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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 Br i t i s h Columbia (Sopron Division), 1959 A THESIS SUBMITTED IN PARTIAL FULFILMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF FORESTRY in the Department of FORESTRY We accept this thesis as conforming to the required standard THE UNIVERSITY OF BRITISH COLUMBIA April, 1961 I n p r e s e n t i n g t h i s t h e s i s i n p a r t i a l f u l f i l m e n t o f t h e r e q u i r e m e n t s f o r a n a d v a n c e d d e g r e e a t t h e U n i v e r s i t y o f B r i t i s h C o l u m b i a , I a g r e e t h a t t h e L i b r a r y s h a l l m a k e i t f r e e l y a v a i l a b l e f o r r e f e r e n c e a n d s t u d y . I f u r t h e r a g r e e t h a t p e r m i s s i o n f o r e x t e n s i v e c o p y i n g o f t h i s t h e s i s f o r s c h o l a r l y p u r p o s e s m a y b e g r a n t e d b y t h e H e a d o f my D e p a r t m e n t o r b y h i s r e p r e s e n t a t i v e s . I t i s u n d e r s t o o d t h a t c o p y i n g o r p u b l i c a t i o n o f t h i s t h e s i s f o r f i n a n c i a l g a i n s h a l l n o t b e a l l o w e d w i t h o u t m y w r i t t e n p e r m i s s i o n . D e p a r t m e n t o f T h e U n i v e r s i t y o f B r i t i s h C o l u m b i a , V a n c o u v e r 8, C a n a d a . D a t e Apn'1 10, 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 . This was done through the observation of populations in the f i e l d , supported by some labora-tory work designed to show, that certain influences are important. The work was 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.) Inter tree differences, 2.) Intra tree variations, 3.) Population changes with time, h») Mortality of the insect. Abundance was affected by extrinsic influences on the trees, such as location and exposure and in 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 factors, resulting in high abundance of the insect i n the pe-ripheral part of the lower crown. The average number of l i v i n g insects decreased with time rec t i l i n e a r l y in generation 1 (Sexuparae and Progredientes) and logarithmically i n gener-ation 2 (Neosistens). A c r i t i c a l period during establishment of generation 2 caused the logarithmic changes. Mortality estimates by direct counts were subject to a large error be-cause many of the dead insects f e l l off. i i i ACKNOWLEDGEMENTS Acknowledgement i s made to the University of British Columbia for the laboratory and education f a c i l i t i e s , which aided materially i n this research. Grateful appreciation i s expressed by the writer to the members of the staff of the Department of Zoology and Faculty of Forestry, expecially to Dr. Kenneth Graham for advice and encouragement throughout the work,and to Dr. J. Harry G. Smith for counsel on s t a t i s t i c a l analyses. Special thanks are due Mr. Jozsef Csizmazia for his help i n evaluating some of the s t a t i s t i c a l analyses, using the ALwac III E electronic computer. i v CONTENTS Page TITLE PAGE i ABSTRACT i i ACKNOWLEDGEMENT i i i CONTENTS iv TABLES v i FIGURES ix INTRODUCTION 1 MATERIALS AND METHODS 3 1. THE INSECT 3 2. THE STAND Ik 3. DESCRIPTION OF SAMPLE TREES AND SAMPLING METHODS. . . . 15" h. PROCESSING OF DATA 18 EXPERIMENTAL RESULTS 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 DISCUSSION 57 1. INTER TREE DIFFERENCES 57 LOCATION OF THE TREES 57 EXPOSURE 58 BUD OPENING TIME 60 TWIG LENGTH 61 2. INTRA TREE VARIATIONS 62 EXPOSURE 62 Page HEIGHT ON LIVING CROWN ~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 7 0 BIBLIOGRAPHY 93 v i TABLES Number Heading Page 1 Relative numbers of sexuparae and progredientes on Douglas f i r in three counts 7 2 Analysis of variance for generation 1 20 3 Analysis of variance for generation 2 21 h Calculation of the equation and correlation coefficient of variables by period and average number of l i v i n g insects per needle 23 5 Calculation of the equation and correlation coefficient of variables by period and average number of dead insects per needle. 2h 6 Calculation of the equation and correlation coefficient of the number of l i v i n g insects per needle at low crown level of the trees on period 26 7 Calculation of the equation and correlation coefficient of the number of dead insects per needle at low level of the trees on period 26 8 Calculation of the equation and correlation coefficient of average number of l i v i n g insects per needle on time. . . . 30 9 Calculation of the equation and correlation coefficient of average number of dead insects per needle on time. . . . 31 10 Calculation of the equation and correlation coefficient of average number of l i v i n g insects per needle on time. . . 32 11 Calculation of the equation and correlation coefficient of average number of dead insects per needle on time. . . . 32 12 Analysis of variance among trees, levels and generations. 33 13 Differences among trees by abundance of Adelges cooleyi. . 3U lU Influence of position of tree i n the stand on abundances of Adelges cooleyi 35 15 Differences among trees by abundance of Adelges cooleyi. The trees were treated as two population, separated by location 36 16 Average numbers of l i v i n g insects per needle, by exposures and levels for the marginal trees 37 v i i Number Heading Page 17 Analysis of variance among exposures and levels 38 18 Differences between differently exposed trees by the abun-dance of Adelges cooleyi 38 19 Influence of 8 factors on the average number of l i v i n g insects per needle kO 20 Calculation of the equation and correlation coefficient of average number of l i v i n g insects per needle on height from the ground (data of June 5) k l 21 Calculation of the equation and correlation coefficient of average number of l i v i n g insects per needle on height from the ground (data of July 25) k3 22 Calculation of the equation and correlation coefficient of average number of l i v i n g insects per needle on height from the ground (data of September 26) kk 23 Calculation of the equation and correlation coefficient of average twig length on height from the ground k6 2k Calculation of the equation and correlation coefficient of average numbers of l i v i n g insects per needle on distances from the stem (data of June 5) k8 25 Calculation of the equation and correlation coefficient of average numbers of l i v i n g insects per needle on distances from the stem (data of July 25) k8 26 Calculation of the equation and correlation coefficient of average number of l i v i n g insects per needle on distances from the stem (data of September 26) k9 27 Calculation of the equation and correlation coefficient of average lengths of twigs on distances from the stem. . . k9 28 Analysis of variance of dead insects among trees, levels and exposures (generation 1) 51 29 Analysis of variance of dead insects among trees, levels and exposures (generation 2) 52 30 Analysis of variance of dead insects among trees, levels and generations 53 31 Variance ratio test between the main factors and f i r s t order interactions 53 32 Percentage of mortality by l o c a l i t y of the trees and generations 5k v i i i Number Heading Page 33 Percentage of mortality by levels and generations. . . . 5k 3 k - 5 k See: L i s t of Appendices 7 0 ix FIGURES Number Heading Page 1 The l i f e cycle of Adelges cooleyi k 2 A, Gallicola migrans with wings closed laying eggs on f i r needle (X20) B, Egg-mass produced by Gallicola or progrediens ( X 3 0 ) . . 6 3 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 h A, Sistens nymph, f i r s t instar B, Progrediens adult 10 5> Number of l i v i n g and dead insects per needle i n generation 1 for different periods 22 6 Number of l i v i n g and dead insects per needle i n generation 1 for different periods at the low level of the l i v i n g crown 2$ 7 Number of l i v i n g and dead insects per needle i n generation 2 for different periods 27 8 Number of l i v i n g and dead insects per needle i n generation 2 for different periods at the low level of the l i v i n g crown 28 9 Transformed curves of f i g . 7 (B) and f i g . 8 (A). . . . 29 10 Relationship between twig lengths and number of l i v i n g insects per needle 39 11 Relationship between number of liv i n g insects per needle and height on the l i v i n g crown U2 12 Relationship between twig length and height on the l i v i n g crown U5 13 Relationship between number of l i v i n g insects per needle and distance from the stem at the low level of the l i v i n g crown. U7 lh Relationship between twig length and distance from the stem at the low level of the l i v i n g crown 50 15 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 Sitka 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 varia-ble i n numbers as to merit examination of the factors influencing i t s abun-dance. The insect and i t s secondary host, Douglas f i r , Pseudotsuga Menziesii var. Menziesii (Mirb.) Franco, offer a unique opportunity for the study of environmental influences on an insect. The insect lends i t s e l f well to observation and sampling because i t i s sessile during most of i t s l i f e , and i s thus suitable for quantitative work. Douglas f i r i s so variable i n i t s phenology that a wide range of phase relations between i t and iU cooleyi may be found. The cooleyi spruce g a l l aphid was named and described by Gillette i n 1907. He called the generations on spruce: Chermes cooleyi, and generations on Douglas f i r Chermes cooleyi var. cowenii. In 191U ChrystaL's (6) experi-ment proved that, Gillette's Chermes cooleyi var. cowenii i s the colonici stage of Chermes cooleyi. Annand changed the name of the subfamily to which the species belongs i n 1928, from Chermisinae to Adelginae, because the former name had been earl i e r used for the psyllids groups (7). No detailed work has been done about the factors influencing the abundance of iU cooleyi on Douglas f i r . The fact 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 just after 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 chiefly on the trees near the margin and was not present i n the centre of the planta-tion. He has also stated that the insects were more frequent on the lower and middle branches of the trees. Both in England and Germany i t has been established that, Pseudotsuga Menziesii var. glauca (Beissn) Franco i s more resistant to cooleyi than the Pseudotsuga Menziesii var. Menziesii (Mirb.) Franco. The main biotic factors - affecting the abundance of A. cooleyi - are the natural enemies. According to the present knowledge the insect i s attacked by comparatively 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 cavity on spruce. The yellow and black 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 on A. cooleyi. Gyorfi (17) has mentioned that Hymenopterous parasites of the family Chalcididae attack certain species of European Adelges. Heavily infested Douglas f i r s are not k i l l e d directly by Adelges, but are so reduced i n vigour that they are more susceptible to attack of second-ary insect and disease. The cumulative effect of the attack probably reduces the rate of growth, reducing the active surface of assimilation, resulting 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 Sitka spruce mixture not be planted on unsuitable site or l o c a l i t i e s . 