o n , meaning that the Adelges cooleyi u s u a l l y i s more abundant on the longer-twigged trees. The factors D.B.H. ( X l ) , height ( X 2 ) , age ( X 3 ) , crown width (Xk), of clear stem (X5), proportion  length  of l i v i n g crown to t o t a l height (X6), bud-  opening time (X7) and l o c a t i o n (X8) of the trees were related to the average number of l i v i n g insects per needle (Y) by analysis of l i n e a r - multiple regression.  This was carried out by the Alwac I I I electronic computer  (Appendix 0) (Table 1 9 ) . Table 19. Influence o f 8 factors on the average number o f l i v i n g insects per needle.  XI  Variable  X2  X3  Means  6.5b U5.20 22.80  Standard deviations  3.39  20.01  Minimum value  0.6  2  12.8 Maximum value 69 • Correlation coefficients 0.131 0.3U3 % of influence of each factor 5.73 28.31 R  2  =  0.625  xu  X5  X6  X7  X8  1U.68  1U.80  68.37  2.0k  k.kk  12.16  19.99  0.9k  3.1k  1  1  5.80  6.60  k  2  0  36.5  29  32  kl  99.9  0.269  0.58  0.125 O.U82 0.513 0.268 -8.82  9 0.261  -U5.57 89.63 12.00 -19.27  (R • multiple c o r r e l a t i o n c o e f f i c i e n t )  SEe = 0.197  . (SEe = standard error of estimate)  Equation: Y = -0.0k2Xl - 0.013X2 - 0.001X3 * 0.03kXk + -  0.025X5 + 0.028X6 -  0.155X7 - 0.076X8 - 0.993  Examining the c o r r e l a t i o n c o e f f i c i e n t s , only two factors show s i g n i f i c a n t  Ul  c o r r e l a t i o n with number of insects: the length of c l e a r stem and the percentage of l i v e crown.  The meaning of these two s i g n i f i c a n t r e s u l t s i s related to the  l o c a t i o n of trees.  In other words the c l e a r stems are longer on the i n t e r i o r  trees, consequently these trees have smaller percentages of l i v i n g crown. 3. I n t r a Tree Variations: The analyses of variance i n table 2 and 3 did not show s i g n i f i c a n t d i f ferences among the cardinal points, meaning that no further analysis i s needed for this factor. S i g n i f i c a n t differences were found among the d i f f e r e n t l e v e l s of the trees (Table 2, 12 and 1 7 ) .  The numbers of l i v i n g insects were summarized  and averaged f o r the three s p e c i a l l y sampled trees (No. U , 10 and 25)  and the  calculated average numbers of l i v i n g insects per needle were plotted by height l e v e l s (Appendix G) ( F i g . 11).  The linear-regression equations were calcu-  lated separately f o r the three sampling times (June 5, J u l y 25, September (Table 20, 21 and  26)  22).  Table 20. Calculation of the equation and c o r r e l a t i o n c o e f f i c i e n t of average number of l i v i n g insects per needle on height from the ground (data of June 5). X  Y  x  o 2  Y "  X Y  12.5  0.22  156.25  0.0U8U  2.750  17.5  0.37  306.25  0.1369  6.U75  22.5  0.27  506.25  O.0729  6.075  27.5  0.19  756.25  0.0361  5.225  32.5  0.10  1056.25  0.0100  3.250  37.5  0.11  LU06.25  0.0121  U.125  U2.5  0.03 0.02 0.02 0.00 1.33  1806.25 2256.25 2756.25 3306.25  0.0009 0.000U 0.000U 0.0000 6.3181  1.275 0.950 1.050 0.000 31.175  U7.5 52.5 57.5 350.0  1U312.50  Fig II  Relationship between number of living insects per needle and height on the living crown  June 5  325  375  42 5  Height from the Ground — Feet  ro  U3  X » height from the ground, Y = number of insects per needle, X =  35.00,  y - 0.13, b - -0.0075, a = 0.39, r - -0.901, t » -6.07, Sb = 0.0013, -0.0105 - i - -0.001;5 Equations Y = Table 21.  -0.0075X+0.39  Calculation of the equation and c o r r e l a t i o n c o e f f i c i e n t of average  number of l i v i n g insects per needle on height from the ground (data of July  25).  *-  Total Ave.  —y2  1  X  12.5  0.32  156.25  0.102U  U.000  17.5  0.27  306.25  0.0729  U.725  22.5  0.11  506.25  0.0121  2.U75  27.5  0.19  756.25  0.0361  5.225  32.5  0.01  1056.25  0.0009  0.975  37.5  0.01  1U06.25  0.0001  0.375  U2.5  o.ou  1806.25  0.0016  1.700  U7.5  0.03  2256.25  0.0009  1.U25  52.5  0.00  2756.25  0.0000  0.000  57.5  0.00  3306.25  0.0000  0.000  350.0  1.00  1U312.50  0.2270  20.900  35.0  0.10  2  XY  X = as i n table 20, Y - as i n table 20, b - -0.0068, a = 0.3U, r - -0.871,  t - -5.03, Sb » 0.001U, -0.100 Equation: Y =  6  i  6  -0.0036  -0.0068X+0.3U  Each of the three analyses (Table 20, 21,  22)  show; s i g n i f i c a n t c o r r e l a -  t i o n between average numbers of l i v i n g insects per needle and height from the ground.  The insect numbers decreased with height i n every instance. *  The change of twig lengths with height on crown was also studied.  The  kk  Table 22.  Calculation of the equation and c o r r e l a t i o n c o e f f i c i e n t of average  number of l i v i n g insects per needle on height from the ground (data of September 26). X2  Y2  XY  156.25  0.032k  2.250  0.20  306.25  0.0k00  3.500  22.5  0.10  506.25  0.0100  2.250  27.5  0.11  756.25  0.0121  3.025  32.5  0.06  1056.25  0.0036  1.950  37.5  0.00  lk06.25  0.0000  0.000  U2.5  0.02  1806.25  o.oook  0.850  U7.5  0.02  2256.25  o.oook  0.930  52.5  0.01  2756.25  0.0001  0.525  57.5  0.00  3306.25  0.0000  0.000  350.0  0.70  lk312.50  0.0990  15.280  35.0  0.07  X  I  12.5  0.18  17.5  X = as i n table 2 0 , I = as i n table 2 0 , b = - 0 . 0 0 k 5 , a = 0.23, t - -6.05, Sb • 0.0007, -0.0061 * i  £  r = -0.907,  -0.0029  Equation; Y » -O.OOk5X+0.23 lengths of twigs were summarized and averaged f o r two trees (No. h and 10) 5 feet i n t e r v a l s (Appendix G).  Then the average twig lengths were plotted by  height l e v e l s from the ground ( F i g . 1 2 ) . rather than a straight l i n e . to logarithms ( F i g . 12)  (Table  by  The plotted points show a curve  For further analysis twig length was  transformed  23).  The c u r v i l i n e a r correlation i s s i g n i f i c a n t between the height l e v e l s and twig lengths (Table The trees No. k,  23). 10 and 25 were also examined h o r i z o n t a l l y at 3-feet  i n t e r v a l s r a d i a l l y from the bole at the middle of the low l e v e l of the l i v i n g  Fig-12 I36r-  Relationship between twig length and height on the living crown L  1*6  Table 2 3 . Calculation  of the equation and  c o r r e l a t i o n c o e f f i c i e n t of average  twig lenth on height from the ground. X  Y  logY  12.5  20.0  1.301  17,5"  20.3  22.5  (logY)*  X(logY)  156.25  1.693  16.26  1.307  306.25  1.708  22.87  19.1*  1.278  506.25  1.633  28.76  27.5  19.5  1.290  756.25  1.661*  35.1*8  32.5  20.6  1.3H*  1056.25  1.727  1*2.71  37.5  19.9  1.299  11*06.25  1.687  1*8.71  1*2.5  20.9  1.320  1806.25  1.71*2  56.10  1*7.5  21.1  1.321*  2256.25  1.752  62.89  52.5  22.2  1.3U6  2756.25  1.812  70.67  57.5  23.0  1.362  3306.25  1.855  78.32  206.9  13.11*1  11*312.50  17.273  1*62.77  20.7  1.311*  350.0 35.0  .  X = height from the ground, Y = average twig length of the height l e v e l , b = 0.00137, a = 1.266, r = O.986, t = 16.08, Sb = 0.0000695, 0.00121 * i * 0.00153 Equation: logY - 0.00137X+1.266 crowns.  The numbers of i n s e c t found were summarized, and  averaged (Appendix G)  Then the average numbers of l i v i n g insects per needle were plotted over the distances from the stem ( F i g . 1 3 ) , and  the regression analyses were carried  out separately f o r the three d i f f e r e n t sampling times (June 5 , July 2 5 , September 26) The  (Tables: 21*, 25 and  correlations  26).  are not s i g n i f i c a n t at the 0.05 l e v e l i n tables 21* and  25, which might also be caused by the small sample s i z e s . l a t i o n was  S i g n i f i c a n t corre-  found i n table 26 between the numbers of l i v i n g insects per needle  and distances from the stem.  hi Fig  13 R e l a t i o n s h i p needle  between number  and distance from  of living insects per  t h e stem a t the low  level of the living c r o w n  Distance from the Stem —Feet  °  U8  Table 2k. Calculation of the equation and c o r r e l a t i o n c o e f f i c i e n t o f average numbers of l i v i n g insects per needle on distances from the stem (data of June 5 ) . X  Y  1.5 U.5  3  XY  <  o.ooou  0.030  . 0.09  20.25  0.0027  o.i405  0.16  56.25  0.0256  1.200  10.5  0.2h  110.25  0.0576  2.520  13.5  0.10  182.25  0.0100  1.310  37.5  0.61  371.25  0.0963  5.505  7.5  0.12  Ave. X - distances = 0.010,  Y  0.02  7.5  Total  o  x2.25  from the stem, Y = numbers of l i v i n g insects per needle, b =  a = O.OU, r = 0.66U, t - 1 . 5 U , Sb = 0 . 0 0 6 7 5 ,  £  -0.011 * i 0 . 0 3 1  Equation: Y = 0.010X+0.0U Table 2 5 . Calculation of the equation and c o r r e l a t i o n c o e f f i c i e n t o f average numbers of l i v i n g insects per needle on distances  from the stem (data of  J u l y 25).  x  y  1.5  0.01  2.25  0.0001  0.015  U.5  0.03  20.25  0.0009  0.135  7.5  o.iU  56.25  0.0196  1.050  10.5  0.10  110.25  0.0100  1.050  13.5  0.15  182.25  0.0225  2.025  37.5  0.U3  371.25  0.0531  U.275  1  Total Ave.  7.5  x — 5  1  XY  0.09  X = as i n table 2 U , Y = as i n table 2 U , b - 0 . 0 1 2 , a = 0 . 0 0 , r - 0.861, t » = 2 . 9 3 , Sb = 0 . 0 0 U 2 1 , - 0 . 0 0 1 * i  s  0.025  Equation: Y = 0 . 0 1 2 X  U9 Table 26. Calculation of the equation and correlation c o e f f i c i e n t of average number of l i v i n g insects per needle on distances from the stem (data of September 26). X  Y  f  1  f  1  XY  1.5  0.03  2.25  U.5  0.03  20.25  0.0009  0.135  7.5  o.m  56.25  0.0196  1.050  10.5  0.12  110.25  0.011*1*  1.260  13.5  0.25  182.25  0.0625  3.375  37.5  0.57  371.25  0.0983  5.865  7.5  0.11  0.0009  o.oh5  X = as .in table 21*, Y = as i n table 21*, b = 0.018, a = -0.03, r = 0.911*, t • - 3.96, Sb = 0.001*35, 0.001* * i * 0.032 Equation: Y = 0.018X-0.03 Table 2 7 . Calculation of the equation and correlation c o e f f i c i e n t o f average lengths of twigs 'on distances from the stem.  Total Ave.  t  Y  1.5  10.1*  2.25  I*.5  12.9  7.5  X  2  1  ^ 1  XY  108.16  15.60  20.25  166.1*1  58.05  17.5  56.25  306.25  131.25  io.5  20.8  110.25  1*32.61*  218.1*0  13.5  22.9  182.25  521* .Ul  309.15  37.5  81*.5  371.25  1537.87  732.1*5  7.5  16.9  X = distances from the stem i n feet, Y = average lengths of twigs, b • 1.096, a - 8.7, r = 0.993, t = 11*.56, Sb = 0.0765, 0.853 - i - 1.339 Equation: Y - 1.096X+8.700  50  Fig-14  Relationship stem  between twig length and d i s t a n c e f r o m  at t h e low level of t h e l i v i n g  crown  the  51  The twig lengths were also measured every 3 feet h o r i z o n t a l l y from the stem, at the middle of the low l e v e l of the l i v i n g crowns (Appendix G).  