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 Sitka spruce i s just after 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 their abundance. Adelges cooleyi prefers two different species «of host-trees i n i t s complete l i f e cycle. In the Pacific Coast the primary host i s Sitka 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" (Fig. l)s Fundatrix Generation: The first-stage larva of fundatrix overwinters on a Sitka spruce terminal twig adpressing i t s body to the twig, and sunking i t s stylet deeply into the tissue through bark fissures. Passing through the winter i n this stage, they become active i n the f i r s t part of April, and begin to feed. The fundatrices are reported to have three la r v a l instars (9, 10). These lar v a l stages are present for a relatively short time i n the spring. The f i r s t - i n s t a r larvae are dark brown with slightly greenish colour. The general appearance the second-instar larvae resemble the adults, having thinner cuticle. 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 April. Gallicola Migrans: The gallicolae migrantes are the gall-forming generation on spruce. This generation i s the progeny of fundatrices. This summer generation hatches about 10 - 15* days after the eggs are deposited by the stem-mothers, and migrates to the inner bases of young needles, which are just breaking from the bud at this time. As soon as they have settled, they begin to feed, and galls begin to 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 varies i n length from 1 - 5 inches, varying with the vigour of the twig attacked (18). 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 instars within the galls (9, 10). The reddish-brown coloured larvae are covered with a fine powdery wax. The wing pads can be seen at the third-instar. They emerge from the galls i n the fourth-instar and settle on the needles, and moult to the adult stage. The parthenogenetic adults are reddish-brown with a well sclerotized thorax. They f l y to the Douglas f i r , where being settled for a few days, they secrete wax from the anterior margin of the head and the abdomen. The wax from the abdomen and the wings protects the eggs which they lay. The average number of eggs l a i d was found to be 65 of 10 counts (9, 10) (Fig. 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, staid on Douglas f i r on the previous summer. Chrystal (6) says that the sistens hatch 6-7 days after the oviposition. This period has been stated by Cumming (10) as about three weeks. In the present study the time required i n I960 was 2 or 2\ weeks. The youngs settle on the lower surface of Douglas-fir needles, and pass the winter i n this stage. This dormant form of sistens generation used to be called neosistens (Fig. h). This generation was one subject i n the experiment, and was sampled 8 times during the summer of I960. This generation w i l l be called generation 2 i n the further part of the thesis. The neosistens are light-brown with a darker prothorax and head. It i s unlikely that this 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 after being hatched, looking for suitable needles to overwinter, where they press their bodies to the underside of the needles, inserting their stylets into the raesophyll tissue. This insertion i s believed to afford additional anchorage, rather than providing food. When settled, 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. The body of the second-instar larvae i s covered with wax. The adults are similar to the second-instar larvae but with more wax. 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 to spruce. The proportion of these two forms varies widely. Ghrystal (6) found that about $0% of 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 in three counts. Date , Number counted Per cent of Sexuparae Progredientes Doubtful J- 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 after oviposition. Hatching begins about the time when the buds start to open. After hatching they move to the needles -•-Larva too small to distinguish form. 8 and develop into 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 gallicolae, but the gallicolae have more glands. Wax can be found on the dorsal and ventral surfaces of the head during the l a r v a l stage. There are four larval instars, three of them similar i n morphology of the two forms. Wing pads are present on the fourth instar 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 no reason to treat them separately for the subject of the experiment. This generation w i l l be called as generation 1 i n the further part of the thesis. Sexuparae t The adults of sexuparae are winged parthenogenetic females (Fig. 3). Wax i s present on a l l parts of the body, except the ventral surface of the abdomen, the adults are heavily sclerotized. 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, settle on the needles, then lay 5-20 eggs (10). The eggs are protected by wax and the folded wings of the female. Progredientes: The adults of progredientes are wingless parthenogenetic females (Fig.U), resembling the sistens. They stay on Douglas f i r . The number of eggs l a i d by Progredientes i s variable. Chrystal (6) has stated that they lay 30 - UO eggs. The average number of eggs i s given as 15 with a range from 3 to 25 i n 11 counts by Cumming (10). Fifteen 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, I960. The average of these was 27, with a range from 5 to k3. Progredientes hatch 2 - 3 weeks after oviposition, and overwinter i n this undeveloped stage (Neosistens). Consequently a subcycle of the l i f e cycle can be found on Douglas f i r (Fig. 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 reproduc-tion occurs. This generation i s not described well i n most of the literature. Chrystal (6) did not observe this generation i n B. C , but later (7) he found the generation i n Britain. S t i U later Annand in California, Cameron (k) i n Britain, Francke-Grosmann (lk) i n Germany and Cumming (10) in Alberta recorded the sexual generation. The most detailed report on this generation was given by Cumming (9, 10). According to him they hatch a week or more after the oviposition of the sexu-parae. Chrystal (6), and Cameron (k) found $ larval instars. Cumming (10) distinguished 3 only. I t i s possible that the f i r s t two instars can be so similar, 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 far as k - 5 years-old twigs. This habit of moving back helps them i n the mating, helping the male find the female. They lay eggs between the old bud scales and twigs. The female secretes wax, which surrounds but does not cover completely the egg. It 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 their shapes and sizes. There are also differences i n the arrangement of the plates on which the gland facets occur. The ventral plates are slightly sclerotized, and always have small round gland facets. The dorsal plates are heavily sclerotized, being i n three pairs, showing a single transverse row on each segment. This row i s arranged from the mesothorax to the sixth or seventh segment of the abdomen. Two rows can be found on the head and prothorax. The central pair of glands on the dorsal surface i s called "mesial", the one beside them "pleural", and the later 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 well outlined except those on the sixth and seventh segments of abdomen. The posterior plates of the adults have a higher propor-tion of round gland facets. They also have angular gland facets which are not so large as those of the sistens and progredientes (10). Gallicola 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 cooleyi. The number of gland areas increases instar after instar. The fourth-instar resembles the adult, but the gland areas are smaller on the abdomen. The gland facets of the dorsal surface of the adult are round. The gland areas of the head and thorax are constant. These areas vary on the abdomen. The mesial areas of the abdomen are fused at the centre l i n e , and the pleural areas are broken up into small groups of gland facets. The hamuli on the hind wing varies from 2 to 5. Sexupara: The f i r s t - i n s t a r larva has setae surrounded by small plates. The adults have large variation 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 (Fig. 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 larva can be distinguished by the appearance of the plates. Two sclerotized areas cover the head which are separated at the centre l i n e . Gland facets can be found on these areas anteriorly and l a t e r a l l y . On each side of the prothorax a large sclerotized area can be found, each of them i s fused from four plates. The mesial, pleural and marginal areas of the mesothorax, metathorax and the abdominal segments^ from 1 to U have rectangular plates. Gland facets can be seen on the f i f t h , sixth and seventh abdominal plates. The gland facets resemble those of the fun-datrix larvae. 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 recticulations are more conspicuous. The separation of the sistens and progredientes can be done by the gland facets on the anterior mesial plate of the prothorax and by the pleural 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 an-tenna 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 . The last-instar female has a wax gland at the posterior end. The adults are smaller than any of the generations. 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 of the trees are 2k - 26 years old, but some 20 and 28 - 29 years old trees are also present. The oldest class seems to be natural regeneration, and the two younger classes are planted. The main material of the plantation was Douglas f i r , with some Sitka spruce. Some other tree species are also present as a result of natural regeneration. The vegetations of three layers of the forest, as tree crown layer (A), shrub layer (B) and ground vegetation (C) are the follows: The Vegetation of the Forest; A level; crown closure; 95$ Pseudotsuga Menziesii Thuja plicata Tsuga heterophylla Picea sitchensis Taxus brevifolia ALnus rubra Acer macrophyllum Prunus sp. B level: closure 20 - 2$% Gaultheria shallon Sarabucus pubens Rubus spectabilis Rubus vite f o l i u s Rosa gymnocarpa Rubus parviflorus Symphoricarpus rivularis Vaccinium parviflorum Physocarpus capitatus C level: 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 triflorum Dicentra formosa Pseudotsuga Menziesii V Tsuga heterophylla Acer macrophyllum In the southern part of the forest most of the B and C level vegetation has been removed. The s o i l of this part i s tramped down, having a thick level of raw humus. Generally the s o i l of the forest has been developed on sandy deposits of the Fraser River, to a podzolic s o i l . The surface layers of the s o i l are: Ao, 1 inch, A l , 2 inches, and A2, I2 inches. The "Bn or depositional layer i s also present but i t was too deep to dig down to "CM or basic layer to measure "B". I t appears that the forest type - indicated by the site 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 things: 1. Location of the treej three classes were established: a. ) Trees growing at the center of the forest. b. ) Trees growing at the different margins of the forest. c. ) Open-growth trees. Eight of the 25 trees were chosen at the center of the forest, 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 differenciate the trees: a. ) Early (E): the buds were open before May 15. b. ) Medium (M): the buds opened from May 15 to May 21. c. ) Late (L): the buds opened later than May 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 for selecting for these v a r i -ables was limited, because the forest consists of planted trees of limited variation i n these factors. Both of the progrediens and sistens (Neosistens) settled on the current year's growth, consequently the new growth of foliage was sampled. The basic sample unit was established as a needle. Samples were drawn from three levels of the l i v i n g crown of each tree. The l i v i n g crowns of the trees were visually separated into three levels (Low, Medium, High), and the samples were drawn from the middle of each level (Appendix E). North, East, South and West sides of the trees were sampled i n each l e v e l . The sample© were drawn from the outer part of the crown. Consequently 12 sample twigs were cut from a normal tree. Some of the trees are not normal in crown shape (Appendix L ) , lacking some part of the crown. Most of the trees on the margins are of this kind. In these cases the 12 samples could not be drawn from one tree. 