The  average twig lengths were plotted over distances from the stem ( F i g . Hi) (Table 27).  S i g n i f i c a n t l y l i n e a r relationships were found between the twig  lengths and distances from the stem. h. M o r t a l i t y : As mentioned, the accurate numbers of dead insects could not be counted, because many of them f e l l o f f the needles, and a large proportion of them was eaten completely by predators.  Assuming that the p r o b a b i l i t y of l o s s of dead  insects was s i m i l a r everywhere, the analysis of data w i l l give some informat i o n about the f a c t o r s influencing mortality. Trees No. 1, 2, 10, 11, 1U and 20 were examined by two analyses of variance, separately f o r generation 1, and generation 2 (Appendix M) (Tables  29).  28 and  Table 28. Analysis of variance of dead insects among trees l e v e l s and exposures (generation 1 ) . Degrees of freedom  Net sum squares  Mean sum squares  Variance ratio (F)  Level (L)  5  0.19  0.095  23.75  H.S.  Exposure (E)  3  0.02  0.007  1.75  N.S.  Tree  5  3.20  0.6L0  L x E  6  0.07  0.012  3.00  S.  L x T  10  0.12  0.012  3.00  S.  E x T  15  0.10  0.007  1.75  N.S.  L x T x E  30  0.13  0.001*  Total  71  3.83  Source  (T)  Significance at 0.05 l e v e l  160.00  Both the analyses of variance (Table 28 and 29) show significant-  H.S.  52  Table 29.  Analysis of variance of dead insects among trees, l e v e l s and  exposures (generation Source  2).  Degrees of freedom  Net sum squares  Mean sum squares  Variance ratio (F)  ,Significance at 0.05 l e v e l :  2  0.032  0.0160  26.67  Exposures (E)  3  0.005  0.0017  2.83  N.S.  Trees (T)  5  0.130  0.0260  U3.33  L x E  6  0.002  0.0003  -  H.S.  L x T  10  O.OLU  0.001L  2.33  S.  E x T  15  0.015  0.0010  1.67  N.S.  L x E x T  30  0.019  0.0006  Total  71  0.217  Levels  (L)  difference within the l e v e l s and within the trees. was  found among the exposures.  ]  H.S.  N.S.  No s i g n i f i c a n t difference  The s i g n i f i c a n t difference i n the interactions  indicates that the e f f e c t s of the factors were not independent.  The L x T  i n t e r a c t i o n maybe s i g n i f i c a n t because the actual heights of the sampling l e v e l s were d i f f e r e n t from tree to tree (Appendix E ) .  These differences  can  be seen well between the marginal and i n t e r i o r trees (e.g. the height of  low  l e v e l of tree No. U was 19 f e e t , and of tree No. 25 was U5 f e e t ) . A l l the 25 sample trees were also examined by an analysis of variance, where the data of exposures were used as r e p l i c a t i o n s (Appendix N) 30).  (Table  A l l of the f a c t o r s , trees, l e v e l s and generations, are s i g n i f i c a n t , and  a l l of the interactions are also s i g n i f i c a n t . These r e s u l t s prompted another analysis, testing whether or not the main e f f e c t s are s i g n i f i c a n t l y greater than the first-rorder interactions (Table 31).  The e f f e c t s of the main factors  were s i g n i f i c a n t l y greater than the f i r s t - o r d e r i n t e r a c t i o n s . The percentages of mortality were tested to determine whether or not marginal trees with many insects had  the  a higher percentage of insect mortality  53  (Table 32) (Appendix PJ. Table 30. Analysis of variance of dead insects among trees, levels! and generations. Variance ratio (F)  Net sum squares  Mean sum squares  2l*  0.86  0.0358  UU.75  H.S.  0.001*86  Levels (L)  2  0.09  0.01*50  56.25  H.S.  0.00087  Gen. (G)  1  0.23  0.2300  287.50  H.S.  0.00298  T xL  1*8  0.09  0.0019  2.38  S.  0.00055  T xG  21*  0.1*0  0.0167  20.87  H.S..  0.00530  L xG  2  0.01  0.0050  6.25  S.  0.00017  1*8  o.ol*  0.0008  11*9  1.72  Trees (T)  T x L xG Total  Significance at 0.05 l e v e l  1 Components o f variance  Degrees of freedom  Source  0.00080  Table 31. Variance r a t i o test between the main factors and f i r s t - o r d e r  inter-  actions . Source  Degrees of freedom  Net sum squares  Mean sum squares  Variance ratio (F)  Significance at 0.05 l e v e l  5.26  H.S.  22.50  H.S.  H*.65  H.S.  Trees  21*  0.86  0.0358  T x L + TxG  72  0.1*9  . 0.0068  2  0.09  0.01*50  50  0.10  0.0020  1  0.23  0.2300  26  0.1*1  0.0157  Levels T x L +L xG Generations T x G +L xG  The degree of mortality (Table 32) was lower i n generation 2 than i n generation 1.  The percentage of mortality was lower on the lower populated  trees i n generation 1, but t h i s was reversed i n population 2. The percentage of mortality by height on the l i v i n g crown was also  5U  calculated (Table 33). Table 32. Percentage of mortality by locality of the trees and generations.  Trees  Average number of dead insects per needle marginal interior  Average number of living+dead insects per needle marginal interior  Percentage of mortality marginal  interior Uo.o  Generation 1  0.16  0.02  0.3U  0.05  1*7.1  Generation 2  0.05  0.01  0.2U  0.03  20.8  • 33.3  Gen. 1 + Gen. 2  0.21  0.03  0.58  0.08  36.2  37.5  Table 33. Percentage of mortality by levels and generations. Levels Generations  Average number of dead insects per needle 2 1  Average number of living +dead insects per needle  Percentage of mortality  1  2  1  2  Low  0.15  0.05  0.35  0.25  U2.9  20.0  Medium  0.12  o.ol*  0.27  0.18  uu.u  22.2  High  0.07  0.02  0.11  0.06  63.6  33.3  The percentage of mortality increased with height (Table 33). 5. Need for Further Studies; A few questions might arise about the sample sizes for further studies, i . How many trees w i l l be required to determine the population of living and dead insects within 5% of the mean nineteen times out of 20? On the average determined from sampling 25 trees, 0.26 living insects were found on one needle, the standard deviation (SD) was 0.3227 and the standard error of the mean (SEM) was 0.0065.  From these the required sample  size w i l l be 2,U6U trees. The mean of the dead insects found was 0.15 per needle with SD = 0.1536 and SEM = 0.0038.  The sample size of trees required to determine the dead  insects was found as 1,63U trees.  55  i i . How many twigs w i l l be required within a l e v e l to determine the population of l i v i n g and dead insects within $% of the mean, nineteen times out of  20?  Four twigs were sampled i n each l e v e l i n this experiment (from North, West, South and E a s t ) .  In an average of 6 trees (No. 1,  2, 10,  11,  l k and 20)  0.35 was the number of l i v i n g insects per needle on a twig, with SD = 0.0565 and SEM = 0.0088.  The required sample size was found as k l twigs per l e v e l to  determine the population of l i v i n g i n s e c t s . The average number of dead insects per needle i n a twig was 0.17 SD =  0.0129 and SEM = 0.00U3.  with  The required sample size was calculated as  9  twigs per l e v e l . i i i . How many needles are required within a twig to determine the population of l i v i n g and dead insects within $% of the mean nineteen times out of 20?  This should be calculated f o r d i f f e r e n t l e v e l s , but here w i l l be shown  only one example. The mean of the number of l i v i n g insects at low l e v e l and North side i n generation 1 of tree No. 1, A t o t a l of  on June 1; was 0.1*0  with SD = 0.52  and SEM = 0.010.  2,70k needles per twig were calculated as the required sample s i z e .  The average number of dead insects per needle was 0.10 time as before, with SD =  0.32 and SEM = 0.0025.  Then  at same place and  16,38k needles per twig  were calculated as the required sample s i z e . These sample s i z e s seem very large, but these numbers would decrease i f the l i m i t s of error were changed. trees was mean.  For example, the required sample size of  2,k6k to determine the population of l i v i n g insects within $% of the  Changing the l i m i t s of error to 10$,  with 20$ e r r o r , only 15k Considerable  616 trees would be required, and  trees would be needed.  influence on the sample also r e s u l t s from changing the  l e v e l s of confidence  e.g. only 150  trees would be required to determine the  population  of l i v i n g insects within 5 % at the 0.20 confidence l e v e l .  Using analyses of variance  to determine the influence of various  factors  may also reduce the need f o r such large numbers because factors themselves provide r e p l i c a t i o n and p a r t i t i o n i n g of sum squares may present a very s e n s i tive e r r o r term.  57  DISCUSSION As the former chapters show,during the summer of  I960, two  main groups of  factors were examined: i n t e r - t r e e differences and i n t r a - t r e e differences.  Two  main groups of f a c t o r s w i l l also be discussed here: the mortality and b i o t i c potencial which were only p a r t l y investigated. 1. Inter Tree Differences: The power of m u l t i p l i c a t i o n of Adelges cooleyi i s affected by external and i n t e r n a l conditions of the tree.  I t i s not always easy to determine  whether external or i n t e r n a l conditions are responsible, but there i s considerable evidence that some trees are more favourable insect than others.  to the m u l t i p l i c a t i o n of the  The inter-tree, differences can be divided i n t o several  groups. Location of the  Trees:  I t has been stated by Chrystal ( 7 )  that i n a dense wood of Douglas f i r ,  Adelges cooleyi occured c h i e f l y on the trees near to the margin of the f o r e s t and the i n s e c t was not abundant i n the centre of the plantation. Twenty-five trees were examined around Totem Park during the summer of I960, 8 of them inside the stand, 13 on the margin and h of them i n the open. I t was  found that by analysis of variance, (Table 12) the trees were populated  d i f f e r e n t l y by Adelges c o o l e y i .  The Student's " t " d i s t r i b u t i o n showed that  generally the s i g n i f i c a n t l y more populated trees are growing at the margin of the f o r e s t , and the s i g n i f i c a n t l y l e s s populated trees are growing at the center of the stand (Table 13).  