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 level and sometimes even i n the medium l e v e l . These levels 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 this time only once per two weeks or once per three weeks. The change of the length of sampling period was suggested by the low activity of insects after August 6, because by this 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 in the lower levels 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 15 - feet - long pieces, working telescopically. This second prunner was not useful over 3 0 - 3 5 feet i n length, because i t was very fl e x i b l e . For sampling the higher trees a ladder was used, which could be extended to 3h feet from which pole prunners could be operated. Beyond the reach of prunners, samples were obtained by climbing the tree. 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 5 were counted on ten randomly chosen needles of each twig. Differentiation between dead and l i v i n g larvae was made with the aid of a microscope. The information was kept separately for l i v i n g and dead insects (Appendix A, B). The different stages and instars 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 July 2 , and the neosistens generation was the material from July 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 tree. The samples by height were drawn from every 5 feet of the outer part 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 in 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 will be discussed in the next chapter. Some of the measured data were not analysed, but they will be used to explain the effect of several factors. 2 0 EXPERIMENTAL RESULTS: Six trees - No. 1 , 2 , 1 0 , 1 1 , lh and 2 0 - of the 25 could be sampled i n each level and at every cardinal points. The abundance of the li v i n g insects on these 6 trees was analyzed by three factors: le v e l , exposure (cardinal points) and bud-opening time (Appendix M), separately for the two generations (Table 2 and 3 ) . Table 2 . Analysis of variance for generation 1 . Source Degrees of freedom Net sum squares Mean sura squares Variance ratio (F) S i g n i f i -cance at 0 . 0 5 level Levels (L) 2 0 . 7 4 0 . 3 7 7.40 H.S.* Exposures (E) 3 0 . 0 3 0 . 0 1 - . N.S. Bud opening (B) 2 1 . 2 1 0.61 1 2 . 2 0 H.S. L x E 6 0 . 0 5 0 . 0 1 - N.S. L x B h 0.14 O.Oij - N.S. E x B 6 0 . 0 8 0 . 0 1 N.S. L x E x B 12 0 . 1 0 0 . 0 1 - N.S. Residual 36 1 . 9 4 0 . 0 5 Total 71 4 . 2 9 * H.S.=highly significant, S i g n i f i c a n t , N.S.=Not significant. The levels and the bud-opening times show significant differences. In table 3 the bud-opening times show also significant difference, the levels do not, but the "F" value of levels i s very close to the significance l e v e l . Examining the factors i n table 2 and 3 , the average numbers of li v i n g insects per needle were greater at the low level (0 . 3 1 i n generation 1 and 0 .39 i n generation 2) than at the medium level (O.2I4 in generation 1 and 0 . 2 8 i n generation 2 ) , and i t was greater at the medium level than at the high level (0.07 i n both generations) of the l i v i n g crown. The average number of li v i n g insects was also greater on the early bud-opening trees (O .38 i n 21 Table 3. Analysis of variance for generation 2. Source Degrees of freedom Net sum squares Mean sum squares Variance ratio (F) S i g n i f i -cance at 0.05 level 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 It generation 1 and 0.56 i n generation 2) than on the medium (0.17 i n generation 1 and 0.09 i n generation 2) and late bud opening trees (0.07 i n generation 1 and 0.09 i n generation 2). Both of the analyses of variance indicate that there i s no significant 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 five sampling time of the 13 covered the generation 1, and the remainder covered the generation 2. a. Population changes with time in generation 1: F i r s t , the correlation between number of insects per needle and time was tested for the 6 trees mentioned. The average number of insects per needle was plotted by time (Fig. 5). The numbers of dead and l i v i n g insects were plotted separately. The points suggested the calculation of linear-regression equa-tions for both numbers of l i v i n g and numbers of dead insects (Table U, 5). 22 June 4 II 18 25 July 2 Sampling Period Fig 5 Number of living and dead insects per needle in generation I for different periods 23 Table lu Calculation of the equation and correlation coefficient 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 coefficient, a = intercept. r = correlation coefficient, t = " t " test for correlation coefficient. Sb • standard deviation for regression, i = confidence intervals. The equation of the regression line i s : Y = -O.OU9X+0.31 The correlation i s significant between the average number of insects and periods. The values of "r" and " t " are highly significant. The average number of insects per needle decreased linearly with time. There i s no significant correlation between the average number of dead insects per needle and time (Table 5), but points f i t well to the l i n e . The unsignificant result probably was caused by the small sample size. Logically, the slope of the mortality line should be the reverse of the line of li v i n g insects, but this i s not so, for two reasons: i . about 50$ of the insects of this generation develop into winged Table 5. Calculation of the equation and correlation coefficient of variables by period and average number of dead insects per needle. t Y —yT~ Y*— XY 0 0.21 0 o.ouui 0.00 1 0.17 1 0.0289 0.17 2 0.20 a O.OiiOO 0.1*0 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 0.20 X = periods, Y = number of dead insects per needle, b " 0 . 0 0 7 , a m 0.19, r - 0.51, t - 1 . 0 2 , Sb • 0.00686, -0.015 6 i * + 0 . 0 2 9 . The equation of the regression line 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 their 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 off, and also many of them were eaten by predators. The correlation between number of insects per needle and period, was extended for every tree sampled. The average numbers of insects at the lower crown level of the trees were used, because they were most abundant there (Fig. 6) (Appendix C) (Table 6, 7 ) . The " t " and " r " show that (Table 6) the correlation i s significant between the number of l i v i n g insects per needle and time. The lack of correlation (Table 7) between the number of dead insects per needle and time probably i s caused by the small sample size. b. Population changes with time i n generation 2: Eight samples out of 13 were drawn from generation 2 . The data of these samples were treated i n the same way as in generation 1 . 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 coefficient of the number of l i v i n g insects per needle at low crown level of the trees on period. X Y XY 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 0.20 Total Ave. X = periods, Y - numbers of insects per needle, b • -0.0U1, a •= 0.28, r - -0.960, t • -5.89, Sb - 0.00775, -0.063 6 1 £ -0.019 Equation: Y = -O.OiaX+0.28 Table 7. Calculation of the equation and correlation coefficient of the number of dead insects per needle at low level of the trees on period. X Y — X 2 " y2 XY 0 0.15 0 0.0225 0.00 1 0.12 1 O . O U 4 U 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 Total Ave. 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 a> E Z a> cr> o > < 0-\ \ \ \ \ \ \ J L Number of living insects Number July 9 16 23 30 Aug 6 13 20 27 Sept 3 10 17 24 Sampling Period F i g 7 N u m b e r 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 2 f o r d i f f e r e n t p e r i o d s 28 j u l / § ife 23 30 Aug^ 13 20 27 Sep t l iO 17 2 4 ~ Sampling Period 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 02 03 0-4 0-5 0-6 07 08 0-9 10 Fig 9 Transformed curves of fig 7 (B) and fig-8(A) 30 time (Fig. 7 ) . Between the first and second and between the second and third sampling periods the line of the living insects is increasing, because these periods were the hatching time. From the third point, the line starts to decrease. The manner of the points suggested to leave out the fi 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 logX logY (logX)2 (logY)2 ' (logX)(logY) 1 0 . 3 3 0 . 0 0 0 -O.U 81 0 . 0 0 0 0 0.2314 0 . 0 0 0 0 2 0 . 3 1 0 . 3 0 1 - 0 . 5 0 9 0 . 0 9 0 6 0 . 2 5 9 1 - 0 . 1 5 3 2 3 0 . 2 2 0 . 4 7 7 - 0 . 6 5 8 0 . 2 2 7 5 0 . 4 3 3 0 - 0 . 3 1 3 9 5 0 . 1 9 0 . 6 9 9 - 0 . 7 2 1 O.U 886 0 . 5 1 9 8 - 0 . 5 0 4 0 8 0 . 1 6 0 . 9 0 3 - 0 . 7 9 6 0 . 8 1 5 4 0 . 6 3 3 6 - 0 . 7 1 8 8 10 0 . 1 7 1 . 0 0 0 - 0 . 7 7 0 1 . 0 0 0 0 0 . 5 9 2 9 - 0 . 7 7 0 0 T# 29 1 . 3 8 3 . 3 8 0 - 3 . 9 3 5 2.6221 2 . 6 6 9 8 - 2 . 4 5 9 9 A* U .83 0 . 2 3 0 . 5 6 3 - 0 . 6 5 6 T* = Total A* m Average X = periods, Y = numbers of living insects per needle, b = - 0 . 3 3 9 , a = O .U65, r - - 0 . 9 6 3 , t = - 6 . 8 0 , Sb - 0 . 0 U 9 , - 0 . 2 0 3 * i 1 8 -O.U65 Equation: logY = -0.3391ogX-0.U65 Both the values of M r " and "t" give significant correlation between the transformed "Y" and "Xn, meaning that the correlation is logarithmic between 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 in the generation 2 (Fig. 7 ) (Table 9 ) . Significant correlation was found between the average.number of dead 31 insects per needle and time. The correlation 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 level of each tree was used, because insects were most abundant there (Appendix C, D). Table 9« Calculation of the equation and correlation coefficient of average number of dead insects per needle on time. X Y p 8- 1 XY 0 0.03 0 0.0009 0 .00 1 o.ou 1 0.0016 o.ou 2 o.o5 a 0.0025 0.10 3 0 . 0 5 9 0.0025 o . i 5 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 . 9U8 , t - 7 . 2 9 , Sb - 0 . 0 0 0 6 9 0 . 0 0 3 * i & 0 . 0 0 7 Equation: Y - 0 . 0 0 5 X + 0 .0U The average numbers of li v i n g insects had to be tested the same way as in table 8. The f i r s t two points were omitted and both X and Y were trans-formed to logarithms (Fig. 8, 9 ) (Table 1 0 ) . A significant correlation was found between X and Y, meaning that the numbers of l i v i n g insects decreased logarithmically with time (Table 1 0 ) . 32 Table 10. Calculation of the equation and correlation coefficient of average number of l i v i n g insects per needle on time. X Y logX logY ( l o g X ) 2 (logY) 2 (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 lw 83 0.20 0.563 -0.721 Total Ave. 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 correlation coefficient of average number of dead insects per needle on time. Total Ave. X Y • ' X 2 ' XY 0 0.011* 0 0.0002 0.000 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 0.0228 2.325 4.5 0.01*9 33 X = periods, Y ° numbers of dead insects per needle, b e 0.00$, a a 0.0026, r = 0.898, t * 5.0U5, Sb - 0.00126, 0.002 * i & 0.008 Equation: Y • 0.005X+0.026 The rsults (Table 11) show significant correlation between X and Y, meaning that the numbers of dead insects per needle increased with time. 2. Inter Tree Differences: The numbers of li v i n g and dead insects were summarized by tree, genera-tion and level i n tree (Appendix N). Analysis of variance was carried out for the l i v i n g insects (Table 12). Table 12. Analysis of variance among trees, levels and generations. Source Degrees of freedom Net sum squares Mean sum squares Variance ratio (F) S i g n i f i -cance at 0.05 level Compo-nents of 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 T x L x G U8 0.190 o.oouo O.OOliOO Total Hi9 5.861 Table 12, shows a highly significant difference among trees, and among levels. These results indicate the pos s i b i l i t y of significant components of variance. The significant interactions indicate that the factors are not independent and there i s no good biological e x p l a n a t i o n of the interactions. The most economical sample numbers could be calculated from the compo-nents of variance for the factors. Minimum variance would be obtained i f we take a large sample in a large stratum, a large sample when the stratum Table 13. Differences among trees by abundance of Adelges cooleyi. Generation 1 . I 1U 22 1* 5 2 15 II 0.62 0.21* 0.23 0 .21 0.21 0.18 III 19.hit ii.37 3.97 3.56 3.56 1.98 17 H.S. S. s. S. s. N.S. V M 0 M M M M I 10 6 1 21 20 9 II O.lit 0.11 0.11 O.OU O.OU o.ou III 0.U0 - 0 . 