Then the trees were separated  into 3 groups  (edge-, i n s i d e - and open-growth) and the groups were tested against each other by " t " t e s t s .  These tests showed abundance to be s i g n i f i c a n t l y greater on the  marginal and open trees than on the inside trees.  No differences were found  at the 0 . 0 5 l e v e l between open and marginal trees. These differences might be explained by several external conditions of  58  the tree.  The differences between crown shapes or percentage of living crown  seem to be the main reason. The proportion of living crown i s much smaller on an interior tree than on a marginal tree (Appendix L and 0). As shown byseveral s t a t i s t i c a l analyses (see chapter: Experimental Results), the abundance of insects i s low at the tops of trees.  Comparing the abundance of  Adelges cooleyi on tree No. U to tree No. 25 (Fig. 15),  the degree of a-  bundance does not show higher level on tree No. ii (edge growth) than on tree No. 25 (inside growth) above UO feet from the ground, where the living crown begins on the interior growth tree.  Consequently i t i s very l i k e l y that the  lack of the lower part of the living crown of the inside-growth trees i s one reason for the differences. The different abundance of the insect on interior and marginal trees might be explained by the dispersal of the Adelges cooleyi.  Chrystal (7) and  Cameron (5) stated that the chief factor of the dispersal of the Adelges cooleyi i s wind. stand (15,  27).  The speed of wind i s always close to zero within a dense The different abundance of the insect might be affected also  by the different wind pattern, i n which the margin of the stand reduces the wind speed and would f i l t e r out the transported insects. Some other climatic factors such as temperature, light and relative humidity differences may influence the different abundance of iL cooleyi on the edge and inside-growth trees. Exposure; Some differences were found between the differently exposed marginal trees (Table 16, 17 and 18). These differences may be the result of chance only, because only three trees were sampled at each exposure.  One thing can  be concluded here, the South and West edges of the forest were less populated than the North and East edges, which might be caused by two reasons. The West and South edges of the forest border on roads, where the needles of the trees  Fig  15  Relationship between the number  of living insects  ( A = interior growth tree , B = marginal growth  a>  per needle  a n d height on t h e living crown  tree)  a> 0 4 r CP  a. * 0-3 CD CO  _c  c? 0 2  o 6  a> > <  01 _L  15-20 20-25 25-30 30-35 35-40 40-45 45-50 50-55 5 60 Height from the Ground on Tree No 25 — Feet  0-5  5-10  10-15  0-5  5-10  10-15 15-20 20-25 25-30 30-35 35-40 40-45 45-50 50-55 55-60 Height from the Ground on Tree No-4 — Feet  60  are covered with a considerable amount of dust, which may be unfavorable to the i n s e c t .  The other reason would be that only suppressed  able f o r study i n the East margin of the f o r e s t .  trees were a v a i l -  The high l e v e l of abundance  on the East side might be caused by the low vigour and the r e l a t i v e l y small height of the t r e e s . The e f f e c t s of p r e v a i l i n g winds - which are NW with a considerable amount of SE i n coastal B.C.  (21)  Adelges cooleyi developed  - should also be examined.  But i n Totem Park  on the summer of I960 only with the subcycle of the  l i f e cycle, i n where the wind was not an important part of the natural d i s persal. Bud-Opening Time; Table 1 5 shows that the s i g n i f i c a n t l y more populated trees were mostly early-opening t r e e s , most of the less-populated trees were late-opening. differences were found both within the edge and i n t e r i o r t r e e s .  These  Only s l i g h t  differences were shown ( s i g n i f i c a n t at 0 . 3 l e v e l ) between e a r l y and l a t e  bud-  opening trees. The reason f o r these differences might be looked f o r i n the seasonal development of Adelges cooleyi .  The hatching of the progeny of sistens gener-  ation was j u s t about over when the early-opening trees started to s w e l l .  The  progredientes and sexuparae s e t t l e more r e a d i l y on the new f o l i a g e , but they could not s e t t l e on the new f o l i a g e of late-opening trees u n t i l 2 - 3 after hatching.  During t h i s 2 - 3  weeks  weeks period the larvae were found on the  surface of the opening buds, waiting f o r the swelling. Many of them died from lack of food, and the lack of s t a b i l i t y on the opening buds.  They were more  exposed to climatic factors on the buds than they were on the lower surface of the needle. One question may be raised here.  Why  i s the l i f e cycle of the Adelges  cooleyi not synchronized to the d i f f e r e n t bud-opening trees?  I t i s true that  61  a l i t t l e displacement i s seen i n the l i f e cycle on the late-opening which (Appendix F) might be caused by the delay i n food supply. l i k e l y that there was  not any synchronization  trees,  I t i s more  between the l i f e cycle of the  insect and the l i f e cycle of the l a t e bud-opened trees, because the sexuparae migrate to S i t k a spruce, and the Sitka spruce has not so wide range i n bud swelling, as Douglas f i r ( 1 6 ) . be synchronized.  For t h i s very reason three l i f e cycles should  Consequently the l i f e cycle of Adelges cooleyi seems to be  determined by two f a c t o r s , the growth cycle of Douglas f i r and the growth cycle of S i t k a spruce, and being the primary host S i t k a spruce^ on which the bisexual forms and reproduction  occur, might be the more i n f l u e n t i a l f a c t o r .  Twig Length; Considerable v a r i a t i o n was of the growing period.  found between trees i n twig length at the  A s l i g h t c o r r e l a t i o n was  end  found between length of twigs  and the abundance of the insects ( F i g . 1 0 ) . The s t y l e t of Adelges cooleyi i s inserted into the spongy mesophyll t i s sue during most of the time spent on Douglas f i r .  I t appears that the  twigged trees have more lush growth, which i s more favorable  longer-  to the insect.  Adelges cooleyi would prefer- the lush growth, because the s t y l e t s would be more e a s i l y inserted into the lush-growth needles.  For another reason the  insect would be favoured by the lush twigs because the solution of the food given by such twigs and needles can be more n u t r i t i o n s f o r the insects. Some general factors as the D.B.H., height, crown width, age, percentage of l i v i n g crown of the t o t a l height, bud-opening time and place of the tree were also examined. These factors were analysed by l i n e a r - multiple - regression (Table 1 9 ) .  Two  analysis  of the eight f a c t o r s , the percentage of l i v i n g crown and  the  length of c l e a r stem^showed s i g n i f i c a n t c o r r e l a t i o n with the abundance of the insect.  Both of these factors mean the same thing: the difference between the  62  marginal or open trees and interior trees.  The trees, having a high percent-  age of living crown (Appendix L) were highly populated by Adelges cooleyi. These trees are marginal or open trees, having comparatively  short clear stem.  The trees having low percentage of living crown were lightly populated by the insect.  These trees are growing at the center of the forest, having compara-  tively high  length of clear stem.  The total height of the trees has a considerable percent of influence (28.32) (Table 1 9 ) , which means that the interior trees are a l i t t l e higher than the outside trees. The location of the trees should be significantly correlated to the abundance of the insect. The lack of significant result was caused by the fact that the trees were not separated as edge and inside trees i n this analysis, but were numbered by different exposures ( 2 - 8 )  and open-growth ( 9 ) and i n -  side-growth ( 1 ) (Appendix 0). The age of the trees as a factor should also show correlation with the abundance, because both Chrystal (7) and Cameron (h, 5) have stated that Adelges cooleyi affects young trees more seriously than older trees.  This  difference could not be shown in this experiment, because the: range in age was very small. 2. Intra Tree Variations: Exposure: The trees were sampled from four sides, North, West, South and East.  No  statistically significant differences appeared between the different sides. This equal abundance of the insect by cardinal points seems to imply only minor microclimatic differences at different cardinal points.  Examining a  single marginal tree, some differences were seen, the open part of the tree had more insects, than the other parts, but i t was more l i k e l y influenced by crown shape than cardinal point.  63  Height on L i v i n g Crown: Chrystal ( 7 ) has stated that the i n t e n s i t y of attack varies by height. This v a r i a t i o n was also observed i n Totem Park.  I t was shown by analyses of  variance, that low, medium and high l e v e l s of l i v i n g crown were s i g n i f i c a n t l y d i f f e r e n t i n abundance of the Adelges cooleyi (Table 2 , 1 2 and 1 7 ) .  Also a  s i g n i f i c a n t c o r r e l a t i o n was found between the abundance of insects and height from the ground.  The number of insects per needle decreased l i n e a r l y with  height (Table 2 0 , 2 1 and 2 2 ) ( F i g . 1 1 ) . Teucher (26) has mentioned, that microclimate i s the main factor a f f e c t ing the i n t e n s i t y of attack, and within the microclimate: temperature and wind are the most important.  Geiger ( 1 5 )  and Botvay ( 3 ) stated that the tempera-  ture during day time i s always higher above and i n the upper crown, than above the f o r e s t f l o o r , and the r e l a t i v e humidity at the same time i s always lower above and i n the upper crown than above the f o r e s t f l o o r . ence i n temperature i s from 0.5°C to 1.0°C, and i n R.H.  The average d i f f e r -  is 3 - h % (15).  These differences to a smaller degree can be observed foot by foot' from the f o r e s t f l o o r to the top of the trees ( 3 ) .  I t might be considered that the  abundance of Adelges cooleyi i s influenced by these microclimatic differences. I t seems to be suggested by the f a c t that the percentage of mortality was higher  i n the medium l e v e l than the low l e v e l , and i n the high l e v e l than  the medium and low l e v e l s (Table 3 3 ) .  The number of unhatched eggs was l a r g e r  i n the medium l e v e l than i n the low l e v e l , and more i n the high l e v e l than i n the medium l e v e l .  Percentages of unhatched eggs were: low l e v e l ; 2 5 . 9 %>  medium l e v e l : 3 3 . 8 % and high l e v e l : 3 7 . 0 %. Changes i n Abundance of Insect i n Horizontal Directions of the Tree Crown: I t was shown by regression analyses that s l i g h t c o r r e l a t i o n can be found between abundance of insect and the distances from the stem at the middle of the low l e v e l of l i v i n g crown (Table 2 U , 2 5 and 2 6 ) ( F i g . 1 3 ) .  The number of  6U  insects per needle increased toward the periphery.  I t may be supposed that  t h i s also resulted from microclimatic differences, or Adelges cooleyi looks i s l e s s abundant on the shaded twigs.  Considerable  differences might be supposed  i n food supply between exposed and shaded twigs, since the needles are noticeably thinner on h e a v i l y shaded branches. Twig Length; As was mentioned, Adelges cooleyi s e t t l e s more abundantly on the needles of longer twigs.  This r e s u l t might be applied to prove the horizontal d i s t r i -  bution of the insect within a t r e e , because as the lengths of the twigs i n creased from the stem to the outer crown (Table 2 7 ) ( F i g . LU), so increased the frequency of the i n s e c t . Examining the v e r t i c a l d i s t r i b u t i o n of the i n s e c t and twig length, the r e s u l t was  opposite.  The length of twigs was  increasing from the bottom of  the l i v i n g crown to the top by a semi-logarithmic  curve (Table 23) ( F i g . 12),  and the numbers of insect per needle were decreasing l i n e a r l y from the bottom to the top of the l i v i n g crown (Table 2 0 , 21 and 2 2 ) ( F i g . 11).  This r e s u l t  seems to show that the twig length, which i s one of the many factors a f f e c t ing  the v e r t i c a l abundance of Adelges c o o l e y i , was not so e f f e c t i v e as others  such as microclimatic f a c t o r s . 3. Mortality; As mentioned, the number of dead insects could not be counted accurately. This unfortunate  f a c t can be seen w e l l i n F i g . 7 and 8 , where the mortality  curves should follow the curves of l i v i n g insect but i n the reverse d i r e c t i o n . I t can be seen from the diagrams that the l o s s of dead insects occurred mostly a f t e r the hatching time, when the mortality showed the highest p o t e n t i a l . This period appears to be the c r i t i c a l stage of generation 2 , when the newly hatched larvae were crawling over needles preparatory to s e t t l i n g down f o r overwintering.  In tfe-is period the larvae were not settled i n a special place,  65  the mouth-parts were not inserted i n t o the mesophyll t i s s u e , and consequently they were more exposed to climatic and b i o t i c e f f e c t s . I t might be considered that the high mortality i n generation 2 was caused by climatic f a c t o r s .  Examining the meteorological records (Appendix K) of  J u l y and August, the temperature was sometimes extremely high i n July, and p r e c i p i t a t i o n low, with 0 . 0 3 inches r a i n i n the whole month. the average temperature was very low i n August. 5 8 . 3 2 ° F , and the average minimum was 5 5 . 3 8 ° F .  Following these  The average maximum was  The average maximum and minimum  temperature f o r August i n a UO-years average up to 1 9 5 5 i n Vancouver was 7 3 ° and 5U°F ( 2 1 ) .  The difference between the average maximums i s about 15°F,  which seems to be high enough to a f f e c t the newly hatched larvae.  The l o s s  of dead insects i n t h i s period can be explained by the f a c t that the larvae were not yet s e t t l e d f o r overwintering. The loss of dead insects cannot be seen i n generation 1 , because one part of t h i s generation was winged.  The decrease of the number of l i v i n g insects  per needle was l i n e a r ( F i g . 5 and 6 ) and the increase of the number of dead insects per needle was also l i n e a r , but the slope of t h i s l i n e i s not proven, because of loss of dead i n s e c t s .  The two l i n e s , the l i n e of l i v i n g insects  and the l i n e of m o r t a l i t y are not balanced, because of the loss of the winged part of the generation, and the l o s s of a part of the dead insects, and the proportion of these two were not or could not be observed. Examining the l i n e of l i v i n g insects, the c r i t i c a l stage did not appear, or i t was not observed, because the observation started only 2 - 3 weeks a f t e r the hatching of generation 1 . I t was shown by analyses of variance (Table 28, 2 9 , 3 0 and 3 1 ) that the number of dead insects per needle was s i g n i f i c a n t l y d i f f e r e n t among trees, among l e v e l s and among generations.  These results indicate that the absolute  number of dead insects was higher i n such a place where the number of l i v i n g  66  insects was higher. A question can be raised at t h i s point, whether the degree of m o r t a l i t y was l a r g e r on the higher-populated  trees or not.  I t was found that (Table 32)  the percentage of mortality was higher within the marginal trees than within the inside trees i n population 1.  The r e s u l t was the reverse i n population 2.  Within a tree, the degree of mortality was increased by height, where the absolute numbers of l i v i n g and dead insects were decreased.  So with one  exception,the percentage of m o r t a l i t y was higher on the places where Adelges cooleyi was l e s s abundant.  I t seems to be that the circumstances o f the l e s s -  populated places were not suitable f o r the insect, because the degree of mortality was higher.  On the other hand the population of Adelges cooleyi  was not so high, even within the heavily populated places that the competition f o r food or place would play an important part i n mortality. The one exception which was found i n population 1, where the percentage mortality was higher on the heavily populated trees, seems to be caused by the l o s s of winged foufis of the generation.  The l i v i n g winged adults were l o s t ,  but the dead ones were counted. k. Natural Enemies; No s p e c i a l studies were conducted on how the natural enemies a f f e c t the abundance of Adelges c o o l e y i .  I t was only noticed that the most important of  the natural enemies were the black and yellow ladybird beetles (Coccinellidae) which frequently feed on Adelges cooleyi. Syphids which are mentioned to be enemies to the Adelges cooleyi ( 1 0 , 11) were found, but they were not so common as the ladybird beetles. One species of Hymenopteran parasite was found, which developed abdomen of both progredientes and sexuparae.  i n the  They were not common, only two  specimens being found during the summer of I 9 6 0 . A large number of Spiders and Red - Spider - Mites occurred around the  67  settled Adelges and l e y i was  not  t h e i r egg  c l u s t e r s , but t h e i r e f f e c t on the Adelges coo-  certain.  5. B i o t i c  Potential;  Since reproduction i s by c y c l i c a l parthenogenesis, the potential rate of reproduction i s not l i m i t e d by the necessity of mating ( 2 , 1 9 ) which i s believed to be a main factor contributing  to the abundance of Adelges cooleyi;  bisexual reproduction i s necessary i n only one  generation out of the  five.  68  SUMMARY The study took place i n the 25-year-old Douglas-fir stand i n Totem Park. Twenty-five trees were sampled during the period June U to September 2k. sampling was repeated 13 times and 12 at each survey.  The  sample twigs were drawn from every tree  The numbers of l i v i n g and dead insects were observed on ten  needles from a twig.  Thus 3,900 twigs were cut down, and on 39,000 needles  the number of insects was observed, and approximately 10,lk0 l i v i n g and 5,850 dead insects were counted. The growth of foliage and the height and D.B.H. of the sample trees were measured.  The crown shape of the 25 trees was studied.  The climatic f a c t o r s ,  e.g. temperature, RH, p r e c i p i t a t i o n , evaporation, wind and t o t a l hours of sunshine,were  also studied from available data, during the period A p r i l to  September. The collected data were examined by s t a t i s t i c a l analyses. Eight analyses of variance, 17 regression analyses, and other s t a t i s t i c a l tests were carried out.  From the r e s u l t s of the s t a t i s t i c a l analyses the inter-and intra-tree  differences i n abundance of Adelges cooleyi ( G i l l . ) were studied.  The mor-  t a l i t y of the insect was also examined i n r e l a t i o n to several factors.  69  CONCLUSIONS  Findings of the.study are summarized as follows: 1. Adelges cooleyi was more frequent on the marginal and open grown trees, than on trees growing i n the center of the stand. 2. The insects were less abundant on trees growing at the edge of roadway than on trees exposed to other d i r e c t i o n s . 3. The insects were s l i g h t l y l e s s abundant on the late budopening trees, than on the e a r l y bud-opening, both on trees grown i n the t e r i o r and on the margin of the stand. k. The frequency of Adelges cooleyi was greater on the long twigged trees. 5. Within a tree the number of l i v i n g insects per needle decreased l i n e a r l y from the bottom to the top of the l i v i n g crown, and decreased l i n e a r l y from the outer part of the crown to the stem. 6. No differences were found i n abundance between North, West, South and East sides of single trees. 7. The mortality of Adelges cooleyi was lower on the higher populated trees (marginal and open) and on the higher-populated parts of the tree (low l e v e l ) .  70  APPENDICES  L i s t of Appendices; Appendix A  Table 3k. The average number of l i v i n g and dead insects per needle of generation  Appendix B  1.  Table 35. The average number o f l i v i n g and dead insects per needle of generation 2.  