7 9 - 0 .79 - 3 . 5 7 - 3 . 5 7 - 3 . 5 7 17 N.S. N.S. N.S. S. s. S. 7 0 M M I I M generation 2 . I lU I* 5 22 3 23 II 1.03 0 .25 0.22 0.22 O.17 0.16 III 22.28 2.97 2.23 2.23 0.99 0.7U 17 H.S. S. s. s. N.S. N.S. 7 M M M 0 M 0 I 11 7 9 10 19 20 II 0.09 o.oi* o.ou o.ou o.ou 0.03 III - 0 . 9 9 - 2 . 2 3 - 2 . 2 3 - 2 . 2 3 - 2 . 2 3 -2 .U9 17 N.S. s. S. s. s. s. 7 0 M M 0 I I feneration 1 + 2 . I 11* U 22 5 2 3 II 1.56 O.U 8 0.U6 o.U3 0 .35 0.32 III 21.55 3 .1a 3.10 2.6U 1.U0 0.93 17 H.S. s. S. s. N.S. N.S. V M M 0 M M M I 6 10 13 7 9 19 II 0.23 0.18 0.17 0.08 0.08 0.07 III -0.1*7 -1.21* - 1 .U0 - 2 . 7 9 - 2 . 7 9 - 2 . 9 5 17 N.S. N.S. N.S. S. S. s. .7 M 0 M M M I 8 13 12 3 11 23 0.17 0.16 0 .15 0 .15 0 . 1 5 O.U* 1.59 1.19 0.79 0.79 0.79 o.uo N.S. N.S. N.S. N.S. • N.S. N.S. M M M M 0 0 7 2U 19 18 16 17 25 o.ou 0.03 0.03 0.03 0.03 0.02 0 .01 -3 .57 - 3 . 9 7 - 3 . 9 7 - 3 . 9 7 - 3 . 9 7 -U.37 -U.76 S. S. S. S. S. S. s. M I I I I I I 2 1 8 6 12 15 o.U* 0.11* 0.13 0.12 0 .10 0 .10 0 .25 0 .25 0 .00 - 0 . 2 5 - 0 . 7 U - 0 . 7 U N.S. N.S. N.S. N.S. N.S. N.S. M M M M M M 21 2U 16 18 13 17 25 0.03 0.03 0.02 0.02 0 .01 0 .01 0 .00 -2 .U9 - 2 .U9 - 2 . 7 2 - 2 . 7 2 - 2 . 9 7 - 2 . 9 7 - 3 . 2 2 S. S. s. S. S. S. S. I I I I M I I 23 8 15 12 1 11 0.30 0 .30 0.28 0 .25 0 .25 0.2U 0.62 0.62 0.31 - 0 . 1 6 - 0 . 1 6 - 0 . 3 1 N.S. N.S. N.S. N.S. N.S. N.S. 0 M M M M 0 20 21 2U 18 16 17 25 0.07 0.07 0.06 0 . 0 5 0 .05 0.03 0 .01 - 2 . 9 5 - 2 . 9 5 - 3 . 1 0 -3.26 -3.26 - 3 . 5 7 - 3 . 8 8 S. S. s. S. S. S. S. I I I I I I I 35 Key for table 13: I " tree No. II = 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 8 5 location of the trees. I = inside growth tree M B margin growth tree 0 e open-growth, tree variance i s Mgh (trees) and a small sample when the stratum cost i s high (level). 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 variation at the ground level of sampling. The differences between trees were examined by the " t " test. Table 13 shows that most of the highly populated trees are edge trees, and the lower populated trees are inside the stand. By these results, the trees were separated into three populations, interior, 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 " test (Table l h ) . Table 1U. Influence of position of tree i n the stand on abundance of Adelges  cooleyi. Values of t Location of trees. Interior Marginal Open Interior — — Marginal 21.1»8 H.S. — — Open 5.19 s. 2.13 N.S. — Table i i * shows that the abundance of the Adelges cooleyi on growing with-i n a stand i s significantly different from those on the stand edge, or grown Table 15. Differences among trees by abundance of separated by location. Generation 1 . Margin growth trees. I 7 9 1 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 . 8 2 - 0 . 7 8 - 0 . 7 8 17 S. S. N.S. N.S. N.S. N.S. 7 L L M E L E Inside growth trees. I 25 17 16 18 19 2lt II 0.01 0.02 0.03 0.03 0.03 0.03 III - 5 . 7 1 - 2 . 8 6 0.00 0 .00 0.00 0.00 17 s. S. N.S. N.S. N.S. N.S. 7 L L E M E L Generation 2 . Margin growth trees. I 13 9 7 15 12 6 II 0.01 O.Oit O.Oit 0.10 0.10 0.12 III - 2 . l j 8 - 2 . 0 6 - 2 . 0 6 - 1 .2U -1 .2 i t - 0 . 9 7 17 s. N.S. N.S. N.S. N.S. N.S. 7 M L L L E E Inside growth trees. I 25 17 18 16 2it 21 II 0.00 0.01 0.02 0.02 0.03 0.03 III - 2 . 1 5 - 1 . 0 7 0.00 0.00 1.07 1.07 17 N.S. N.S. N.S. N.S. N.S. N.S. 7 L L M E L E Generation 1 + 2 . Margin growth trees. I 9 7 13 6 1 12 II 0.08 0.08 0.17 0.23 0 .25 0 .25 III - 2 . 2 1 - 2 . 2 1 - 1 . 5 3 -1.07 -0.£2 - 0 . 9 2 17 S. S. N.S. N.S. N.S. N.S. 7 L L M E M E Inside growth trees. I 25 17 16 18 2it 21 II 0.01 0.03 0 .05 0 .05 0.06 0.07 III -It.60 - 2 . 3 0 0.00 0 .00 1.15 2 .30 17 S. N.S. N.S. N.S. N.S. N.S. 7 L L E M L E Adelges cooleyi. The trees were treated as two population, 13 8 15 5 2 It lit 0.16 0.17 0.18 0.21 0.21 0.23 0.62 •0.52 -0.26 0.00 0.78 0.78 1.30 11.Ul N.S. N.S. N.S. N.S. N.S. N.S. H.S. M E L. E L E E 20 21 O.Oit 0.0U 2.86 2.86 S. S. M E 8 1 2 3 5 it lit 0.13 O.llt O.llt 0.17 0.22 0 .25 1 .03 •0.83 - 0 . 6 9 - 0 . 6 9 - 0 . 2 7 O.ltl 0.83 11.58 N.S. N.S. N.S. N.S. N.S. N.S. H.S. E M L L E E E 20 19 0.03 O.Oit 1.07 2 .15 N.S. N.S. M E 15 8 3 2 5 U U4 0.28 0 .30 0.32 0 .35 0.U3 0.U8 1 .65 •0.69 - 0 . 5 3 - 0 . 3 8 - 0 . 1 5 0.U6 0.8lt 9.77 N.S. N.S. N.S. N.S. N.S. N.S. H.S. L E L L E E E 20 19 Key: I, I I , III, 17 = as in tal 0.07 0.07 7,=.Bud opening time 2.30 2 .30 E = Early N.S. N.S. M = Medium M E L = Late i n the open. No significant difference was found between open and margin growth trees. Then the trees were separated into two groups, interior and edge trees, omitting the open trees and differences between trees were sought, differences were indicated by the " t " test (Table 1$). Slight differences can be seen between trees within the separated tree populations (Table 15). Most of the trees, which were significantly more populated than the average were early bud-opening trees, and most of those which were significantly less populated were late bud-opening trees. Slight differences were shown by " t n test between the late and early-opening trees (significant at 0.3 level) i n respect to the abundance of Adelges cooleyi. The test 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 relation to exposures and levels, 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 lev-els for 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 levels and exposures show significant differences (Table 17). The relationship between number of l i v i n g insects per needle and height level 38 Table 17 . Analysis of variance among exposures and levels. Source Degrees . of freedom Net sum squares Mean sum squares Variance ratio • (F) Signifi-cance at 0 .05 level 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 x E 6 0.0566 0.009k Total 11 1.0010 on the living crown wi l l be examined later. 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 Ranges Significantly different from North 0.69-1.17 S, E South 0.39-0.87 N, E West 0.5U-1.02 E East 1.86-2.3k N, S, W The 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 .0023 , a - 0 . 2 2 5 , r - 0.228, t = 0 . 1 1 2 , Sb = 0 .02013, - 0 .019 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 relatively small number of insects on i t . Another calculation 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 calculation shows significant correlation, meaning that the Adelges cooleyi usually i s more abundant on the longer-twigged trees. The factors D.B.H. (Xl), height (X2) , age (X3) , crown width (Xk), length of clear stem (X5), proportion of l i v i n g crown to total height (X6), bud-opening time (X7) and location (X8) of the trees were related to the average number of livi n g insects per needle (Y) by analysis of linear - multiple -regression. This was carried out by the Alwac III electronic computer (Appendix 0) (Table 19). Table 19. Influence of 8 factors on the average number of li v i n g insects per needle. Variable XI X2 X3 xu X5 X6 X7 X8 Means 6.5b U5.20 22.80 1U.68 1U.80 68.37 2.0k k.kk Standard deviations 3.39 20.01 5.80 6.60 12.16 19.99 0.9k 3.1k Minimum value 0.6 2 k 2 0 36.5 1 1 Maximum value 12.8 69 29 32 k l 99.9 9 • Correlation coefficients 0.131 0.3U3 0.269 0.125 O.U82 0.513 0.268 0.261 % of influence of each factor 5.73 28.31 0.58 -8.82 -U5.57 89.63 12.00 -19.27 R2 = 0.625 (R • multiple correlation coefficient) 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 correlation coefficients, only two factors show significant Ul correlation with number of insects: the length of clear stem and the percentage of l i v e crown. The meaning of these two significant results i s related to the location of trees. In other words the clear stems are longer on the interior trees, consequently these trees have smaller percentages of l i v i n g crown. 3. Intra Tree Variations: The analyses of variance i n table 2 and 3 did not show significant d i f -ferences among the cardinal points, meaning that no further analysis i s needed for this factor. Significant differences were found among the different levels of the trees (Table 2, 12 and 1 7 ) . The numbers of l i v i n g insects were summarized and averaged for the three specially 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 levels (Appendix G) (Fig. 11). The linear-regression equations were calcu-lated separately for the three sampling times (June 5, July 25, September 26) (Table 20, 21 and 22). Table 20. Calculation of the equation and correlation coefficient 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 2  o 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 1806.25 0.0009 1.275 U7.5 0.02 2256.25 0.000U 0.950 52.5 0.02 2756.25 0.000U 1.050 57.5 0.00 3306.25 0.0000 0.000 350.0 1.33 1U312.50 6.3181 31.175 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 r o 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 = -0.0075X+0.39 Table 21. Calculation of the equation and correlation coefficient of average number of l i v i n g insects per needle on height from the ground (data of July 25). *- 1 X 2 —y2 XY 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 Total Ave. 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 6 i 6 -0.0036 Equation: Y = -0.0068X+0.3U Each of the three analyses (Table 20, 21, 22) show; significant correla-tion 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 correlation coefficient of average number of l i v i n g insects per needle on height from the ground (data of September 26). X I X2 Y2 XY 12.5 0.18 156.25 0.032k 2.250 17.5 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 = as i n table 2 0 , I = as i n table 2 0 , b = - 0 . 0 0 k 5 , a = 0.23, r = -0.907, t - -6.05, Sb • 0.0007, -0.0061 * i £ -0.0029 Equation; Y » -O.OOk5X+0.23 lengths of twigs were summarized and averaged for two trees (No. h and 10) by 5 feet intervals (Appendix G). Then the average twig lengths were plotted by height levels from the ground (Fig. 1 2 ) . The plotted points show a curve rather than a straight l i n e . For further analysis twig length was transformed to logarithms (Fig. 12) (Table 23). The curvilinear correlation i s significant between the height levels and twig lengths (Table 23). The trees No. k, 10 and 25 were also examined horizontally at 3-feet intervals radially from the bole at the middle of the low level of the l i v i n g Fig-12 Relationship between twig length and height on the living crown I36r- L 1*6 Table 23 . Calculation of the equation and correlation coefficient of average twig lenth on height from the ground. X Y logY (logY)* X(logY) 1 2 . 5 20 .0 1.301 156.25 1.693 16.26 17,5" 20 .3 1.307 306 .25 1.708 22.87 2 2 . 5 19.1* 1.278 506.25 1.633 28 .76 27 .5 19.5 1.290 756.25 1.661* 35.1*8 3 2 . 5 20 .6 1 .3H* 1056.25 1.727 1*2.71 3 7 . 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 78.32 350.0 . 