Appendix C  Table 36. Average numbers of l i v i n g insects per needle by period and tree at -the low l e v e l on the l i v i n g crown.  Appendix D  Table 37- Average numbers of dead insects per needle by period and tree at the low l e v e l on the l i v i n g  crown.  Appendix E  Table 38. Heights of sampling l e v e l s of trees.  Appendix F  Table 39. Changes i n development of Adelges c o o l e y i .  Appendix G  Table UO. Number o f l i v i n g insects per needle and average twig lengths change by height on the l i v i n g  crown.  Table h i . Number of l i v i n g insects per needle and average twig lengths change h o r i z o n t a l l y within the l i v i n g crown, at the middle o f low l e v e l of the trees. Appendix H  Table h2. The growth of foliage by l e v e l and time on trees No. Ik and  Appendix I  15.  Table k3. Calculation of the equation and c o r r e l a t i o n c o e f f i c i e n t of average number of l i v i n g insects per needle on average twig lengths.  Appendix J  Table kk. The average length of twigs by l e v e l and tree.  Appendix K  Tables k5, k6, k7, k8, k9 and 50. Climatological Station Report.  Appendix L  Crown shapes o f three t y p i c a l trees.  Appendix M  Table 51. No.  Average l i v i n g and dead insects per needle, on trees  1, 2, 10, 11,  lk and  20.  71  Appendix N  Table 52. Average numbers of l i v i n g and dead insects per needle by l e v e l and tree.  Appendix 0  Table 53. The average number of l i v i n g insects per needle, related to 8 factors: D.B.H., height, age, crown width, c l e a r stem, % of l i v i n g crown, bud opening time and l o c a t i o n of the trees.  Appendix P  Table 5U. Percentages of mortality by generation and tree.  72  Table 3k. The average number of l i v i n g and dead insects per needle of generation 1. (The numbers are averaged o f 5 sampling periods, from June k to July 2). Level Low Medium High Tree N E N ~s F~ E | N S E W No. No. o f l i v i n g insects. T572TT "OTIS" ~o7nr "OTEH o.io T "0750" 0.02 7J72TF "0702 "0730" 0.U2 0.k8 0 . 5 0 o.ok 0.16 0 . 3 8 o.ik 0.3k 2 . 0 0 0.02 0 . 0 2 0.02 0.2k 0.20 0.3k 3 . 0 0 0.00 0.12 o . i k 0.22 0.10 0.02 0.12 0.3k . 0 0 0.00 k 0 . 1 6 0 . 0 8 0 . 0 5 0 . 0 5 0.3k 0.58 0.26 0.22 0 . k 6 .02 0.00 0.28 0.22 0.32 5 0 . 0 0 0.06 . 0 0 0.00 0.12 0.12 0.12 0.02 0 . 0 0 6 0.1k 0.16 0 . 3 k 0.05 0.06 0 . 0 2 . 0 0 0.00 0 . 0 8 0 . 0 0 7 o.ok 0.00 0.02 0.12 8 0 . 3 0 0 . 2 0 0 . k 2 0.1k 0.22 0.18 0.2k 0.16 0 . 0 0 0.00 . 0 0 0.02 0.06 0 . 1 0 0.06 .02 0.02 9 0.02 0 . 0 0 o.ok 0.02 0 . 0 5 0 . 1 6 o.ko 0.12 .00 0.05 10 0.16 0.08 0.06 0 . 2 8 0.2k 0 . 0 0 0.00 0.36 0.2k o.kk 0.36 0.06 0.0k o.ok 0.12 0 . 0 0 0 . 1 0 11 . 0 5 0.00 0.26 0.22 12 0.60 0.06 0 . 0 8 0.12 0 . 0 0 0 . 0 0 . 0 5 0.00 0.3k 0.18 13 0.28 0.10 o.ik 0.02 0.22 o.ok 0 . 0 0 0.10 0 . 8 2 0.66 0 . k 6 0.78 1.0k lk 0.76 0.38 0 . 0 0 0 . 3 0 0.k2 0.86 0.12 0.12 15 0.26 o.ik 0.16 0 . 5 8 o.ko 0.06 0.32 0.30 0.06 0.08 0.0k 0.02 0.02 16 0 . 0 0 0.00 0.0k 05 0.00 0.06 0.0k 0 . 0 0 0.06 0 . 0 0 0 . 0 0 0.02 0 . 0 0 0 . 0 0 0 . 0 0 17 .00 0 . 0 5 18 0.0k 0.0k 0.02 0.00 0.00 0 . 0 0 0 . 0 5 0.10 0 . 0 0 0 . 0 0 .00 0.00 0.06 0.0k o.ok 0 . 0 0 0 . 0 5 0 . 0 5 0 . 0 5 0 . 0 0 0 . 1 0 0.0k .00 0.00 19 0.08 0.08 0 . 0 0 0.06 o.ok .00 0.00 20 0.02 0 . 0 0 0 . 0 0 0.02 0 . 0 0 0 . 0 8 0.06 0.0k 0.0k 0.02 o.ok 0 . 0 5 0 . 0 0 .00 21 0.00 0.00 0.28 0.22 o.ik 0.36 0 . k 6 o.ok 0 . 3 0 o.ik o.o5 .12 0.18 22 0.02 0.12 0.28 0.12 0.08 0.1k .08 0.12 23 0 . 3 0 0.16 0.12 0 . 0 8 0 . 0 2 0.02 0.0k 0.06 0.02 0 . 0 0 2k top 0.02 0.2k 25 o.ok 0.02 0.02 o.oo 0 . 0 0 0 . 0 0 0.02 0 . 0 0 0.00 broken . 0 0 0 . 0 0 No. of dead insects, 0 . 0 0 0 "0720""0715""07W "OTT? "0778""0706" 0 . 0 5 0.06 "OToTT 75b~"0702" 1 .08 0.02 0.18 0.26 0.20 0.08 0.12 0.16 0.12 0.06 0.02 2 0.18 .02 0.02 0.16 0.12 0.20 0.08 0.02 0.26 0.06 o.ok 3 o.ok 0.18 0.06 0.16 0.18 0.20 .05 0.05 0.00 0.00 k 0.22 0.28 0.02 0.22 0.26 o.ik 0.3k 0.00 .02 o.ok 5 0 . 1 0 0 . 1 0 0.02 0 . 0 0 0.05 0.10 0.10 0.12 o.ok .00 o.ik 6 0.12 0.06 0 . 0 5 0 . 0 0 0.00 0.3k 0.12 0.18 .00 o.ik 0.22 7 0.18 0.12 0 . 0 8 0.00 0 . 2 2 0.26 0.2k . 0 0 o.ik 0.16 0 . 0 5 0.10 8 0.08 0.00 0 . 0 0 0.00 0.02 . 0 0 o.ok o.ok o.ok 0 . 0 0 9 0.08 0 . 2 0 0.30 0.22 0.10 0.06 o.ko 0.12 0 . 0 5 0 . 0 0 . 0 0 0.00 10 0.20 0.30 0.22 0.22 0.06 0.10 0.12 0.06 0.05 .05 0.05 11 0.08 0 . 0 0 0 . 0 0 0.06 0.10 0.18 0.02 o.ik 0.16 12 .00 0 . 0 5 0.16 0 . 0 5 0 . 1 0 0.12 0.16 o.ok 0.32 0 . 2 0 0.06 13 0.6k 0.k2 0.80 0.66 0 . 1 0 0.56 0.66 0.68 0.58 0.7k 0.8k 0.72 lk 0.38 0.72 0.10 o.ok 0.06 0 . 2 0 0.18 0.12 0.18 15 0.02 0.1k 0.00 0.02 . 0 0 0.00 O.Ok 0.10 0 . 0 5 0.02 o.ok 16 0.06 o.ok 0.00 0.00 0.00 0.00 0 . 0 0 0 . 0 0 0 . 0 5 0.00 05 0.00 17 0.00 0.00 0.00 0.00 0.02 0 . 0 5 o.ok o.ok 0.02 . 0 0 0.00 18 0.05 0.20 0 . 0 5 0.06 o.ok 0.00 0.06 0 . 0 0 0.06 0 . 0 5 0 . 0 0 .00 19 0.00 0.02 o.ok 0.02 0.08 0.08 0.02 0.10 0 . 0 8 0 . 0 0 o.ok .02 20 0.02 0 . 0 5 0 . 0 0 0.00 O.Ok 0.02 0.00 0 . 0 0 o.ok O.Ok 0 . 0 0 .00 21 0.00 0 . 1 0 0 . 2 0 0.16 0.10 0.16 0.06 0 . 0 0 0.12 .16 0.12 22 0.26 0 . 1 0 0.08 0.10 0.10 0.16 0.08 0.1k .lk 0.10 23 0.02 0.02 0.12 0 . 0 0 0.02 0 . 0 8 broken 0.02 o.ok o.ok 2k 0.02 0 . 0 0 0.00 0.02 0.00 0.02 0 . 0 0 0 . 0 5 top 0 . 1 0 0 .00 0.00 25 0.05 0.05  Appendix A  -  073? "o7nr  750""  73 Appendix B Level Tree No.  1 2 3 u 5 6 7 8 9 10 11 12 13  lU  15 16 17 18 19 20 21 22 23 2U 25 1 2 3 u 5 6 7 8 9 10 11 12 13 lU 15 16 17 18 19 20 21 22 23 2U 25  N  Table 35. The average number of l i v i n g and dead insects per needle of generation 2. (The numbers are averaged of 8 sampling period, from J u l y 9 to September 21*). Low Medium High W E j N S ST E~ E N W  033 0.39 0.18 O.UU 0.36 0.16 0.05 0.25  0.26 0.29  0.16 0.10 0.05 0.20 0.06 0.06 0.05 0.23 0.15 0.15 0.11* 0.01 0.00 1.51 1.75 0.05 0.11 o.ou 0.03 0.00 o.ou 0.01 0.05 0.06 0.06 0.06 o.ou 0.03 0.25 0.15 0.09 o.ou 0.03 0.03 0.01 "TJ720"  0.09 0.09 0.09 0.05 0.08  0.03  0.08  0.05 0.09  0.05  0.00  0.15 0.03  0.01 0.01 0.03 0.03 0.01 0.08 0.00 0.01  0.05 0.09 o.ou 0.08 0.00 0.08 0.00 0.03 o.o5 0.01 0.00 0.20 0.05 0.03 0.01 0.00 0.01 0.00 0.05 0.08 o.ou 0.03 0.00  0715 0.31  1^  0.20  0.59  0.60 0.61 0.28  O.lU 0.53  o.iU 0.05 0.11 0.25 0.50 0.05 0.7U 1.76 0.2U 0.05 0.03 0.01 0.01 0.03 0.01 0.05 o.ou 0.06 o.ou 0.03 0.05 0.20 0.19  0.09 0.10 0.23  o.ou 0.01  0.05 0.00  No. of l i v i n g insects. 0723 0713 "oTiT 0715 0.18 0.10 o.io 0.11 0.10 0.16 0.19 0.11 0.13 0.21 0.2U 0.15 o . i 5 o.ui 0.09 0.10 0.2U 0.08 o.ou 0.01 0.05 0.06 0.05 0.13 0.16 0.01 o.ou 0.03 0.00 0.03 0.01 o.ou 0.03 0.06 0.03 0.00 o.ou o.ou o.ou 0.03 0.06 o.ou 0.01 0.00 1.78 1.5U 1.11 0.76 0.13 0.08 0.13 0.00 0.01 o.ou o.ou 0.00 0.00 0.00 0.10 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.10 0.03 0.01 0.03 0.03 0.00 0.00 0.00 0.00 0.3U 0.29 0.31 0.23 0.25 0.26 0.20 0.29 0.00 0.03 0.01 0.00 0.00 0.00 0.00 0.00  No. of dead insects "OTOT 0709 "07IF 0.10 ~ o n r UToT 0.06 0.09 0.10 0.06 0.09 0.09  0.11 0.13 0.16 0.08 0.05 0.10 o.ou 0.01 0.01 o.ou 0.03 0.05 o.ou 0.09  0.05  0.08  0.05 0.08 0.03 o.ou  0.05  o.ou 0.11 0.10 0.10 0.03 0.12 o.o5 0.01 0.00 0.03 0.03 0.00 0.00 0.01 0.01 0.08 0.06 0.03 0.00  o.ou  0.05 0.03 0.00 0.00 0.00 0.03 0.01 0.01 O.OU 0.05 o.ou 0.01 0.03 0.01 0.10 0.16 O.IU 0.13 0.11 0.23 0.11 0.00 0.08 0.00 0.03 0.00 0.00 0.01 0.00 0.01 0.00 0.00 0.10 0.00 0.00 0.00 0.00 0.01 0.00 0.10 0.00 0.00 0.03 0.03 o.ou 0.00 0.00 0.00 o.ou 0.10 o.oU 0.01 0.01 0.01 0.03 0.00 0.00 0.00 0.00 0.00 0.08 o.ou 0.08 0.08 0.05 0.05 0.08 0.10 0.10 0.11 0.03 0.01 0.01 0.00 0.01 0.00 0.00 0.01 0.00 0.00 0.00 0.00  O T O T  0.06 0.00 0.00 0.00 0.00 0.00 0.00 0.03 0.00 0.00 0.00 0.00 o.U5 0.05 0.00 0.00 0.20 0.00 0.01 0.20 0.2U 0.10 0.00  .01 .01 .01 00 00 .00 .00 .00 .01 00 .10 .00 .00 .U3  o.ou 0700 0.01 0.00 o.ou 0.00 0.00 0.10 0.00 0.20 0.20 0.00 0.00 0.00 0.10 0.00 o.ou 0.03 0.00 0.00 0.00 0.00 0.00 0.00 0.10 0.16  o.Ui  .09  .00 0.00 .00 0.00 .00 0.00 .20 0.00 .01 0.01 .00 0.00 .13 0.13 .10 0.10  0.08 0.00 0.00 0.00 0.00 0.00 0.00 0.13 O.IU  top broken  0 .00 0.00 0.00  0.01 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.10 0.08 o.ou 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.01 0.03 0.00 0.00 0.05 0.05 0.05 0.08 top broken 0.00 0. 00 0.00 0.00  "oToT 0.01 0.00 0.00 0.00 0.00 0.00 0.00 0.01 0.00 0.10 0.00 0.00 0.13 o.o5 0.00 0.00 0.00 0.00 0.00 0.00 0.05 0.05  0. w 0. 01 0. 00 0. 00 0. 00 0. 00 0. 00 0. 00 0. 00 0. 00 0. 00 0. 00 0. 00 0. 11 0. 09 0. 00 0. 10 0. 00 0. 00 0. 00 0. 00 0. 06 0. 08  0.00 0.03 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00  Appendix C Table 36. Average numbers of l i v i n g insects per needle by period and tree at the low l e v e l on the l i v i n g crown. Tree No.  Generation 2 Sampling periods (week"T  Generation 1 Sampling periods (week)  0 0.2« 1 2 0.53 3 0.50 0.50 u o.uo 5 0.20 6 0.07 7 o.U5 8 o.i3 9 0.38 10 0.U5 11 0.70 12 0.U3 13 0.95 iu 0.03 0.10 15 0.08 16 0.03 17 18 0.05 0.10 19 20 o.o5 21 0.27 22 o.i5 23 0.03 2U 0.03 Ave. 25 "0T2T  1 "oTTF 0.55 0.27 0.60 o.uo 0.23 0.03 0.25 0.03 0.30 0.U3 0.57 0.23 0.73 0.20 0.00 0.03 0.00 0.05 0.05 0.05 0.33 0.15 0.03 0.05 0.23  F  2 0718 O.UO 0.30 0.55 0.33 0.23 0.27 0.13 0.13 0.13 0.53 0.33 0.37 0.83 0.30 0.10 0.03 0.05 0.03 0.05 0.08 0.27 0.05 0.08 0.00 0.23  1  u~  ~G~TT "0718" ~G~W  0.28 0.07 0.30 0.20 0.20 0.03 0.28 0.10 0.15 0.25 0.13 0.23 0.60 0.20 0.03 0.08 0.03 0.05 0.05 0.03 0.07 0.05 0.03 0.03  "amr  0.15 0.17 o.35 0.23 0.20 0.03 0.23 0.00 0.10 0.23 0.07 0.07 0.33 0.10 0.07 0.00 0.03 0.05 0.03 0.00 0.13 0.15 0.03 0.00 0.12  0.38 0.10 0.75 0.27 0.23 0.00 0.35 0.00 0.00 0.30 0.U7 0.03 1.83 0.00 0.00 0.00 0.00 0.00 0.00 0.03 0.23 0.00 0.00 0.00 0.21  0.60 0.U7 0.75 0.50 0.27 0.10 0.38 0.07 0.00 0.35 0.53 0.00 1.63 0.00 0.03 0.03 0.03 0.05 0.03 0.05 0.33 0.00 0.00 0.00 0.26  T57UT 0.35 0.50 0.65 0.73 0.17 0.13 0.28 0.10 0.08 0.23 0.50 0.03 1.93 0.30 0.10 0.00 0.03 0.08 0.13 0.10 o.uo 0.20 0.03 0.00 0.30  0725 W  0.28 0.U0 0.25 0.U3 0.17 0.10 0.18 0.13 0.08 0.20 0.10 0.03 1.90 0.33 0.00 0.05 0.05 0.13 0.13 0.03 0.23 0.20 0.08 0.03 0.23  0.20 0.17 0.60 0.23 0.15 0.10 0.15 0.00 0.13 0.20 0.00 0. 07 1. U3 0.07 0.13 0.03 0.00 0.00 0.05 0.03 0.20 0.30 0.10 0.03 0.1B  9  TOO" TJ7I6 " O J -  0.13 0.23 0.30 0.30 0.13 0.10 0.U3 0.23 0.23 0.20 0.13 0.00 1.05 0.13 0.03 0.00 0.03 0.08 0.08 0.03 0.17 0.10 0.08 0.00 0.17  0.18 0.37 0.U5 0.27 0.07 0.03 0.18 0.00 0.08 0.13 0.27 0.00 0.98 0.17 0.00 0.00 0.03 0.03 0.03 0.00 0.17 0.10 0.00 0.00 0.15  0.23 0.27 o.uo 0.27 0.27 0.10 0.20 0.00 0.08 0.15 0.10 0.00 1.00 0.07 0.00 0.03 0.03 0.05 0.03 0.03 0.10 0.05 0.03 0.05 "07TF -4 •Cr-  Appendix D Table 37.  