206.9 13.11*1 11*312.50 17.273 1*62.77 35 .0 20.7 1.311* X = height from the ground, Y = average twig length of the height lev 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 insect 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 (Fig. 1 3 ) , and the regression analyses were carried out separately for the three different sampling times (June 5 , July 25 , September 26) (Tables: 21*, 25 and 26). The correlations are not significant at the 0 .05 level i n tables 21* and 25, which might also be caused by the small sample sizes. Significant corre-lation was found i n table 26 between the numbers of l i v i n g insects per needle and distances from the stem. hi F i g 13 R e l a t i o n s h i p b e t w e e n n u m b e r o f l i v i n g i n s e c t s p e r n e e d l e a n d d i s t a n c e f r o m t h e s t e m a t t h e l o w l e v e l o f t h e l i v i n g c r o w n ° Distance from the Stem —Feet U8 Table 2k. Calculation of the equation and correlation coefficient of average numbers of l i v i n g insects per needle on distances from the stem (data of June 5). X Y o x-Y3  XY < 1.5 0.02 2.25 o.ooou 0.030 U.5 . 0.09 20.25 0.0027 o.i405 7.5 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 Total Ave. X - distances from the stem, Y = numbers of l i v i n g insects per needle, b = = 0 . 0 1 0 , 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 correlation coefficient of average numbers of l i v i n g insects per needle on distances from the stem (data of July 25). x1 y x 5 — 1 XY 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 7.5 0.09 Total Ave. X = as in table 2 U , Y = as in 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 . 0 2 5 Equation: Y = 0 . 0 1 2 X U9 Table 26. Calculation of the equation and correlation coefficient 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 0.0009 o.oh5 U.5 0.03 20 .25 0.0009 0 .135 7 . 5 o.m 56.25 0.0196 1.050 1 0 . 5 0.12 110 .25 0.011*1* 1.260 1 3 . 5 0 .25 182.25 0.0625 3 .375 3 7 . 5 0.57 371.25 0.0983 5 .865 7 . 5 0 .11 X = as .in table 21*, Y = as in 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 27 . Calculation of the equation and correlation coefficient of average lengths of twigs 'on distances from the stem. t Y X 2 1 ^ 1 XY 1.5 10.1* 2.25 108.16 15.60 I*.5 12.9 20.25 166.1*1 58.05 7.5 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 Total 37.5 81*.5 371.25 1537.87 732.1*5 Ave. 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 F i g - 1 4 R e l a t i o n s h i p b e t w e e n t w i g l e n g t h a n d d i s t a n c e f r o m t h e s t e m a t t h e l ow l e v e l o f t h e l i v i n g c r o w n 51 The twig lengths were also measured every 3 feet horizontally from the stem, at the middle of the low level of the l i v i n g crowns (Appendix G). The average twig lengths were plotted over distances from the stem (Fig. Hi) (Table 27). Significantly linear relationships were found between the twig lengths and distances from the stem. h. Mortality: As mentioned, the accurate numbers of dead insects could not be counted, because many of them f e l l off the needles, and a large proportion of them was eaten completely by predators. Assuming that the probability of loss of dead insects was similar everywhere, the analysis of data w i l l give some informa-tion about the factors influencing mortality. Trees No. 1, 2, 10, 11, 1U and 20 were examined by two analyses of variance, separately for generation 1, and generation 2 (Appendix M) (Tables 28 and 29). Table 28. Analysis of variance of dead insects among trees levels and expo-sures (generation 1). Source Degrees of freedom Net sum squares Mean sum squares Variance ratio (F) S i g n i f i -cance at 0.05 level Level (L) 5 0.19 0.095 23.75 H.S. Exposure (E) 3 0.02 0.007 1.75 N.S. Tree (T) 5 3.20 0.6L0 160.00 H.S. 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 Both the analyses of variance (Table 28 and 29) show significant-52 Table 29. Analysis of variance of dead insects among trees, levels and exposures (generation 2). Source Degrees of freedom Net sum squares Mean sum squares Variance ratio (F) ,S i g n i f i -:cance at 0.05 level Levels (L) 2 0.032 0.0160 26.67 ] H.S. Exposures (E) 3 0.005 0.0017 2.83 N.S. Trees (T) 5 0.130 0.0260 U3.33 H.S. L x E 6 0.002 0.0003 - N.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 difference within the levels and within the trees. No significant difference was found among the exposures. The significant difference i n the interactions indicates that the effects of the factors were not independent. The L x T interaction maybe significant because the actual heights of the sampling levels were different from tree to tree (Appendix E). These differences can be seen well between the marginal and interior trees (e.g. the height of low level of tree No. U was 19 feet, and of tree No. 25 was U5 feet). A l l the 25 sample trees were also examined by an analysis of variance, where the data of exposures were used as replications (Appendix N) (Table 30). A l l of the factors, trees, levels and generations, are significant, and a l l of the interactions are also significant. These results prompted another analysis, testing whether or not the main effects are significantly greater than the first-rorder interactions (Table 31). The effects of the main factors were significantly greater than the first-order interactions. The percentages of mortality were tested to determine whether or not the marginal trees with many insects had a higher percentage of insect mortality 53 (Table 32) (Appendix PJ. Table 30. Analysis of variance of dead insects among trees, levels! and generations. Source Degrees of freedom Net sum squares Mean sum squares Variance ratio (F) S i g n i f i -cance at 0.05 level 1 Compo-nents of variance Trees (T) 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 x L 1*8 0.09 0.0019 2.38 S. 0.00055 T x G 21* 0.1*0 0.0167 20.87 H.S.. 0.00530 L x G 2 0.01 0.0050 6.25 S. 0.00017 T x L x G 1*8 o.ol* 0.0008 0.00080 Total 11*9 1.72 Table 31. Variance ratio test between the main factors and first-order inter-actions . Source Degrees of freedom Net sum squares Mean sum squares Variance ratio (F) S i g n i f i -cance at 0.05 level Trees 21* 0.86 0.0358 5.26 H.S. T x L + T x G 72 0.1*9 . 0.0068 Levels 2 0.09 0.01*50 22.50 H.S. T x L + L x G 50 0.10 0.0020 Generations 1 0.23 0.2300 H*.65 H.S. T x G + L x G 26 0.1*1 0.0157 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 this was reversed i n population 2. The percentage of mortality by height on the li v i n g crown was also 5U calculated (Table 33). Table 32. Percentage of mortality by locality of the trees and generations. Average number of dead insects per needle Average number of living+dead insects per needle Percentage of mortality Trees marginal interior marginal interior marginal interior Generation 1 0.16 0.02 0.3U 0.05 1*7.1 Uo.o 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 Average number of dead insects per needle Average n living insects p umber of +dead er needle Percentage of mortality Generations 1 2 1 2 1 2 Low Medium High 0.15 0.12 0.07 0.05 o.ol* 0.02 0.35 0.27 0.11 0.25 0.18 0.06 U2.9 uu.u 63.6 20.0 22.2 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 will 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 will 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 level 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 level i n this experiment (from North, West, South and East). In an average of 6 trees (No. 1, 2, 10, 11, lk and 20) 0.35 was the number of li 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 level to determine the population of l i v i n g insects. The average number of dead insects per needle i n a twig was 0.17 with SD = 0.0129 and SEM = 0.00U3. The required sample size was calculated as 9 twigs per leve l . i i i . How many needles are required within a twig to determine the population of li v i n g and dead insects within $% of the mean nineteen times out of 20? This should be calculated for different levels, but here w i l l be shown only one example. The mean of the number of li v i n g insects at low level and North side i n generation 1 of tree No. 1, on June 1; was 0.1*0 with SD = 0.52 and SEM = 0.010. A total of 2,70k needles per twig were calculated as the required sample size. The average number of dead insects per needle was 0.10 at same place and time as before, with SD = 0.32 and SEM = 0.0025. Then 16,38k needles per twig were calculated as the required sample size. These sample sizes seem very large, but these numbers would decrease i f the limits of error were changed. For example, the required sample size of trees was 2,k6k to determine the population of livi n g insects within $% of the mean. Changing the limits of error to 10$, 616 trees would be required, and with 20$ error, only 15k trees would be needed. Considerable influence on the sample also results from changing the levels 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 level. Using analyses of variance to determine the influence of various factors may also reduce the need for such large numbers because factors themselves provide replication and partitioning of sum squares may present a very sensi-tive error term. 57 DISCUSSION As the former chapters show,during the summer of I960, two main groups of factors were examined: inter-tree differences and intra-tree differences. Two main groups of factors w i l l also be discussed here: the mortality and biotic potencial which were only partly investigated. 1. Inter Tree Differences: The power of multiplication of Adelges cooleyi i s affected by external and internal conditions of the tree. I t i s not always easy to determine whether external or internal conditions are responsible, but there i s consider-able evidence that some trees are more favourable to the multiplication of the insect than others. The inter-tree, differences can be divided into 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 chiefly on the trees near to the margin of the forest and the insect 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 in the open. It was found that by analysis of variance, (Table 12) the trees were populated differently by Adelges cooleyi. The Student's " t " distribution showed that generally the significantly more populated trees are growing at the margin of the forest, and the significantly less populated trees are growing at the center of the stand (Table 13). Then the trees were separated into 3 groups (edge-, inside- and open-growth) and the groups were tested against each other by " t " tests. These tests showed abundance to be significantly greater on the marginal and open trees than on the inside trees. No differences were found at the 0 . 0 5 level 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 is much smaller on an interior tree than on a marginal tree (Appendix L and 0). As shown by-several statistical analyses (see chapter: Experimental Results), the abundance of insects is 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 is very likely that the lack of the lower part of the living crown of the inside-growth trees is 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 is wind. The speed of wind is always close to zero within a dense stand (15, 27). The different abundance of the insect might be affected also by the different wind pattern, in 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 F i g 15 R e l a t i o n s h i p b e t w e e n t h e n u m b e r o f l i v i n g i n s e c t s p e r n e e d l e a n d h e i g h t o n t h e l i v i n g c r o w n ( A = i n t e r i o r g r o w t h t r e e , B = m a r g i n a l g r o w t h t r e e ) a> a> 0 4 r CP a. * 0-3 CD CO _c c? 0 2 o 6 a> > < 01 _L 0-5 5-10 10-15 15-20 20-25 25-30 30-35 35-40 40-45 45-50 50-55 5 Height from the Ground on Tree No 25 — Feet 60 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 6 0 are covered with a considerable amount of dust, which may be unfavorable to the insect. The other reason would be that only suppressed trees were avail-able for study i n the East margin of the forest. The high level of abundance on the East side might be caused by the low vigour and the relatively small height of the trees. The effects of prevailing winds - which are NW with a considerable amount of SE i n coastal B.C. ( 2 1 ) - should also be examined. But i n Totem Park Adelges cooleyi developed 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 dis-persal. Bud-Opening Time; Table 1 5 shows that the significantly more populated trees were mostly early-opening trees, most of the less-populated trees were late-opening. These differences were found both within the edge and interior trees. Only slight differences were shown (significant at 0 . 3 level) between early and late bud-opening trees. The reason for these differences might be looked for i n the seasonal development of Adelges cooleyi. The hatching of the progeny of sistens gener-ation was just about over when the early-opening trees started to swell. The progredientes and sexuparae settle more readily on the new foliage, but they could not settle on the new foliage of late-opening trees u n t i l 2 - 3 weeks after hatching. During this 2 - 3 weeks period the larvae were found on the surface of the opening buds, waiting for 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 different bud-opening trees? It i s true that 61 a l i t t l e displacement i s seen in the l i f e cycle on the late-opening trees, which (Appendix F) might be caused by the delay in food supply. I t i s more l i k e l y that there was not any synchronization between the l i f e cycle of the insect and the l i f e cycle of the late bud-opened trees, because the sexuparae migrate to Sitka spruce, and the Sitka spruce has not so wide range in bud swelling, as Douglas f i r ( 16 ) . For this very reason three l i f e cycles should be synchronized. Consequently the l i f e cycle of Adelges cooleyi seems to be determined by two factors, the growth cycle of Douglas f i r and the growth cycle of Sitka spruce, and being the primary host Sitka spruce^ on which the bisexual forms and reproduction occur, might be the more influential factor. Twig Length; Considerable variation was found between trees i n twig length at the end of the growing period. A slight correlation was found between length of twigs and the abundance of the insects (Fig. 1 0 ) . The stylet 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 longer-twigged trees have more lush growth, which i s more favorable to the insect. Adelges cooleyi would prefer- the lush growth, because the stylets would be more easily 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 nutritions for 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 total height, bud-opening time and place of the tree were also examined. These factors were analysed by linear - multiple - regression analysis (Table 1 9 ) . Two of the eight factors, the percentage of l i v i n g crown and the length of clear stem^showed significant correlation 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 a-bundance of the insect. The lack of significant result was caused by the fact that the trees were not separated as edge and inside trees in this analysis, but were numbered by different exposures (2- 8 ) and open-growth ( 9 ) and in-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 likely influenced by crown shape than cardinal point. 63 Height on Living Crown: Chrystal ( 7 ) has stated that the intensity of attack varies by height. This variation was also observed i n Totem Park. It was shown by analyses of variance, that low, medium and high levels of l i v i n g crown were significantly different i n abundance of the Adelges cooleyi (Table 2 , 12 and 1 7 ) . Also a significant correlation was found between the abundance of insects and height from the ground. The number of insects per needle decreased linearly with height (Table 2 0 , 2 1 and 2 2 ) (Fig. 1 1 ) . Teucher (26) has mentioned, that microclimate i s the main factor affect-ing the intensity 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 forest floor, and the relative humidity at the same time i s always lower above and i n the upper crown than above the forest floor. The average d i f f e r -ence i n temperature i s from 0.5°C to 1 .0°C, and i n R.H. i s 3 - h % ( 1 5 ) . These differences to a smaller degree can be observed foot by foot' from the forest floor 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 fact that the percentage of mortality was higher in the medium level than the low level, and i n the high level than the medium and low levels (Table 3 3 ) . The number of unhatched eggs was larger in the medium level than i n the low leve l , and more in the high level than i n the medium l e v e l . Percentages of unhatched eggs were: low level; 2 5 . 9 %> medium level: 3 3 . 8 % and high level: 3 7 . 0 %. Changes in Abundance of Insect in Horizontal Directions of the Tree Crown: I t was shown by regression analyses that slight correlation can be found between abundance of insect and the distances from the stem at the middle of the low level of l i v i n g crown (Table 2 U , 25 and 2 6 ) (Fig. 1 3 ) . The number of 6U insects per needle increased toward the periphery. I t may be supposed that this also resulted from microclimatic differences, or Adelges cooleyi looks i s less abundant on the shaded twigs. Considerable differences might be supposed in food supply between exposed and shaded twigs, since the needles are notice-ably thinner on heavily shaded branches. Twig Length; As was mentioned, Adelges cooleyi settles more abundantly on the needles of longer twigs. This result might be applied to prove the horizontal d i s t r i -bution of the insect within a tree, because as the lengths of the twigs i n -creased from the stem to the outer crown (Table 27) (Fig. LU), so increased the frequency of the insect. Examining the ve r t i c a l distribution of the insect and twig length, the result 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) (Fig. 12), and the numbers of insect per needle were decreasing linearly from the bottom to the top of the l i v i n g crown (Table 2 0 , 21 and 2 2 ) (Fig. 11). This result seems to show that the twig length, which i s one of the many factors affect-ing the v e r t i c a l abundance of Adelges cooleyi, was not so effective as others such as microclimatic factors. 3. Mortality; As mentioned, the number of dead insects could not be counted accurately. This unfortunate fact can be seen well i n Fig. 7 and 8 , where the mortality curves should follow the curves of l i v i n g insect but i n the reverse direction. I t can be seen from the diagrams that the loss of dead insects occurred mostly after the hatching time, when the mortality showed the highest potential. 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 settling down for overwintering. In tfe-is period the larvae were not settled i n a special place, 6 5 the mouth-parts were not inserted into the mesophyll tissue, and consequently they were more exposed to climatic and biotic effects. I t might be considered that the high mortality in generation 2 was caused by climatic factors. Examining the meteorological records (Appendix K) of July and August, the temperature was sometimes extremely high i n July, and precipitation low, with 0 . 0 3 inches rain i n the whole month. Following these the average temperature was very low in August. The average maximum was 5 8 . 3 2°F, and the average minimum was 5 5 . 3 8°F. The average maximum and minimum temperature for 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 affect the newly hatched larvae. The loss of dead insects i n this period can be explained by the fact that the larvae were not yet settled for overwintering. The loss of dead insects cannot be seen i n generation 1 , because one part of this generation was winged. The decrease of the number of l i v i n g insects per needle was linear (Fig. 5 and 6 ) and the increase of the number of dead insects per needle was also linear, but the slope of this line i s not proven, because of loss of dead insects. The two lines, the line of li v i n g insects and the line of mortality are not balanced, because of the loss of the winged part of the generation, and the loss of a part of the dead insects, and the proportion of these two were not or could not be observed. Examining the line of li 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 after 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 significantly different among trees, among levels and among generations. These results indicate that the absolute number of dead insects was higher in such a place where the number of l i v i n g 66 insects was higher. A question can be raised at this point, whether the degree of mortality was larger on the higher-populated trees or not. It 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 result 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 mortality was higher on the places where Adelges  cooleyi was less abundant. I t seems to be that the circumstances of the le s s -populated places were not suitable for 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 for 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 loss of winged foufis of the generation. The l i v i n g winged adults were lost, but the dead ones were counted. k. Natural Enemies; No special studies were conducted on how the natural enemies affect the abundance of Adelges cooleyi. 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 (10 , 11) were found, but they were not so common as the ladybird beetles. One species of Hymenopteran parasite was found, which developed i n the abdomen of both progredientes and sexuparae. They were not common, only two specimens being found during the summer of I 960 . A large number of Spiders and Red - Spider - Mites occurred around the 67 settled Adelges and their egg clusters, but their effect on the Adelges coo-l e y i was not certain. 5. Biotic Potential; Since reproduction i s by cyclical parthenogenesis, the potential rate of reproduction i s not limited by the necessity of mating ( 2 , 1 9 ) which i s be-lieved 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. The sampling was repeated 13 times and 12 sample twigs were drawn from every tree at each survey. 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 factors, e.g. temperature, RH, precipitation, evaporation, wind and total hours of sunshine,were also studied from available data, during the period April 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 results 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 relation 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 directions. 3. The insects were slightly less abundant on the late bud-opening trees, than on the early bud-opening, both on trees grown in the te 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 de-creased linearly from the bottom to the top of the l i v i n g crown, and decreased linearly 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 level). 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 1. Appendix B Table 35. The average number of li 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 level 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 level on the li v i n g crown. Appendix E Table 38. Heights of sampling levels of trees. Appendix F Table 39. Changes in development of Adelges cooleyi. Appendix G Table UO. Number of li v i n g insects per needle and average twig lengths change by height on the livi n g crown. Table h i . Number of livi n g insects per needle and average twig lengths change horizontally within the l i v i n g crown, at the middle of low level of the trees. Appendix H Table h2. The growth of foliage by level and time on trees No. Ik and 15. Appendix I Table k3. Calculation of the equation and correlation coefficient of average number of li v i n g insects per needle on average twig lengths. Appendix J Table kk. The average length of twigs by level and tree. Appendix K Tables k5, k6, k7, k8, k9 and 50. Climatological Station Report. Appendix L Crown shapes of three typical trees. Appendix M Table 51. Average l i v i n g and dead insects per needle, on trees No. 