Average numbers of dead insects per needle by period and tree at the l i v i n g crown. Tree Generation 2 Generation 1 No. Sampling periods (week) Sampling periods (week) 1 0 0 2 i u— 1 "oToT 0.10 0.15 0.15 008" 0.0b ~OJ5F "07lB~ 0.05 0.10 "7J30 0.15 2 0.18 0.18 0.15 0.15 0.25 0.05 0.05 0.03 0.10 0.00 0.08 0.13 3 0.10 0.20 0.20 0.23 0.03 0.07 0.13 0.10 0.07 0.10 0.17 0.20 0.00 0.00 0.10 0.10 0.10 0.10 0.25 U 0.23 0.15 0.15 0.20 0.25 0.33 0.00 0.03 0.07 0.10 0.03 0.13 0.13 5 0.23 0.23 0.33 0.23 6 0.13 0.10 0.17 0.13 0.07 0.07 0.07 0.10 0.07 0.07 0.03 0.17 0.27 0.20 0.10 0.30 0.00 0.00 0.03 0.03 0.00 0.03 0.03 7 0.23 8 0.15 0.18 0.25 0.13 0.03 0.05 0.03 0.10 0.08 0.18 0.08  low l e v e l on the  -  -  9 10 11 12 13 Hi 15 16 17 18 19 20 21 22 23 2ii 25  Ave.  0.23 0.07 0.33 0.38 0.23 0.13 0.63 0.03 0.10 0.00 0.03 0.03 0.05 0.03 0.07 0.05 0.03 0.03 0.15  0.03 0.08 0.20 0.17 0.03 0.65 0.07 0.00 0.00 0.05 0.08 0.00 0.00 0.10 0.15 0.05 0.00 0.12  0.00 0.20 0.15 0.13 0.33 0 80 0 10 0 03 0 ,00 0 05 0 08 0 10 0 00 0 20 0 15 0 00 0.00 0715" oTiF ~o7ic"  0.00 0.23 0.18 0.23 0.23 0.20 0.07 0.23 0.30 0.70 0.78 0.23 0.10 0.03 0.10 0.08 05 0.03 00 0.03 08 10 10 05 05 03 13 10 05 0,08 03 0.00 0.03 0.03  0.00 0.00 0.00 0.00 0.00 0.10 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00  O.OHi  0.03 0.00 0.03 0.07 0.00 0.03 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.019  0.00 0.03 0.05 0.03 0.00 0.10 0.03 0.03 0.03 0.00 0.00 0.05 0.08 0.03 0.05 0.03 0.03 0.050  11  oHF  0.15 0.10 0.20 0.17 0.03 0.07 0.05 0.00 0.10 0.10 0.07 0.03 0.23 0.00 0.00 0.00 0.00 0.00 0.03 0.05 0.20 0.10 0.00 0.00  0.00 0.05 0.08 0.08 0.10 0.10 0.03 0.00 0.18 0.30 0.10 0.03 0.10 0.00 0.00 0.00 0.00 0.00 0.05 0.05 0.00 0.03 0.00 0.03 0.03 0.03 0.15 0.00 0.03 0.03 0.00 0.00 o.ouu 0.0U8 0.067 0.07U TJ^7l  0.00 0.03 0.05 0.03 0.00 0.08 0.00 0.00 0.00 0.03 0.05 0.00 0.03 0.07 0.05 0.03 0.00  0.00 0.05 0.08 0.00 0.00 0.20 0.03 0.00 0.03 0.03 0.03 0.05 0.00 0.13 0.10 0.03 0.00  0.03 0.05  -~0  Appendix E Table 38. Heights of sampling l e v e l s of trees. Tree No.  1 2  3  h  5 6 7  8  9  10 11 12  13 Hi  15 16 17 18 19 20 21 22 23 2U 25  Height of l e v e l s i n feet Low Medium High  21  16  15 19 17 21 16 20 10  19 17 16  8  5 6  25  h3  36 39  21  Ul  0.6  2  19 U5  27  26  27  36 36 39 33 36 16 37 32 31 12  11 10  39 53 42  33 35 39 53  55  57  50 52 22  55  47 46 16 17 lit 5u 63  47  53 1.3  65  h  27 57  LE  LE, :LS, ME LE, ME LE LE LN, MN LW  LW, MW, HW LW, MW, LN  HW  U8  h3  30  Levels and directions where could not be found l i v i n g branches  39  2  6 —  LN LW, LE  61  Key: L = Low l e v e l , M = Medium l e v e l , H = High l e v e l W = West, N = North, E = East, S = South  77  Appendix F Table 39. Changes i n development of Adelges cooleyi. Tree No. i 2 3  1* 5 6  7 8  9  10 11 12 13 LU  15 16  17 18  19  20 21 22 23 2U  25  June  U  FSN2 PSN1+2 PSN1  PSN3  PSN2 PSN1+2 PSN1 PSN1+2 PSN1+2 PSN1+2 PSN3" PSN2+3 PSN2+3 PSN2+3 PSN1+2 PSN2 PSN1+2 PSN1 PSN2+3 PSN2+3 PSN2+3 PSN2+3 PSN1+2 PSN1 PSN1+2  June 11 PSN3+A+E PSN3+A PSN2+3 PSN3+A+E PSN3+A+E PSN3+A PSN2+3 PSN3+A+E PSN3+A PSN3+A PSN3+A+E PSN3+A+E  PSN3+A+E PSN3+A+E PSN3+A PSN3+A PSN2+3+A PSN2+3 PSN3+A+E PSN3+A+E PSN3+A+E PSN3+A+E PSN3+2 PSN2+3 PSN3+A  June 18 A+E A+E PSN3+A+E A+E A+E A+E PSN3+A+E A+E A+E A+E A+E A+E A+E A+E A+E PSN3+A+E A+E PSN3+A+E A+E A+E A+E A+E PSN3+A+E PSN3+A+E A+E  Date June  25  A+E+H A+E A+E A+E A+E A+E A+E A+E A+E A+E A+E A+E A+E A+E A+E A+E A+E A+E A+E+H A+E A+E+H A+E A+E A+E A+E  July 2 A+H A+E+H A+E A+E+H A+E+H A+E+H A+E+H A+E+H A+E+H A+E+H A+E+H A+E+H A+E+H A+E+H A+E+H A+E A+E+H A+E A+E+H A+E+H E+H A+E+H A+E A+E E+H  July  9  H E+H E+A+H E+H E+H E+A+H A+E+H E+H E+H E+H E+H E+H A+E+H E+H E+H A+E+H H A+E+H A+H+E E+H E+H E+H A+E+H E+H H  July 16 H H E+H H H H E+H H H H H E+H H H E+H E+H H E+H H H H H E+H H H  Fall • Winter H H H H H H H H H H H H H H H H H H H H H H H H H  Key: N = Larva, A = Adult, PS = Progrediens and Sexupara, E = Egg, 1,2,3 = Instars (1 means f i r s t and second instars), H = Neosistens  Appendix G Table kO. Number of l i v i n g insects per needle and average twig lengths change by height on the l i v i n g crown. (Averaged data of tree No. k, 10 and 2 5 ) . Date 0-5  5-10  June 5 J u l y 25 Sept. 26 Sept. 26  10-15 0.22 0.32 0.18  -  20.0  Height l e v e l s i n f e e t 15-20 20-25 25-30 30-35 35-kO U0-U5 k 5 - 5 0 50-55 55-60 Number of insects per needle 0.37 0.27 0.19 0 . 1 0 0.11 0.03 0.02 0.02 0 . 0 0 0.27 O.lo 0.19 0 . 0 3 0.01 0.0k 0.03 0.00 0.00 0.20 0.10 0.11 0.06 0.00 0.02 0.02 0.01 0.00 Average twig length i n cm. 20.3 19 .U 19.5 20.6 19.9 20.9 2 1 . 1 22.2 23.0  Table k l . Number of l i v i n g insects per needle and average twig lengths change h o r i z o n t a l l y within the l i v i n g crown, at the middle of low l e v e l of the trees. (Averaged date, of tree No. k, 10 and 25). Date  Distances from the stem i n f e e t 3-6 6-9 9-12 12-15 15-18 Number of insects per needle 0.02 0.09 0.16 0.2U 0.10  0-3  June 5 July 25 Sept. 26 -  Sept. 26  0.01 0.03  lO.k  0.03 0.03  0.1k 0.1k  0.10 0.12  0.15 0.25  Average twig length i n cm.  12.9  17.5  20.8  22.9  Appendix H Table U2. The growth of foliage by l e v e l and time on trees No. LU and 15Tree No.  Level June U  June  11  June 18  June  25  July  2  Date July  9  July  16  July  23  July  30  Aug.  6  Aug. Sept.  20  3  8.5  8.6  Sept.  2U  Twig lengths i n cm.  LU  15  Low  3.7  5.6  7.0  7.U  7.9  8.0  8.1  8.3  8.3  Med.  U.6  5-5  6.8  7.6  8.5 9.6  9-7  9.8  9-8  High  5.8  8.1  9.U  12.5  13.6 15.U  16.8  16.9  17.2  17.3  17 .h  17-5  17.5  Low  1.9  2.6  3.8  U.U  U.8  5.6  6.5  6.9  7.0  7-0  7.2  7.2  7.2  Med.  2.3  2.8  3.7  U.8  5.7  6.6  7.9  9.U  9.6  9.6  9.8  9.9  9.9  High  3.1  U.U  5.8  6.6  8.0 9.2  10.7  11.2  11.3  8.U  8.6  10.0 1 0 . l 1 0 . l 10.1  ll.U  11.6  11.7  11.8  -o  80  Appendix I Table k3. Calculation of the equation and c o r r e l a t i o n c o e f f i c i e n t of average number of l i v i n g insects per needle on average twig lengths. Tree No.  1 2 3 k 5 6 7 8 9 10 11 12 13  m  15 16 17 18 19 20 21 22 23 2k 25 Total Ave.  X  Y  b.k 6.0 6.k 15.9 8.7 7.7 6.3 9.8 9.3 23.6 13.6 7.3 6.8 7.2 6.1 k.l  <U2 0.65 0.61 0.98 0.69 0.39 0.15 0.55 0.1k 0.29 0.59 0.62 0.29 2.13 0.30 0.10 0.05 0.05 0.10 0.12 0.07 0.k2 0.23 0.08 0.03 10.05 0.k02  a.2  3.8 6.0 5.8 5.1 2.9 8.7 3.1 5.1* 192.8 7.71  52  70.56 36.00 k0.96 252.81 75.69 59.29 39.69 96.0k 86 .U9 556.96 l8k.96 53.29 k6.2k 51.8k 37.21 22.09 17.6k lk.kk 36.00 33.6k 26.01 8.kl 75.69 9.61 29.16 1960.72  52 0.176k 0.k225 0.3721 0.960k 0.k76l 0.1521 0.0225 0.3025 0.0196 0.08kl 0.3k8l 0.38kk 0.08kl k.5369 0.0900 0.1000 0.0025 0.0025 0.0100 o.oikk 0.00k9 0.176k 0.0529 0.006k 0.0009 8.7127  XY  3.$2B 3.900 3.90k 15.582 6.003 3.003 0.9k5 5.390 1.302 6.8kk 8.02k k.526 1.972 15.336 1.830 0.U70 0.210 0.190 0.600 0.696 0.357 1.218 2.001 0.2k8 0.162 88.2kl  X = average twig length at the low l e v e l of the trees, Y = number of l i v i n g insects per needle, b -  0.023, a - 0.22k7> r - 0.228, t - 0.112, Sb - 0.02013,  -0.019 - i - 0.065 The c a l c u l a t i o n without trees No. 10 and l k :  b - 0.058, a - -0.08, r - 0.680, t - k . 2 5 , Sb - 0.01375, 0.030 * i * 0.086 Equation: Y -  0.058X-0.08  81  Appendix J Table UU. The average length of twigs by l e v e l and tree. Tree No. Level 1 2  Low  3.0  0.6  June U Medium  3.9  3.5  0.7 0.U 5.2  6 7 8 9 10  U.5  5.6  12 13  3.4  3 U 5  11 Hi 15  16 17 18 19 20 21 22  23 2U 25  o.U  2.1  0.0 3.5 2.5  3.7 6.1 3.2  5.0 2.1 1.8 0.0 1.0 2.1  1.2 2.0  1.7 1.0 0.0 2.1  3.3 0.0 3.5 3.1  4.0  6.7 3.7  U.O  5.6  2.4  2.3 0.0 1.1  3.0  1.9 2.7 2.5 1.3 0.0 2.6  High  6.8 1.0 0.7 7.1 U.5 6.2 0.0  3.6  3.1 5.1  7.5  U.9 3.8 6.3 2.8 3.U 0.0  l.U 3.7 2.8 3.6  September 26 Medium High 8.6 8.U 18.6  Low  6.0  6.U  15.9  8.7 7.7 6.3 9.8 9.3  23.6  13.6 7.3 6.8 7.2 6.1  U.7  U.2  3.8  9.6 9.3 12.U 10.6 8.9  2U.7 17.2 9.1 6.5 8.0  8.3  7.0  6.7 5.3  6.2  5.1  5.2  5.8  6.8  11.3 10.8 20.1 11.7  21.9 18.7 11.1 8.5  26.6 19.9 10.6 6.1 13 .U 10.7 13.3 12.7 9.8 9.U 10.2  8.7  U.3 12.7  6.9 6.8 17.7  5.U  6.6  io.U  2.9  0.0  3.1  3.2  15.0  6.0  3.0  1.7  6.1  7.6  3.5  -  82  Appendix K Table U5. Climate-logical Station Report (April I960, Vancouver U.B.C.). Date  Air temperature F Max.  1 2 3  U  5 6 7 8 9 10 11 12 13  11*  15 16 17 18 19 20 21 22 23 21* 25 26  27 28 29 30 Jl  Sum Mean  1*6.5 1*9.7 53.7 59.7 59.8 50.1* 55.6 61*.1* 57.8 51.7 51.8 53.6 52.3 1*8.0 U8.3 50.3 1*9.5 53.0 1*6.6 1*9.2 1*9.1* U7.U 52.8 58.0 57.3 57.5 60.1* 63.0 62.2  59.0  Min.  3H.9  1*3.8 1*8.8 1*5.9  1*1*.9  U2.0 1*6.8 50.1* 1*0.2 36.1* l*l*.l 1*2.0 1*0.3 1*1.2 37.5 37.3 1*2.1* 1*1.3 1*0.9 U2.5 36.7 38.5 1*2.8 1*3.2 1*2.0 1*5.8 1*3.3 1*7.9 1*8.6 U6.5  RH AM  93 96 90 85 93 86 68 79 79  71*  98 90 95 77 85 82  90 88  81*  79 87 85 71 88 86 89 77 73 83 88  PM  7U 96 70 80 85 75 59 87 56 65 66 70 77 70 78 60 1*8 93 86 77 73 53 56 61* 61 55 1*9 52 78 71*  2087 1618.9 1282.9 2538 53.96 1*2.76 81*. 60 69.57  Rain EvapoWind Sunshine inches ration mileage total past 21+ hours hours  O.ll* 0.19 0.10  0.100 0.021 0.022 0.106 0.090 0.090 0.128  o.iu  0.07 o.ol* 0.13 0.11* 0.17 0.07 0.08 0.1*1 0.28  0.06 0.01 0.01  0.177  0.01*3  0.110 0.115 0.083 0.071 0.06U 0.106 0.137  0.081*  0.09U 0.050  0.028  0.090 0.115 0.078 0.036 0.120 0.0U6 0.187  0.231*  2.01*  0.2 8.1  U.5  6.3  9.U  9.1* 0.9 9.8 8.9 6.8 2.8 U.U  9.2  U.U  9.5  1.5 6.0 10.2  7.U  0.5 0.9 5.6 10.1 13.1  13.3  0.177 0.135  10.7  2.81*7  178.8 5.96  0.068 0.095  U.9  83  Appendix K Table  1*6. Climatological Station Report (May i960, Vancouver U.B.C.). Date  1 2 3 I* 5 6 7 8 9 10 11 12 13 Hi 15 16 17 18 19 20 21 22 23 2li 25 26 27 28 29 30 31 Sum Mean  Air  RH  temperature F  Max.  Min.  