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 level 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, clear stem, % of l i v i n g crown, bud opening time and location of the trees. Appendix P Table 5U. Percentages of mortality by generation and tree. 72 Appendix A Table 3k. The average number of l i v i n g and dead insects per needle of generation 1. (The numbers are averaged of 5 sampling periods, from June k to July 2). Low Level Medium ~s F~ No. of High Tree No. N E N T 2 3 k 5 6 7 8 9 10 11 12 13 lk 15 16 17 18 19 20 21 22 23 2k 25 T572TT 0.U2 0.2k 0.3k 0.26 0.1k 0.06 0 .30 0.16 0.36 0.26 0.3k 0.82 0.12 0.06 0.0k 0.0k 0.08 0.0k 0.08 0.02 o.ok "OTIS" 0.k8 0.20 0.22 0.16 0.02 0.20 0.06 o.ko 0.2k 0.22 0.18 0.66 0.12 0.06 0.0k 0.0k 0.06 0.08 0.0k 0.28 0.1k 0.02 0.02 ~o7nr 0.50 0.3k 0.58 0 .k6 0.3k 0.12 0.k2 0.10 0.12 o.kk 0.k6 0.08 0 .00 0.02 0.0k 0 .00 0.02 0.22 0.0k 0.02 "OTEH o.ok E | N S living insects. W E 0.1k 0.06 0.16 0.36 0.60 0.28 0.78 0.26 0.0k 0.06 0.00 o.ok 0.06 o.ok o.ik 0.06 o.oo o.io 0.16 0.12 0.16 0.28 0.12 o.ok 0.22 0.08 0.06 0.06 0.10 1.0k o.ik 0.02 0 .00 0.00 0.00 o.ok 0.05 0.36 0.28 0.02 0 .00 0 .38 o . i k 0.08 0.22 0.12 0.00 0.18 0.02 0.06 0.0k 0.08 o.ik 0.86 0.16 0.02 0.00 0.00 0 .05 0.02 0 .00 0 .k6 0.12 0.00 0.00 0.02 o.ik 0.22 0.3k 0.32 0.12 0.02 0.2k 0.00 0.28 o.ok 0.02 0.58 0.02 0 .05 0 .05 0.00 o.ok 0.02 0.30 0.2k 0.02 0 .05 7J72TF 0.3k 0.10 0.02 0.08 0.16 o.ok 0.2k 0.12 0.12 0.22 0.76 o.ko 0.00 0 .00 0.10 0 .05 0.00 0.00 0 .30 0.16 0.02 0.00 "0702 -0.02 0.02 0 .05 0 .00 0 .00 0.00 0 .00 0.02 0.00 0.00 0.00 o.ok 0.38 0.06 0.00 0.00 0.00 0 .00 0.02 0.08 o.ik 0.12 top 0.00 "0730" 0.02 0.12 0 .05 0.06 0 .05 0.00 0 .05 0.00 0 .10 0 .00 0.00 0.00 0.32 0.0k 0 .00 0.00 0 .10 0.00 0.06 o.o5 0.12 0.08 broken 0.00 0 750"" .00 .00 .00 .02 .00 .00 .00 .02 .00 .05 .05 0.30 05 .00 .00 .00 .00 .00 .12 .08 dead "o7nr 0.18 o.ok insects, "0750" 0.02 0.00 0.00 0.00 0.00 0.00 0.02 0.02 0.05 0.00 0.00 0.10 0.k2 0.30 0.00 0 .05 0.00 0.00 0.00 0.00 0.18 0.12 .00 0 .00 "0720" 0.18 0.12 0.18 0.22 o.ik o.ik 0.10 0.08 0.20 0.18 0.16 0.7k o.ok 0.06 o.ok 0.06 0.08 o.ok 0.10 0.02 0.02 "0715" 0.26 0.20 0.3k 0.12 0.18 0.26 o.ok 0.20 0.30 o.ik 0.32 0.8k 0.06 0.10 o.ok o.ok 0.06 0.08 O.Ok 0.16 0.10 o.ok 0.00 "07W 0.20 0.26 0.20 0.26 0.10 0.3k 0.2k o.ok 0.30 0.22 0.58 O.Ok 0.00 0.02 o.ok 0.02 0.02 0.10 o.ok 0.00 "OTT? 0.08 "0778" 0.12 0.16 0.18 0.22 o.ok 0.22 0.16 0.10 0.06 0.06 0.12 0.6k 0.18 0.00 0.00 0.05 0.00 0.10 0.00 0.16 0.16 0.02 0.00 "0706" 0.16 0.06 0.06 o.ik 0.10 0.12 0.22 0.08 0.06 0.10 0.10 o.ok 0.k2 0.12 o.ok 0.00 0.05 0.00 0.02 0.00 0.26 0.12 0.02 0.05 No. of 073? 0.06 0.06 o.ok 0.00 0.00 0.02 0 .05 0 .05 0.00 0 .05 0 .00 0 .05 0 .10 0.72 0.1k 0 .05 0 .00 0 .05 0.00 O.Ok 0.00 0.08 0.08 top 0.05 "OToTT 0.02 0.02 0 .00 0.02 0 .00 0.00 0.12 0.02 0.00 0 .05 0 .00 0.10 0.56 0.10 0 .05 0.00 0.00 0 .00 o.ok 0.00 0.12 0.1k broken 0.10 0 75b~ .08 .02 .05 .02 .00 .00 .00 .00 .00 .05 .00 "0702" 0.02 0.02 0.05 o.ok 0.05 0.00 0.00 o.ok 0.00 0 .05 0 .05 0.06 0.68 0.18 0.00 0.00 0.00 0.00 0.02 0.00 0.12 0.10 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 o.ik 0.00 0.22 0.22 0.16 0.20 0.72 0.20 0.02 0.00 0.02 0.06 0.08 0 .00 0.06 0.02 0.02 0.12 0.08 0.16 0.28 0.10 0.12 0.18 0.00 o.ko 0.12 0.02 0.80 0.02 0.00 0.00 0 .20 0.02 0 .05 0 .10 0 .10 0 .00 0 .00 0.10 0.06 0.08 0 .00 0.12 0.06 0.08 0.16 0.66 0.38 0.02 0 .00 0 .00 0 .05 o.ok 0.00 0.20 0.08 0.02 0 .05 0.66 .00 05 .00 .00 .02 .00 .16 .lk .00 0.00 73 Appendix B Table 35. The average number of l i v i n g and of generation 2. (The numbers are averaged from July 9 to September 21*). dead insects per needle of 8 sampling period, Level Low Medium High Tree No. N 0.26 0.29 0.16 0.10 0.05 0.20 0.06 0.05 0.15 0.11* 0.00 1.75 0.11 o.ou 0.00 0.01 0.06 0.06 0.03 0.25 0.09 0.03 0.01 W E j N S ST E~ No. of l i v i n g insects. N W 1^ 0.20 "oTiT o.io 0.19 0.2U o.ui 0.2U 0.05 0.16 0.03 o.ou 0.00 0.03 0.76 o.ou 0.00 0.00 0.00 0.03 0.00 0.31 0.20 0.01 0.00 E 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 0 3 3 0.39 0.18 O.UU 0.36 0.16 0.05 0.25 0.06 0.23 0.15 0.01 1.51 0.05 0.03 o.ou 0.05 0.06 o.ou 0.15 o.ou 0.03 0715 0.31 0.59 0.60 0.61 0.28 O.lU 0.53 0.09 0.10 0.23 0.7U 0.05 0.01 0.03 0.05 0.06 0.03 0.20 o.ou 0.01 o.iU 0.05 0.11 0.25 0.50 0.05 1.76 0.2U 0.03 0.01 0.01 o.ou o.ou 0.05 0.19 0.05 0.00 0723 0.18 0.10 0.13 0.15 0.09 o.ou 0.05 0.03 0.06 o.ou 0.01 1.5U 0.08 0.01 0.00 0.00 0.00 0.03 0.00 0.3U 0.25 0.00 0.00 0713 0.10 0.16 0.21 o. i5 0.10 0.01 0.13 o.ou 0.01 0.03 o.ou 0.00 1.11 0.13 o.ou 0.00 0.00 0.00 0.01 0.00 0.29 0.26 0.03 0.00 0715 0.11 0.11 0.08 0.06 0.01 0.00 0.03 o.ou 0.06 o.ou 1.78 0.13 0.00 0.10 0.00 0.10 0.03 0.00 0.23 0.29 0.00 0.00 dead 0.10 0.06 0.05 0.08 0.06 0.03 0.00 o.o5 0.01 0.03 0.00 0.01 0.01 0.13 0.00 0.01 0.00 0.10 0.00 0.01 0.00 0.08 0.10 0.00 0.00 insects ~ o n r 0.09 o.ou 0.11 0.10 0.10 0.03 0.12 0.00 0.03 0.00 0.01 0.11 0.00 0.00 0.00 0.00 0.01 0.00 0.05 0.10 0.01 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 top 0.00 0 o.ou 0.01 o.ou 0.00 0.00 0.20 0.00 0.10 o.ou 0.00 0.00 0.00 0.16 0.00 0.00 0.00 0.00 0.01 0.00 0.13 0.10 .01 .01 .01 00 00 .00 .00 .00 .01 00 .10 .00 .00 .U3 .09 .00 .00 .00 .20 .01 .00 .13 .10 broken .00 0.00 0700 0.00 0.00 0.10 0.20 0.00 0.00 0.00 0.03 0.00 0.00 0.00 0.10 o.Ui 0.08 0.00 0.00 0.00 0.00 0.00 0.00 0.13 O.IU 0.00 No. of "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 "OTOT 0.06 0.11 0.13 0.16 0.08 0.05 0.10 0.01 o.ou 0.05 0.10 0.03 0.00 0.00 0.03 o.ou 0.03 0.08 0.03 0.00 "07IF 0.10 0.05 0.08 0.05 0.08 0.03 o.ou 0.03 0.01 O.OU 0.03 O.IU 0.08 0.00 0.10 0.00 o.ou o.oU 0.00 0.08 0.08 0.01 0.00 UToT 0.09 o.ou "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 top 0.00 0. 0.00 0.03 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.08 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 o.ou 0.00 0.00 0.00 0.00 0.03 0.00 0.05 0.08 0.00 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 0709 0.09 o.ou 0.01 0.03 o.ou 0.09 0.01 0.16 0.00 0.00 0.00 0.01 0.03 0.10 0.00 o.ou 0.01 0.01 0.05 0.03 0.00 0.00 0.00 0.01 0.05 o.ou 0.23 0.11 0.01 0.00 0.00 0.00 0.01 0.00 0.05 0.11 0.00 0.00 0. 0. 0. 0. 0. 0. 0. 0. 0. 0. 0. 0. 0. 0. 0. 0. 0. 0. 0. 0. 0. 0. 0. w 01 00 00 00 00 00 00 00 00 00 00 00 11 09 00 10 00 00 00 00 06 08 0.00 0.00 0.00 0.00 0.01 0.00 0.05 0.05 broken 00 0.00 Appendix C Table 36. Average numbers of livi n g insects per needle by period and tree at the low l e v e l on the li v i n g crown. Tree No. 1 2 3 u 5 6 7 8 9 10 11 12 13 iu 15 16 17 18 19 20 21 22 23 2U 25 Ave. Generation 1 Sampling 0 1 periods F 2 0.2« 0.53 0.50 0.50 o.uo 0.20 0.07 o.U5 o.i3 0.38 0.U5 0.70 0.U3 0.95 0.03 0.10 0.08 0.03 0.05 0.10 o.o5 0.27 o. i5 0.03 0.03 "0T2T "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 (week) 1 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 0.23 ~G~TT 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 u ~ "0718" 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 ~G~W 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 0.60 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 Generation 2  Sampling periods (week"T 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 0725 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 W 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 TOO" 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 0.21 0.26 0.30 0.23 0.1B 0.17 9 TJ7I6-0.18 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 " O J 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 low level on the living crown.  Tree No. 1 2 3 U 5 6 7 8 9 10 11 12 13 Hi 15 16 17 18 19 20 21 22 23 2ii 25 Ave. Generation 1 Sampling periods (week)-0 "oToT 0.18 0.23 0.15 0.23 0.13 0.23 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 0.10 0.18 0.10 0.15 0.23 0.10 0.27 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 0.15 0.15 0.15 0.12 0.15 0.20 0.20 0.33 0.17 0.20 0.18 0.03 0.23 0.23 0.20 0.23 0.70 0.23 0.03 0.08 0.03 0.03 10 05 03 10 0,08 0.00 0715" 0.15 0.20 0.25 0.23 0.13 0.10 0.25 0.00 0.18 0.23 0.07 0.30 0.78 0.10 0.10 05 00 08 10 05 13 05 03 0.03 oTiF 008" 0.25 0.23 0.20 0.33 0.07 0.30 0.13 0.00 0.20 0.15 0.13 0.33 80 10 03 ,00 05 08 10 00 20 15 00 0 0 0 0 0 0 0 0 0 0 0 0.00 ~o7ic" Generation 2 0 0.0b 0.05 0.03 0.00 0.00 0.07 0.00 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 . O H i 1 ~OJ5F 0.05 0.07 0.00 0.03 0.07 0.00 0.05 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 Sampling periods 2 i u — (week) 0.019 "07lB~ 0.03 0.13 0.10 0.07 0.10 0.03 0.03 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 0.05 0.10 0.10 0.10 0.10 0.07 0.03 0.10 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 o.ouu 0.10 0.00 0.07 0.10 0.03 0.07 0.00 0.08 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 0.0U8 "7J30 -0.08 0.10 0.10 0.13 0.03 0.03 0.18 0.03 0.05 0.08 0.10 0.03 0.18 0.10 0.10 0.00 0.00 0.05 0.00 0.00 0.03 0.15 0.03 0.00 0.067 11 0.15 0.13 0.17 0.25 0.13 0.17 0.03 0.08 0.00 0.05 0.08 0.10 0.00 0.30 0.03 0.00 0.00 0.00 0.05 0.03 0.03 0.03 0.00 0.03 0.00 0.07U 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 TJ^7l -~0 Appendix E Table 38. Heights of sampling levels of trees. Tree Height of levels Levels and directions No. in feet where could not be found Low Medium High l i v i n g branches 1 21 27 33 2 16 26 35 3 15 27 39 LE h 19 36 53 LE, : LS, ME 5 17 36 55 LE, ME 6 21 39 57 LE 7 16 33 50 LE 8 20 36 52 9 10 16 22 LN, MN 10 19 37 55 11 17 32 47 12 16 31 46 LW 13 8 12 16 LW, MW, HW Hi 5 11 17 15 6 10 lit LW, MW, HW 16 25 39 5u LN 17 h3 53 63 18 36 42 U8 19 39 h3 47 20 21 30 39 21 Ul 53 65 22 0.6 1.3 2 LN 23 2 h 6 LW, LE 2U 19 27 — 25 U5 57 61 Key: L = Low leve l , M = Medium leve l , H = High level W = West, N = North, E = East, S = South 77 Appendix F Table 39. Changes in development of Adelges cooleyi. Date Tree June June June June July July July Fall • No. U 11 18 25 2 9 16 Winter i FSN2 PSN3+A+E A+E A+E+H A+H H H H 2 PSN1+2 PSN3+A A+E A+E A+E+H E+H H H 3 PSN1 PSN2+3 PSN3+A+E A+E A+E E+A+H E+H H 1* PSN3 PSN3+A+E A+E A+E A+E+H E+H H H 5 PSN2 PSN3+A+E A+E A+E A+E+H E+H H H 6 PSN1+2 PSN3+A A+E A+E A+E+H E+A+H H H 7 PSN1 PSN2+3 PSN3+A+E A+E A+E+H A+E+H E+H H 8 PSN1+2 PSN3+A+E A+E A+E A+E+H E+H H H 9 PSN1+2 PSN3+A A+E A+E A+E+H E+H H H 10 PSN1+2 PSN3+A A+E A+E A+E+H E+H H H 11 PSN3" PSN3+A+E A+E A+E A+E+H E+H H H 12 PSN2+3 PSN3+A+E A+E A+E A+E+H E+H E+H H 13 PSN2+3 PSN3+A+E A+E A+E A+E+H A+E+H H H LU PSN2+3 PSN3+A+E A+E A+E A+E+H E+H H H 15 PSN1+2 PSN3+A A+E A+E A+E+H E+H E+H H 16 PSN2 PSN3+A PSN3+A+E A+E A+E A+E+H E+H H 17 PSN1+2 PSN2+3+A A+E A+E A+E+H H H H 18 PSN1 PSN2+3 PSN3+A+E A+E A+E A+E+H E+H H 19 PSN2+3 PSN3+A+E A+E A+E+H A+E+H A+H+E H H 20 PSN2+3 PSN3+A+E A+E A+E A+E+H E+H H H 21 PSN2+3 PSN3+A+E A+E A+E+H E+H E+H H H 22 PSN2+3 PSN3+A+E A+E A+E A+E+H E+H H H 23 PSN1+2 PSN3+2 PSN3+A+E A+E A+E A+E+H E+H H 2U PSN1 PSN2+3 PSN3+A+E A+E A+E E+H H H 25 PSN1+2 PSN3+A A+E A+E E+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 living 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 Height levels i n feet 0-5 5-10 10-15 15-20 20-25 25-30 30-35 35-kO U0-U5 k5-50 50-55 55-60 Number of insects per needle June 5 July 25 Sept. 26 0.22 0.37 0.27 0.19 0.10 0.11 0 .03 0.02 0.02 0 . 0 0 0.32 0.27 O.lo 0.19 0 .03 0 . 0 1 0.0k 0.03 0 .00 0 . 0 0 0.18 0 .20 0.10 0.11 0.06 0 . 0 0 0.02 0.02 0.01 0 . 0 0 Average twig length i n cm. Sept. 26 - 2 0 . 0 20.3 19 .U 19.5 20.6 19.9 20.9 21.1 22.2 2 3 . 