AM  Rain inches PM  57.8 47.4 73 8i 47.3 89 57.7 94 94 86 53.5 U5.3 94 51.0 1*5.8 87 54.1 1*8.0 60 89 98 96 59-3 51.4 1*6.6 86 59.3 63 1*2.2 78 56.1* 65 62 61* 5U.9 47.4 53.2 93 94 71.7 50.1 92 57.9 85 50.6 91* 55 55.1 60.2 45.0 79 57 U8.2 72 57 59.3 62 57.6 1*3.8 79 1*5.8 98 74 56.9 54.1 95 76 42.3 1*3.2 90 57 53.3 96 7U 58.3 46.5 62 1*9.0 li5.8 74 1*1.2 53.3 89 65 1*2.0 58 87 53 .4 60 71 1*0.3 55.5 58.8 68 66 U7.5 60.1 60 67 49.5 87 75 61.7 48.5 57.6 90 72 U6.3 62 49.0 78 55.9 1*8.6 82 79 60 .U 50.8 82 94 56.3 60.6 1*9.0 81 69 2207 1770.7 11*1*8.6 2609 57.12 71.2 1*6.73 81*.2  0.090 0.032 0.370 0.380 0.080 0.1*20 0.1*1*3  0.215 0.160 0.070  Wind Sunshine Evapor a t i o n mileage t o t a l past 21* hours hours  0.090 0.086 0.055 0.017 0.01*8 0.108 0.032 0.163 0.188 0.130 0.021* 0.023 0.177 0.181* 0.158 0.090 0.080 0.057 0.152  0.750 0.205 0.133 0.017 0.11*1 0.178 0.220 0.180 0.270 0.068 0.01*5 0.029 0.081* 0.165 0.050 3.762 0.1213  O.lli*  0.011 0.051 3.071 0.0991  120 100 97 50 120 100 89 68 67 66 69 59 80 82.5 69.5 59.5 86 77 76.5 110 92 105 74 112.5 106 123.5 75.5 97 70.5 108 11*2 2752.0 88.8  0.8 0.1  6.8 0.3 9.4 6.2 6.1 0.9 7.4 10.0 10.1 5.6 2.1* 6.6 6.5 10.6 9.1 9.7 12.0 10.0 3.7 1.2 1.5 3.4 0.9 7.1 11*8.1* 4.79  81*  Appendix K Table  1*7. Climatological Station Report (June I960, Vancouver U.B.C.). Date  1 2 3  1*  5 6 7 8 9 10 11 12 13  U* 15 16 17 18  19 20 21 22 23  21* 25  26  27  28 29 30 J>X  Sum Mean  A i r temperature F Max. Min.  62.2 65.8 68.5 61.3 63.3  66.7  59.8 58.5 63.0 66.6 65.7 65.0 69-7 63.9 59.7 58.1 62.6 63.3  62.5  59.3 60.7 59.8 66.6 72.6 67.0 6 3 . 1* 61.1 67.3 69.0 71.9  1*9-8 52.2 5U.0 2*5.2 53.8 50.8 51.0 U5.5 1*9.8 ' 53.5 52.5 51.6 55.5 52.1*  50.1* 51.1*  1*9.9  1*8.8 1*7.9 1*7.2 1*9.1 U8.I4 53.6 57.1  51*.3  53.8 55-9 52.5  51*.2 55.5  1 9 2 5 . 8 151*7.6 51.59 6U.19  RH  Rain inches  AM  PM  81*  72  86 91 67 68  61* 71* 82  68 67 89 83 70 81  91* 69 75 70  77 92 85 79  72 82 81* 83 82  71* 75 81  231*8 78.3  71*  70 72  62 58 70 65 57 77 72  72  70 96 96 61 57 57 75 68 77 71 75 78 79 83 70 56 59 79  2128 70.0  0.170 0.170  0.01*1  0.017  Wind Sunshine Evapor a t i o n mileage t o t a l past 2l* hours hours 101 0.182 3.7 0.116 2.1* 102.5 0.121 79 11.5 0.208 128 5.6 o.U*5 67.5 7.1 0.218 111 l l * .1* 11*0 0.269 11* .1* 0.186 78 13.1* 0.238 60 11*.2 3.6 0.267 75 0.133 83 10.3 9.8 O.I89 61*.5 0.230 83.5 1.1* 96 0.123 0.3 0.1 0.055 55-5 135 11-9 10.1 0.271 81*.5 101* 0.186 3.3 0.118 9.8 73 0.209 1.0 96.5 88 3.0 0.103 0.110 56 li*.3 O.198 68 12.6 88.5 0.21*7 U.l 0.109 95.5 0.093 1*3 2.2 0.075 1*6.5 0.156 1*1* 11*.3 0.222 12.6 72.5 101* 0.261 0.6  212.0 0.398 5.038 2 5 2 3 . 5 81*. 12 7.0 0.0132 0.1679  Appendix K Table L8. Climatological Station Report ( J u l y I960, Vancouver U.B.C.). Date  1 2 3  4  5 6 7 8 9 10 11 12 13 lit 15 16 17 18  19 20 21 22 23 24  25 26 27 28 29 30 31  Sura Mean  Max.  63.3 61.7 61.6 68. U 68.6 75.9 81.5 72.1 65.8 67.9 66.2 72.8 82.2  71.5  70.8  77.1 81.4 81.4  73.8 68.6 69.8 72.0 68.2 67.4 67.5 71.8 77.0 77.2 78.0  Min.  52.8  51.7 52.3 54.6 60.3 61.5  57.4 49.9  55.0 50.3 54.1 56.8 58.9 57.6 56.4  60.3 60.3 59.5 59. h 57.4  55.3 52.0 5l.o 52.0 56.8 59. u 63.8 60.6 62.8  65.6 58.3 76.5 221*5.4 176U.1 72.43 56.91 87.4  Rain inches  RH  A i r temperature F AM  85  78 80 87 87 65 78 76 95 78 71 70 79  65 66 63 69 84 81 83 81 78 80  77 87 71 66 84  PM  96 68 68 73 56 44  6U 60 65 69 58 59 65 56 59 49 73 70 70 68 64 67 58 64 75 55 50 47  68 47 50 64 83 71 1938 2379 62.5 76.7  Wind Sunshine Evapor a t i o n mileage t o t a l past 24 hours hours  0.122 0.030 0.077 0.178 0.2U6 0.254 0.310 0.282 0.302 0.243 0.285 0.166 0.200 0.266 0.263 0.265 0.182 0.340 0.291 0.289 0.290 0.2ii5 0.233 0.234 0.253 0.191 0.313 0.259 0.2Hi 0.302 0.285 0.342 0.030 7.722 0.2491  105.5 109.5 65 74.5 117 99 58 158.5 141  78 86 32 84 85 75 70 85 60 140 130 85 95 78 82 75.5 105 90.5 49.5  61.5 67 140 2782.0 89.74  1.1 13.8  14.6 14.1 14.4  LU.5 lit.2  14.5 14.4  8.7 7.0 11.3 9.5 12.0 10.2 12.6 14.3 14.1 14.1  Hi.O 13.9 11.9 13.9 lu.l  1U.1 12.U 8.U 13.8 13.4 8.1 10.8 378.2 12.2  86  Appendix K Table I49. Climate-logical Station Report (August Date  A i r temperature F Max. Min.  1 2  59.2 60.6  3  59-9 58.9 61.0 61.1 63.3  a  5 6 7 8 9 10 11 12 13  ia  15 16 17 18 19 20 21 22 23  2a 25 26  27 28 29 30 31  Sum Mean  7a.0 7a.5  61.a  61.3 59.8 57.6 53.2 56.0 55.0 59.3 61.9 58.8 61.7 50.2 52.0 53.8 53.5 5k.8  5a.0 55.3 53.5  5a.2  5a.0  56.9 58.7 58.2 55.0 57.2  .a  58 60.6 58.8 60.2 58.0 57.1 56.7  5a.5  62.a  52.2  5a.7  58.0 60.0 57.1 59.9  a9.5  RH PM  87 89 90 79 79 85 85  72 78 68 63 69 71 39  a3  21  83  67  77  63 73 51 89 73 93  sa 83  9a  78 98  27  93  82  89  69  90  6a  89 96  56  9a  51.0 51.9 53.0 53.3 53.2 53.2 53.0  9a  87 96  60  5o.a  77  70 98 56  89  96 91  9a  93 52.9 80 50.9 2 6 2 8 1807.9 1716.9 8a.8 58.32 55.38  5a.1  Rain inches  AM  ao  I960,  65 91 72 91 71 98  215a  69.5  Vancouver U.B.C.).  Wind Sunshine Evapor a t i o n mileage t o t a l past 2a hours hours 0.183  o.ia6  98.5  85 0.118 56.5 96 0.175 0.116 61 0.188 91 0.281 109 0.305 108.5 0.3a2 61.0 97 0.238 H»3 0.22a 73 0.229 88.5 0.005 95 73 o.oa3 0.031 0.163 0.122 96.5 0.230 0.013 120 0.077 75.5 0.172 72.5 0.165 68.5 0.220 0.110 99 0.310 0 . 1 0 8 110.5 0.108 88.5 0.360 0.006 62 0.060 0 . 1 2 6 101 0.001 83 0.a35 0.087 83 0.015 0.176 105.5 0.380 0.075 129 103.5 0.155 o.oa9 103 o.6ao 0.018 2837.0 3.013 583 0.0972 0.1LJ78 91.52  0.3a9  o.2a5  a.  0.8  2.5  a.2  a.a  13.6 13.3 13.7 13.6  13.5 13.0 12.9 8.5 8.5 0.2 8.1  6.9 9.2 8.5  i.a o.a 5.0  a.3  2.2 6.0  11.0 182.7  5.9  87  Appendix K Table $0. Climate-logical Station Report (September I960, Vancouver U.B.C.). Date  1 2 3  U  5 6 7 8 9 10 11 12 13  H*  15 16 17 18 19 20 21 22 23 21* 25 26 27 28 29 30  A i r temperature F Max. 50.3 53.8 55.8 56.0 51*.2 52.8 53.1 55.3 58.9 59.1 61.1 62.1 56.1 55.7  51*.5  56.7 51.8 5b.7 55.5 1*9.5 1*8.3 51.5 52.5 5i.o 53-7 52.1 5U.0 52.2 5U.l 53.8  Min. 1*8.0 50.5 51.0 52.3 51.0 1*9.7 1*9.3 53.5 52.0 52.7 51*.7 57-5 55.0 55.1 5U.3 56.1 51.8 53.5 55.2 1*7.2 1*7.2 50.3 51.9 50.2 52.8 50.9 51.9 50.5 52.3 53.0  RH AM 81* 80 73 79 80 82 76 86 63 66 67 75 93 96 98 96 100 93 98 81* 92 92 96 9b 91* 92 87 89 89 9b  Wind Sunshine Evaporation mileage t o t a l past 2b hours hours 0.11*0 52 11.1 0.11*8 11.6 7b.5 0.127 68.5 9.3 0.100 0.11*6 112.5 1.1 96 0.135 0.075 10.3 0.126 9.6 62.5 11.b 0.153 93.5 0.176 ll.l 93.5 0.176 11.1 83.5 0.153 10.9 53.5 0.161 b6 10.7 0.11*6 2.0 52.5 68 0.01*9 9-1 0.093 b2.5 1.2 0.070 37 7.8 0.087 59 7.1 0.123 56 0.010 0.036 99.5 0.1*05 0.017 116 6.6 0.120 0.155 8.8 16b. 5 0.006 0.129 52 9.b 0.023 0.01*6 57.5 0.31*0 93 0.9 0.337 0.01*8 85 0.030 0.056 73.5 7.b 0.101 122.5 9.7 0.10b 98.5 9.5 0.09b 115 9.5 0.117 123 9.6 0.131 61.0 8.6  Rain inches PM 71 77 77 91 62 68 83 68 1*3 b9 39 86 80 93 79 81 82 91 61* 81* 85 91 81* 95 80 83 72 78 67 73  -31  Sum Mean  1630.2 1560.9 2588 51*.31* 52.03 86.3  2276 75.9  1.506 3.183 2412.0 0.0502 0.1061 80.b  215.6 7.19  Appendix  L  C r o w n - s h a p e s o f three differently located trees  Open growth tree (No- 10)  Interior growth tree (No 25)  88  89  Appendix M Table 51.  Average l i v i n g and dead insects per needle, on trees No. 1,  2,  10,  11, Hi and 20. Living Generation 1 Tree Low l e v e l Medium l e v e l N E No. W N E W IM 072U 07T6 0.11* OTllT 0.10 0.22 0.02 "072U 2L 0.1*2 0.1*8 0.50 0.01* 0.16 0.38 0.11* 0.31* lhE 0.81* 0.66 0.1*6 0.78 1.01* 0.86 0.58 0.76 20M 0.08 0.08 0.00 0.06 0.01* 0.02 0.01* 0.00 10L 0.16 0.1*0 0.12 0.16 0.08 0.06 0.28 0.21* 11E 0.36 0.31* 0.1*1* 0.36 0.06 0.01* o.ob 0.12 Generation 2 IM 0.33 0.23 0715 "072T 072T 0713 "6715""5715" 2L 0.39 0.26 0.31 0.20 0.18 0.10 0.10 0.11 IUE 1.51 1.75 0.7U 1.76 1.5U 1.11 0.76 1.77 20M 0.01 0.03 0.03 0.06 0.06 0.06 O.OU 0.03 10L 0.01 o.ou 0.03 0.06 0.05 0.10 0.11 0.03 11E 0.23 0.03 0.01 o.ou 0.25 0.23 0.18 0.06 Dead" Generation 1 "075B" 0 . 0 6 "0732 "OTHT IM 0.20 0.10 o.oa 0.12 0.16 0.12 0.18 2L 0.26 0.08 0.18 0.20 IUE 0.7U 0.61* 0.58 0.72 0 . 6 U 0.U2 0.80 0.66 20M 0.08 0.08 0.02 0.08 0.10 0.02 0.02 O.OU 10L 0.08 0.20 0.30 0.22 0.10 0.06 O.UO 0.12 11E 0.20 0.30 0.22 0.22 0.06 0.10 0.12 0.06 Generation 2 0.20 077J6~7J709" "0709 "IF TJ7HT 0.11 7J70T 2L 0.09 0.05 0.06 0.09 0.06 0.09 0.09 IUE 0.15 0.20 0.10 0.16 0.10 0.13 0.11 0.23 20M 0.03 0.00 o.ou 0.01 o.iU 0.01 0.01 0.01 10L 0.05 0.03 o.ou 0.03 o.ou 0.03 0.03 0.00 11E 0.05 o.ou 0.03 0.00 0.00 0.01 0.09 -  •urn*  0.05  0.01  N 0702 0.02 0.38 0.02 0.00 0.00  High l e v e l W  0.00 0.02 0.32 0.06  0.00 0.00 0.30 0.00 O.OU 0.00 0.00 0.02  "oToT 0.01 0.06 0.U5 0.01  0.01 0.U3 0.01 0.00 0.01  0.06 0.06  O.OU 0.02 0.56 O.OU 0.00 0.02  o.oo o.oo  0.72 O.OU 0.02 0.00  1570T 0.01 0.13 0.00 0.00 0.01  o.ou  0.01 0.11 0.00 0.00 0.00  o.ou 0.01 0.16 0.01 0.00 0.00 0706  0.08 0.66 0.02 0.00 0.02  E  0.00 0.02 0.U2 0.00 0.02 0.00 0.01 0.00 o.Ui  0.00 0.00 0.00  0702" 0.02 0.68 0.02 0.00 0.02  ~o7bT0.01 0.00 0.00 0.08 0.10 0.01 0.03 0.00 0.00 0.00 0.00  Key: E = e a r l y bud-opened tree, M = medium bud opened tree, L = late budopened tree, N = North, S = South, W = West, E = East.  90  Appendix N Table 52. Average numbers of l i v i n g and dead insects per needle by l e v e l and tree. Level Low T. No _ L D  1 2 3  k  5 6  7  8 9 10 11 12 13 lk  15 16  17  18 19 20 21 22 23 2k 25 Key: L  Generation 1 Medium D  High  D  Low _L  Generation 2 Medium D  "OF 0.13 "oTIT 0.20 0.01 0^5 "072lT 0.11 0.16 0.36 0.18  0.26 0.19  0.ij6 0.19 0.31 0.27 0.21 0.12 0.07 0.22 0.27 0.19 0.07 0.03 0.21 0.20 0.2k 0.38 0.16 0.36 0.23 0.27 0.72 0.69 0.17 0.10 0.06 0.05 0.03 0.0k 0.03 0.03 0.06 0.05 0.07 0.06 0.03 o.ok 0.13 0.21 0.10 0.11 0.03 o.ok 0.01 0.02 = number of  0.26 0.16  0.15 0.20 0.27 0.10 o.ok 0.20 0.02 0.17 0.07 0.07 0.15  0.81  0.23 0.02 0.01 o.ok 0.03 0.03 0.03 0.36 0.20 0.02 0.01  0.09 0.13 0.21 0.09 0.13 0.16  0.03  0.17 0.09 0.07 0.11 0.66 0.22 0.02 0.00 0.03 0.06 0.05 0.01 0.18 0.12 0.02 0.03  0.02 o.ok 0.03 0.05 0.01 0.00 0.03 0.02 o.ok 0.01 0.01 0.05 0.36 0.13 0.01 0.01 0.03 0.00 0.02 o.ok o.ik 0.10 0.00  0.05 0.03 0.01 0.05 0.03 0.01 0.09 0.02 0.01 o.ok 0.03 0.09 0.66 0.16 0.03 0.01 0.01 0.00 0.03 0.00 0.12 0.12  0.29 0.35 0.52 0.38 0.18 0.08  0.28  0.07  0.08  0.21 0.26 0. 02 1. kk 0.13 o.ok 0.01 0.02 0.05 0.06 0.03 0.21 0.12 0.0k o.ok 0.01  0.07 0.10 0.11 0.08 0.08 0.03 0.07 0.01 0.03 0.06 0.05 0.00 0.15 0.03 0.02 0.01 0.01 0.02 0.02 0.02 0.06 0.06 0.02 0.00  0.12 o.ik 0.19 0.2k 0.13 o.ok 0.09 0.02 0.03 0.03 o.ok 0.02 1.30 0.11 0.02 0.03 0.00 0.03 0.02 0.00 0.29 0.25 0.01 0.00  0.08 0.08  o.ok 0.09  0.07 0.06 0.02  o.ok 0.00 0.02 0.01  o.ok 0.03  0.15  0.06 0.01 0.03 0.03 0.03 0.02 0.00 0.06 0.10 0.01 0.00  l i v i n g i n s e c t s , D = number of dead insects.  High  0.02 0.02 0.01 0.03 0.05 0.05 0.00 0.03 0.03 0.00 0.03 0.00 0.00 0.36 0.07 0.00 0.00 0.05 0.05 0.01 0.05 0.16 0.11  D  0.01 0.01 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.03 0.00 0.00 0.10 0.03 0.00 0.03 0.00 0.00 0.01 0.00 0.05 0.06  0.00 0.00  91  Appendix 0 Table 5 3 . The average number of living insects per needle related to 8 factors. T.No . XI 1 5.5 2 5.U 3 7.9 U 8.7 9.6 5 6 10.8 7 10.3 8 12.1 9 u.u 10 12.8 11 11.1 12 9.7 13  LU  15 16 17 18 19 20 21  22 23 2U 25  3.3 3.3  2.7 6.6 5.8 7.5  U.9 5.2 6.8 0.6 2.2 3.7 2.5  x3 2U 20 20  X2 37 Uo UU  62  26  63  25 29  66 58 59 25 63 53 5U  28 26  27  22 28 25 25 20 20 2U 2U 25 2U  19  21 17 61 67 52 U9 U5 69  26 25 U  2 8  7 22  31 65  2U  x5 18  XU 19  X6  51.3 72.5 79.5 82.3  10 13 16 20 17 19  11 9 11 7  10 32 2U  7  72.0  9 9 6 2  83.0 83.3  12  8  12  22  10  22 9 11  u  10 18 1U 17  18 38 33 37 17 3U 00  11 11 1U  2  u8  8U.1 68.U  90.5 76.5 U7.5 U3.3 36.5 2U.5  1  62.2 50.7 99.9 92.2  Ul  36.9  16  1U  88.9 81.8 86.2 79.7  U8.U  X7 2 3 3  1 3  1 3  1 3 3  1 1 2 1 3  1 3  2 1 2  1 1 3 3 3  XB  u u u 3 3  8 8 8 2 9 9 7 5 5  6 1 1 1 1 1 1 9 9 1 1  Y  0.25 0.35 0.32 0.U8 0.U3 0.23 0.08 0.30 0.08 0.18 0.2U 0.25 0.17 1.65  0.28  0.05 0.03 0.05 0.07 0.07 0.07 0.U6 0.30 0.06 0.01  Key: Bud opening time: 1 = early opened tree, 2 ° medium opened tree, 3 = late opened tree. Location of the tree: 1 = inside growth, 2 = faced South, 3 faced West, U = faced North, 5 faced East, 6 = faced East-North, 7 = faced SouthEast, 8 = faced South-West, 9 open growth. XI = D.B.H. i n inches, X2 = height i n feet, X3 = age, XU = crown width in feet, X5 = clear stem i n feet, X6 = % of living crown, X7 bud opening time, X8 = location of tree, Y = average number of living insects per needle. =  =  e  =  a  92  Appendix P Table 5U. Percentages of mortality by generation and tree. Tree No.  1 2 3 1* 5 6 7 8 9 10 11 12 13 11* 15 16 17 18 19 20 21 22 23 2U 25  Ave.  Generation 1 L+D l^ofD D 0711 0.21  L  Generation 2 "D L + 5 %olB  ~07lT "072TT "51772- ~o~7Hr 0.13 0.31* 38.2 o.U*  0.15 0.10 0.23 0.11 0.21 0.18 0.11 0.08 o.ou 0.12 0.17 0.15 o.oU 0.03 o.U* 0.13 o.i5 0.12 0.15 0.09 0.16 O.U* 0.62 0.68 0.18 0.16 0.03 0.03 0.02 0.01 0.03 0.02 0.03 O.OU o.oU o.o5 o.ou 0.01 0.2U O.U* O.U* 0.11 0.03 0.03 0.01 0.03 0.13 0.11  0.25  0.17 0.31* 0.25 0.39 U6.2 0.22 0.19 U2.1 0.12 0.16 75.0 O.OU 0.32 U6.9 0.13 0.07 1*2.9 o.ou 0.27 1*8.1 O.OU 0.27 1*1* .1* 0.09 0.2U 37.5 0.10 0.30 1*6.7 0.01 1.30 52.3 1.03 0.3U 1*7.1 0.10 0.06 50.0 0.02 0.03 33.3 0.01 0.05 1*0.0 0.02 0.07 57.1 o.ou 0.09 55.6 0.03 0.05 20.0 0.03 0.38 36.8 0.22 0.25 1*1*.0 0.16 0.06 50.0 0.03 o.oU 75.0 0.00 "072TT TToTF 0.13 Uo.o 32 .U  0.05 0.05 0.07 0.05 0.05 0.02 o.ou  0.00 0.02 0.03 0.03 0.01 0.13 0.05 0.01 0.02 0.01 0.02 0.02 0.01 0.06 0.07 0.02 0.00  o.ou  "072T "3371 0.19 26.3 0.22 22.7 0.32 21.9 0.27 18.5 0.17 29.1* 0.06 33.3 0.17 23.5 o.ou 00.0 0.06 33.3 0.12 25.0 0.13 23.1 0.02 50.0 1.16 11.2 0.15 33.3 0.03 33.3 0.03 66.7 0.03 33.3 0.06 33.3 0.05 UO.O o.ou 25.0 0.28 21 .U 0.23 30.U 0.05 UO.O 0.00 00.0 "o7lo~ "2B7J  Generation 1+2 D  "072T 0.35 0.32 0.U8 0.U3 0.23 0.08 0.30 0.08 0.18 0.2U 0.25 0.17 1.65 0.28 0.05 0.03 0.05 0.07 0.07 0.07 0.U6 0.30 0.06 0.01  IT+D  %6fD  0.20 "oTUT 0.18 0.53 31*.0 0.15 0.U7 31.9 0.18 0.66 27.3 0.23 0.66 3U.8 0.13 0.36 36.1 o.u* 0.22 63.6 0.19 0.U9 38.8 0.03 0.11 27.3 0.15 0.33 U5.5 0.15 0.39 38.5 0.12 0.37 32 .U 0.15 0.32 U6.9 0.81 2.U6 32.9 0.21 O.U9 U2.9 o.ou 0.09 UU.U 0.03 0.06 50.0 0.03 0.08 37.5 0.06 0.13 U6.2 0.07 o.U* 50.0 0.02 0.09 22.2 0.20 0.66 30.3 0.18 0.U8 37.5 0.05 0.11 1*5.5 0.03 O.OU 75.0 0.15 o.ui U0.6  Key: L = number of l i v i n g insects per needle, D' = number of dead insects per needle.  93  BIBLIOGRAPHY 1. Balch, R. E., and G. R. Underwood. 1 9 5 0 . The l i f e - h i s t o r y of Pineus p i n i f o l a e (Fitch) (Homoptera: Phylloxeridae) and i t s e f f e c t on white pine. Can. Ent. 82j 117 - 1 2 3 . 2 . Balch, R. E. 1 9 5 2 . Studies of the Balsam woolly aphid, Adelges piceae (Ratz) and i t s e f f e c t s on Balsam f i r , Abies balsamea (L.) M i l l . Dept. of Agr., Ottawa, Canada. 3 . Botvay, K. 1953..  Az altalanos meteorologia, idojarastan es eghajlattan a l a p j a i . Unpublished, U n i v e r s i t y of Sopron.  k. Cameron, A. E. 1936. Adelges cooleyi G i l l e t t e (Hemiptera, Adelgidae) of the Douglas f i r i n B r i t a i n Completion of i t s l i f e cycle. Ann. Appl. B i o l . 2 3 : 585 - 6 0 5 . 5. Cameron, A. E. 1936. The present of status of the Douglas f i r woolly aphids (Adelges cooleyi G i l l e t t e ) i n B r i t a i n . Forestry 1 0 : 133 - 3l*2. 6 . Chystal, R. N. 1916. The l i f e h i s t o r y of Chermes cooleyi G i l l e t t e i n Stanley Park, Vancouver, B.C. Ent. Soc. Ont. k 6 t h Ann. Rept.: 123 - 1 3 0 . 7 . Chrystal, R. N. 1 9 2 2 . The Douglas f i r chermes (Chermes c o o l e y i ) . For. Comm. B u l l . k. 8. Cochran, W. G., and G. M. Cox. 1957. Experimental Designs. 2nd Ed., John Wiley and Sons, Inc., New York, pp. 601. 9 . Cumming, M. E. P. 1 9 5 6 . The Adelginae of Alberta, i d e n t i f i c a t i o n and investigation of l i f e h i s t o r i e s . I . Adelges cooleyi ( G i l l . ) . Unpublished report. For. B i o l . Lab. Calgary, A l t a . 10.  Cumming, M. E. P. 1 9 5 9 . The Biology of Adelges cooleyi ( G i l l . ) (Homoptera: Phylloxeridae). Can. Ent. 9 1 : 601 - 6 1 7 .  1 1 . Forest Club. U.B.C. 1 9 5 9 . Forestry Handbook f o r B r i t i s h Columbia. 2nd Ed. The Forest Club, U.B.C., Vancouver 8 , B.C. Canada, pp. 8 0 0 . 12.  Forestry Commission. 1 9 k 6 . Adelges attacking Spruce and other conifers. L e a f l e t No. 7 .  13.  Forestry Commission. 19U7. Adelges cooleyi an insect pest of Douglas f i r and S i t k a spruce. L e a f l e t No. 2 .  Ik. Francke - Grosmann, H. 1 9 5 0 . Die Douglasienlaus G i l l e t t e e l l a cooleyi ( G i l l . ) C.B. als Schadling der S i t k a f i c h t e . ForstWissenschaftliches Centralblatt. 69 ( 8 ) . k83 - U 9 3 . 15. Geiger, R. 1 9 5 0 . The Climate near the ground (Tanslated from the second German e d i t i o n of Das Klima der Bodennahen L u f t s c h i c h t ) . Harvard U n i v e r s i t y Press Cambridge,  9k  Massachusetts,  pp. U82.  16. G r i f f i t h , B. G. I960. Growth of Douglas f i r at the University o f B r i t i s h Columbia Research Forest as related to climate and s o i l . U.B.C, For. B u l l . 2. 17. Gyorfi, J . 1957.  Erdeszeti rovarton. pp.  Akademiai Kiado, Budapest.  670.  18. Herrick, G. W., and T. Tanaka. 1926.  The Spruce Gall-aphid.  Cornell  University, Agr. Exp. Sta. B u l l . k5k. 19. Imms, A. D. 1925.  A General Textbook of Entomology. 9th Ed.  and Co. Ltd., London,  20.  Methuen  pp. 886.  J e f f e r s , J . N. R. i960. Experimental design and analysis i n f o r e s t research. Almqvist & Wiksell, Stockholm, pp. 16k.  21. Kendrew, ¥. D., and D. Kerr. 1955- The Climate of B r i t i s h Columbia and the Yukon T e r r i t o r y . Edmond C l o u t i e r , C.M.G., O.A., D.S.P., Ottawa, pp. 222. 22.  Laing, F. 1929.  23.  Smith, D. N. 1956. Studies on Adelges cooleyi G i l l , i n r e l a t i o n to two-year Douglas f i r seedlings. Unpublished report. For. B i o l . Lab. V i c t o r i a , B.C.  2k.  Snedecor, G. W. 1956.  The Douglas f i r Adelges.  S t a t i s t i c a l methods. 5th Ed.  College Press, Ames, Iowa. 25.  Scot. For. J . k3: k - 10.  Iowa State  pp. 53k.  Steel, R. G. D., and J . H. Torrie. I960. P r i n c i p l e s and procedures of S t a t i s t i c s with special reference to the b i o l o g i cal sciences. McGraw-Hill Book Company, Inc., New York, Toronto, London, pp. k 8 l .  26. Teucher, G. 195k. Die Douglasienwollaus. ( G i l l e t t e e l l a cooleyi Gill.). I n s t i t u t f u r Forstwissenschaften Eberswalde. Merkblatt. Nr. 15. 27. Uvarov, B. P. 1931.  Insects and Climate.  The transactions of the  Entomological Society of London. 79: pp. 2k2. 28. Varty, I. W. 1956.  Adelges insects of S i l v e r f i r s .  B u l l . 26.  For. Comm.  Appendix  L  C r o w n - s h a p e s of three differently located trees  Interior growth tree (No- 25)  Fig-15 Relationship between the number of living insects per needle and height on the living crown a>  T3  c o CD  (A = interior growth tree, B = marginal growth tree)  z 0-4r CD CL  % 0-31CD co  _c  2 0-2 o 01 6  CD  >  <  15-20 20-25 25-30 30-35 35-40 40-45 45-50 5 Height from the Ground on Tree No-25 — Feet  0-5  5-10  10-15  0-5  5-10  10-15 15-20 20-25 25-30 30-35 35-40 40-45 45-50 50-55 55-60 Height from the Ground on Tree No- 4 — Feet  50  U7 Fig-13  Relationship between number needle a n d d i s t a n c e from  of living insects per  t h e stem a t the low  Distance from the Stem — Feet  F i g - II  Relationship between number  of living insects per needle a n d height on the living  June  32-5  375  5  42-5  Height from the Ground — Feet  crown  Fig-10  Relationship between twig lengths and of l i v i n g insects per  needle  number  Log Sampling Period 01  02  0-3  0-4  0-5  0-6  07  0-8  0-9  10  0-7  08  0-9  10  Log- Sampling Period 0 1  Fig  9  0-2  0-3  Transformed  0-4  0-5  0-6  curves of fig- 7 ( B ) a n d f i g - 8 ( A )  1 July 9 Fig-8  ° i6 Number  23  30  Aup 13 20 Sampling Period  27  Sept-3  10  17  24  o f l i v i n g a n d d e a d i n s e c t s p e r n e e d l e in g e n e r a t i o n  for different periods a t the low level o f t h e living c r o w n  2  27  \  \  \  \  \  \  \  \ Number of living insects  dead \rv Number o*  July 9  7  16  Number  23  I  30  I  Aug 6  I  L  13 20 27 Sampling Period  Sept 3  of living a n d dead insects p e r n e e d l e  for different  periods  10  17  in generation  24  2  2$  June 4  il  18  25  July 2  Sampling Period Fig- 6  Number  o f l i v i n g a n d d e a d i n s e c t s p e r n e e d l e in  generation  I f o r different periods at the low level  of t h e l i v i n g  crown  June 4 Fig- 5  Number  II  18 Sampling Period  25  July 2  o f living a n d d e a d insects p e r n e e d l e in  generation I f o r different  periods  10  Fig- 4  A,  Sistens nymph,first  B,  Progrediens  adult  instar  (After  Chrystal)  Fig- 2  A,  Gallicola  eggs B,  migrans with wings closed laying  on f i r n e e d l e  Egg-mass  progrediens  (X  20)  p r o d u c e d by Gallicola (X  30 )  or  Fig-1  The  life c y c l e of  Adelges  cooleyi  

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