0 Table k l . Number of livi n g insects per needle and average twig lengths change horizontally within the l i v i n g crown, at the middle of low level of the trees. (Averaged date, of tree No. k, 10 and 25). Date Distances from the stem i n feet 0-3 3-6 6-9 9-12 12-15 15-18 Number of insects per needle June 5 July 25 Sept. 26 0.02 0.09 0.16 0.2U 0.10 0.01 0.03 0.1k 0.10 0.15 0.03 0.03 0.1k 0.12 0.25 - Average twig length i n cm. Sept. 26 lO.k 12.9 17.5 20.8 22.9 Appendix H Table U2. The growth of foliage by level and time on trees No. LU and 15-Tree No. Level Date June June June June July July July July July Aug. Aug. Sept. Sept. U 11 18 25 2 9 16 23 30 6 20 3 2U Twig lengths i n cm. LU Low Med. High 3.7 5.6 7.0 7.U 7.9 8.0 8.1 8.3 8.3 8.U 8.5 8.6 8.6 U.6 5-5 6.8 7.6 8.5 9.6 9-7 9.8 9-8 10.0 1 0 .l 1 0 .l 10.1 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 15 Low Med. High 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 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 3.1 U.U 5.8 6.6 8.0 9.2 10.7 11.2 11.3 l l . U 11.6 11.7 11.8 -o 80 Appendix I Table k3. Calculation of the equation and correlation coefficient of average number of l i v i n g insects per needle on average twig lengths. Tree No. X Y 52 52 XY 1 b.k <U2 70.56 0.176k 3.$2B 2 6.0 0.65 36.00 0.k225 3.900 3 6.k 0.61 k0.96 0.3721 3.90k k 15.9 0.98 252.81 0.960k 15.582 5 8.7 0.69 75.69 0.k76l 6.003 6 7.7 0.39 59.29 0.1521 3.003 7 6.3 0.15 39.69 0.0225 0.9k5 8 9.8 0.55 96.0k 0.3025 5.390 9 9.3 0.1k 86 .U9 0.0196 1.302 10 23.6 0.29 556.96 0.08kl 6.8kk 11 13.6 0.59 l8k.96 0.3k8l 8.02k 12 7.3 0.62 53.29 0.38kk k.526 13 6.8 0.29 k6.2k 0.08kl 1.972 m 7.2 2.13 51.8k k.5369 15.336 15 6.1 0.30 37.21 0.0900 1.830 16 k.l 0.10 22.09 0.1000 0.U70 17 a.2 0.05 17.6k 0.0025 0.210 18 3.8 0.05 lk.kk 0.0025 0.190 19 6.0 0.10 36.00 0.0100 0.600 20 5.8 0.12 33.6k o.oikk 0.696 21 5.1 0.07 26.01 0.00k9 0.357 22 2.9 0.k2 8.kl 0.176k 1.218 23 8.7 0.23 75.69 0.0529 2.001 2k 3.1 0.08 9.61 0.006k 0.2k8 25 5.1* 0.03 29.16 0.0009 0.162 Total 192.8 10.05 1960.72 8.7127 88.2kl Ave. 7.71 0.k02 X = average twig length at the low level 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 calculation without trees No. 10 and lk: b - 0.058, a - -0.08, r - 0.680, t - k .25, 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 level and tree. Tree No. June U September 26 Level Low Medium High Low Medium High 1 3.0 3.9 6.8 8.U 8.6 18.6 2 0.6 0.7 1.0 6.0 6.1 11.3 3 o.U 0.U 0.7 6.U 7.6 10.8 U 3.5 5.2 7.1 15.9 15 .0 20.1 5 2.1 3.3 U.5 8.7 9.6 11.7 6 U.5 5.6 6.2 7.7 9.3 21.9 7 0 .0 0.0 0 .0 6.3 12.U 18.7 8 3 . 5 3.5 3.6 9.8 10.6 11.1 9 2 . 5 3.1 3.1 9.3 8.9 8.5 10 3.7 4 .0 5.1 23.6 2U.7 26.6 11 6.1 6.7 7.5 13.6 17.2 19.9 12 3.4 3.7 U.9 7.3 9.1 10.6 13 3.2 U.O 3.8 6.8 6.5 6.1 Hi 5.0 5.6 6.3 7.2 8.0 13 .U 15 2 .1 2.4 2.8 6.1 8.3 10.7 16 1.8 2.3 3.U U.7 7.0 13.3 17 0 .0 0 .0 0 .0 U.2 6.7 12.7 18 1.0 1 .1 l.U 3.8 5.3 9.8 19 2.1 3.0 3.7 6.0 6.2 9.U 20 1.2 1.9 2.8 5.8 6.8 10.2 21 2.0 2.7 3.6 5.1 5.2 6.9 22 1.7 2 . 5 3.0 2.9 U.3 6.8 23 1.0 1.3 1.7 8.7 12.7 17.7 2U 0 .0 0 .0 0 .0 3.1 3.5 -25 2 .1 2.6 3.2 5.U 6.6 io.U 82 Appendix K Table U5. Climate-logical Station Report (April I960, Vancouver U.B.C.). Date Air tempera- RH Rain Evapo- Wind Sunshine ture F inches ration mileage total past 21+ hours Max. Min. AM PM hours 1 1*6.5 3H.9 93 7U O.ll* 0.100 0.2 2 1*9.7 1*3.8 96 96 0.19 0.021 3 53.7 1*8.8 90 70 0.10 0.022 8.1 U 59.7 1*5.9 85 80 0.106 U.5 5 59.8 1*1*.9 93 85 0.090 6.3 6 50.1* U2.0 86 75 0.090 9.U 7 55.6 1*6.8 68 59 0.128 9.1* 8 61*.1* 50.1* 79 87 0.177 0.9 9 57.8 1*0.2 79 56 o.iu 0.01*3 9.8 10 51.7 36.1* 71* 65 0.110 8.9 11 51.8 l * l * . l 98 66 0.07 0.115 6.8 12 53.6 1*2.0 90 70 o.ol* 0.083 2.8 13 52.3 1*0.3 95 77 0.13 0.071 11* 1*8.0 1*1.2 77 70 0.11* 0.06U U.U 15 U8.3 37.5 85 78 0.17 0.106 9.2 16 50.3 37.3 82 60 0.137 U.U 17 1*9.5 1*2.1* 90 1*8 0.07 0.081* 9.5 18 53.0 1*1.3 88 93 0.08 0.09U 19 1*6.6 1*0.9 81* 86 0.1*1 0.050 1.5 20 1*9.2 U2.5 79 77 0.28 0.028 6.0 21 1*9.1* 36.7 87 73 0.090 10.2 22 U7.U 38.5 85 53 0.115 7.U 23 52.8 1*2.8 71 56 0.078 0.5 21* 58.0 1*3.2 88 61* 0.06 0.036 0.9 25 57.3 1*2.0 86 61 0.01 0.120 5.6 26 57.5 1*5.8 89 55 0.01 0.0U6 10.1 27 60.1* 1*3.3 77 1*9 0.187 13.1 28 63.0 1*7.9 73 52 0.231* 13.3 29 62.2 1*8.6 83 78 0.177 10.7 30 59.0 U6.5 88 71* 0.135 U.9 J l Sum 1618.9 1282.9 2538 2087 2.01* 2.81*7 178.8 Mean 53.96 1*2.76 81*. 60 69.57 0.068 0.095 5.96 83 Appendix K Table 1*6. Climatological Station Report (May i960, Vancouver U.B.C.). Date Air tempera- RH Rain Evapo- Wind Sunshine ture F inches ration mileage total past 21* hours Max. Min. AM PM hours 1 57.8 47.4 8i 73 0.090 120 0.8 2 57.7 47.3 94 89 0.086 100 3 53.5 U5.3 94 86 0.090 0.055 97 0.1 I* 51.0 1*5.8 94 87 0.032 0.017 50 5 54.1 1*8.0 89 60 0.01*8 120 6.8 6 59-3 51.4 98 96 0.370 0.108 100 0.3 7 59.3 1*6.6 86 63 0.380 0.032 89 9.4 8 56.1* 1*2.2 78 65 0.163 68 6.2 9 5U.9 47.4 61* 62 0.188 67 6.1 10 71.7 53.2 93 94 0.080 0.130 66 0.9 11 57.9 50.1 92 85 0.1*20 0.021* 69 12 55.1 50.6 91* 55 0.1*1*3 0.023 59 7.4 13 60.2 45.0 79 57 0.177 80 10.0 Hi 59.3 U8.2 72 57 0.181* 82.5 10.1 15 57.6 1*3.8 79 62 0.158 69.5 5.6 16 56.9 1*5.8 98 74 0.215 0.090 59.5 2.1* 17 54.1 42.3 95 76 0.160 0.080 86 6.6 18 53.3 1*3.2 90 57 0.070 0.057 77 6.5 19 58.3 46.5 7U 96 0.152 76.5 20 1*9.0 li5.8 74 62 0.750 110 10.6 21 53.3 1*1.2 89 65 0.205 0.133 92 9.1 22 53 .4 1*2.0 87 58 0.017 0.11*1 105 9.7 23 55.5 1*0.3 71 60 0.178 74 12.0 2li 58.8 U7.5 68 66 0.220 112.5 10.0 25 60.1 49.5 67 60 0.180 106 3.7 26 61.7 48.5 87 75 0.270 0.068 123.5 1.2 27 57.6 U6.3 90 72 0.01*5 0.029 75.5 1.5 28 55.9 49.0 78 62 0.081* 97 3.4 29 60 .U 1*8.6 79 82 O.lli* 70.5 30 56.3 50.8 94 82 0.165 0.011 108 0.9 31 60.6 1*9.0 81 69 0.050 0.051 11*2 7.1 Sum 1770.7 11*1*8.6 2609 2207 3.762 3.071 2752.0 11*8.1* Mean 57.12 1*6.73 81*. 2 71.2 0.1213 0.0991 88.8 4.79 81* Appendix K Table 1*7. Climatological Station Report (June I960, Vancouver U.B.C.). Date Air tempera- RH Rain Evapo- Wind Sunshine ture F inches ration mileage total Max. Min. AM PM past 2l* hours hours 1 62.2 1*9-8 81* 72 0.182 101 3 .7 2 65.8 52.2 86 71* 0.116 1 0 2 . 5 2.1* 3 6 8 . 5 5U.0 91 70 0.121 79 11.5 1* 6 1 . 3 2*5.2 67 72 0.208 128 5.6 5 6 3 . 3 53.8 68 62 o.U*5 6 7 . 5 7.1 6 66.7 5 0 . 8 61* 58 0.218 111 l l * .1* 7 5 9 . 8 51.0 71* 70 0.269 11*0 11* .1* 8 5 8 . 5 U 5 . 5 82 65 0.186 78 13.1* 9 6 3 . 0 1*9.8 68 57 0.238 60 11*.2 10 6 6 . 6 ' 5 3 . 5 67 77 0.267 75 3 . 6 11 65.7 5 2 . 5 89 72 0.133 83 10.3 12 6 5 . 0 51.6 83 72 O.I89 61*.5 9 . 8 13 69-7 55 .5 70 70 0.230 8 3 . 5 1.1* U* 6 3 . 9 52.1* 81 96 0.123 96 0 . 3 15 59.7 50.1* 91* 96 0.170 0.055 55-5 0.1 16 5 8 . 1 51.1* 69 61 0.170 135 11-9 17 6 2 . 6 1*9.9 75 57 0.271 81*.5 1 0 . 1 18 6 3 . 3 1*8.8 70 57 0.186 101* 3 . 3 19 62.5 1*7.9 77 75 0.118 73 9 . 8 20 59.3 1*7.2 92 68 0.01*1 0.209 9 6 . 5 1 .0 21 6 0 . 7 1*9.1 85 77 0.103 88 3 . 0 22 5 9 . 8 U8.I4 79 71 0.110 56 l i * . 3 23 6 6 . 6 53.6 72 75 O.198 68 12.6 21* 72.6 57.1 82 78 0.21*7 8 8 . 5 U.l 25 6 7 . 0 51*.3 81* 79 0.017 0.109 9 5 . 5 26 6 3 . 1* 53.8 83 83 0.093 1*3 27 6 1 . 1 55-9 82 70 0.075 1*6.5 2.2 28 6 7 . 3 5 2 . 5 71* 56 0.156 1*1* 11*.3 29 6 9 . 0 51*.2 75 59 0.222 7 2 . 5 12 .6 30 71 .9 55 .5 81 79 0.261 101* 0 . 6 J>X Sum 1925.8 151*7.6 231*8 2128 0.398 5.038 2523.5 212.0 Mean 6U.19 51.59 78.3 7 0 . 0 0.0132 0.1679 81*. 12 7 .0 Appendix K Table L8. Climatological Station Report (July I960, Vancouver U.B.C.). Date Air tempera- RH Rain Evapo- Wind Sunshine ture F inches ration mileage total past 24 hours Max. Min. AM PM hours 1 63.3 52.8 85 96 0.122 105.5 1.1 2 61.7 51.7 78 68 0.030 0.077 109.5 13.8 3 61.6 52.3 80 68 0.178 65 14.6 4 68. U 54.6 87 73 0.2U6 74.5 14.1 5 68.6 60.3 87 56 0.254 117 14.4 6 75.9 61.5 65 44 0.310 99 LU.5 7 81.5 57.4 78 6U 0.282 58 lit.2 8 72.1 49.9 76 60 0.302 158.5 14.5 9 65.8 55.0 95 65 0.243 141 14.4 10 67.9 50.3 78 69 0.285 78 8.7 11 66.2 54.1 71 58 0.166 86 7.0 12 72.8 56.8 70 59 0.200 32 11.3 13 82.2 58.9 79 65 0.266 84 9.5 lit 71.5 57.6 65 56 0.263 85 12.0 15 70.8 56.4 66 59 0.265 75 10.2 16 77.1 60.3 63 49 0.182 70 12.6 17 81.4 60.3 69 73 0.340 85 14.3 18 81.4 59.5 84 70 0.291 60 14.1 19 73.8 59. h 81 70 0.289 140 14.1 20 68.6 57.4 83 68 0.290 130 Hi.O 21 69.8 55.3 81 64 0.2ii5 85 13.9 22 72.0 52.0 78 67 0.233 95 11.9 23 68.2 5l.o 80 58 0.234 78 13.9 24 67.4 52.0 77 64 0.253 82 lu.l 25 67.5 56.8 87 75 0.191 75.5 1U.1 26 71.8 59. u 71 55 0.313 105 12.U 27 77.0 63.8 66 50 0.259 90.5 8.U 28 77.2 60.6 84 47 0.2Hi 49.5 13.8 29 78.0 62.8 68 47 0.302 61.5 13.4 30 87.4 65.6 64 50 0.285 67 8.1 31 76.5 58.3 83 71 0.342 140 10.8 Sura 221*5.4 176U.1 2379 1938 0.030 7.722 2782.0 378.2 Mean 72.43 56.91 76.7 62.5 0.2491 89.74 12.2 8 6 Appendix K Table I49. Climate-logical Station Report (August I 9 6 0 , Vancouver U.B.C.). Date Air tempera- RH Rain Evapo- Wind Sunshine ture F inches ration mileage total Max. Min. AM PM past 2a hours hours 1 59.2 56.9 87 72 0 . 1 8 3 98.5 0 . 8 2 60.6 58.7 89 78 o.ia6 85 2.5 3 59-9 58.2 90 68 0.118 56.5 a.2 a 58.9 55.0 79 63 0.175 96 a.a 5 61.0 57.2 79 69 0.116 61 13.6 6 61.1 58 .a 85 71 0.188 91 13.3 7 63.3 60.6 85 39 0.281 109 13.7 8 7a.0 58.8 ao 2 7 0.305 108.5 13 . 6 9 7a.5 60.2 a3 21 0.3a2 6 1.0 13.5 10 61.a 58.0 8 3 6 7 0.3a9 97 13.0 11 61.3 57.1 77 63 0.238 H»3 12.9 12 59.8 56.7 sa 73 0.22a 73 8.5 1 3 57.6 5a.5 8 3 51 0.229 8 8.5 8.5 ia 53.2 62.a 9a 89 0.005 o.2a5 95 0.2 15 56.0 52.2 78 73 o.oa3 0.031 73 8.1 16 55.0 5a.7 98 93 0.163 0.122 9 6.5 17 59.3 58.0 9 3 8 2 0.230 0.013 120 18 61.9 6 0 . 0 89 69 0.077 75.5 6.9 19 58.8 57.1 9 0 6a 0.172 72.5 9.2 20 61.7 59.9 89 56 0.165 68.5 8.5 21 50.2 a9.5 96 9a 0.220 0.110 99 i.a 22 52.0 51.0 9a 65 0.310 0 . 1 0 8 110.5 a.3 23 53.8 51 . 9 8 9 91 0 . 1 0 8 8 8.5 o.a 2a 53.5 53.0 96 72 0.360 0.006 62 5.0 25 5k.8 53.3 91 91 0.060 0 . 1 2 6 101 2 6 5 a .0 53.2 9a 7 1 0.a35 0.001 83 2.2 27 55.3 53.2 87 60 0.087 83 6.0 28 53.5 53.0 96 9 8 0.015 0.176 105.5 29 5a.2 5o.a 7 7 70 0.380 0.075 129 30 5a.0 52.9 93 98 0.155 o.oa9 103.5 31 5a.1 50.9 80 56 o.6ao 0.018 103 11.0 Sum 1807.9 1716.9 2 6 2 8 215a 3.013 a. 583 2837.0 1 8 2 . 7 Mean 58.32 55.38 8a.8 69.5 0.0972 0.1LJ78 91.52 5.9 87 Appendix K Table $0. Climate-logical Station Report (September I960, Vancouver U.B.C.). Date Air tempera- RH Rain Evapo- Wind Sunshine ture F inches ration mileage tot a l past 2b hours Max. Min. AM PM hours 1 50.3 1*8.0 81* 71 0.11*0 52 11.1 2 53.8 50.5 80 77 0.11*8 7b.5 11.6 3 55.8 51.0 73 77 0.127 68.5 9.3 U 56.0 52.3 79 91 0.100 0.11*6 112.5 1.1 5 51*.2 51.0 80 62 0.135 0.075 96 10.3 6 52.8 1*9.7 82 68 0.126 62.5 9.6 7 53.1 1*9.3 76 83 0.153 93.5 11.b 8 55.3 53.5 86 68 0.176 93.5 l l . l 9 58.9 52.0 63 1*3 0.176 83.5 11.1 10 59.1 52.7 66 b9 0.153 53.5 10.9 11 61.1 51*.7 67 39 0.161 b6 10.7 12 62.1 57-5 75 86 0.11*6 52.5 2.0 13 56.1 55.0 93 80 0.01*9 68 9-1 H* 55.7 55.1 96 93 0.093 b2.5 1.2 15 51*.5 5U.3 98 79 0.070 37 7.8 16 56.7 56.1 96 81 0.087 59 7.1 17 51.8 51.8 100 82 0.123 56 18 5b.7 53.5 93 91 0.010 0.036 99.5 19 55.5 55.2 98 61* 0.1*05 0.017 116 6.6 20 1*9.5 1*7.2 81* 81* 0.120 0.155 16b. 5 8.8 21 1*8.3 1*7.2 92 85 0.006 0.129 52 9.b 22 51.5 50.3 92 91 0.023 0.01*6 57.5 23 52.5 51.9 96 81* 0.31*0 93 0.9 21* 5i.o 50.2 9b 95 0.337 0.01*8 85 25 53-7 52.8 91* 80 0.030 0.056 73.5 7.b 26 52.1 50.9 92 83 0.101 122.5 9.7 27 5U.0 51.9 87 72 0.10b 98.5 9.5 28 52.2 50.5 89 78 0.09b 115 9.5 29 5 U .l 52.3 89 67 0.117 123 9.6 30 -31 53.8 53.0 9b 73 0.131 61.0 8.6 Sum 1630.2 1560.9 2588 2276 1.506 3.183 2412.0 215.6 Mean 51*.31* 52.03 86.3 75.9 0.0502 0.1061 80.b 7.19 A p p e n d i x L C r o w n - s h a p e s o f t h r e e d i f f e r e n t l y l o c a t e d t r e e s 88 Open growth tree (No- 10) Interior growth tree (No 25) 89 Appendix M Table 51. Average li v i n g and dead insects per needle, on trees No. 1, 2, 10, 11, Hi and 20. Living Generation 1 Tree No. Low level Medium level High level N W E N W E N W E IM 2L lhE 20M 10L 11E 072U 0.1*2 0.81* 0.08 0.16 0.36 07T6 0.1*8 0.66 0.08 0.1*0 0.31* 0.11* 0.50 0.1*6 0.00 0.12 0.1*1* OTllT 0.01* 0.78 0.06 0.16 0.36 0.10 0.16 1.01* 0.01* 0.08 0.06 0.22 0.38 0.86 0.02 0.06 0.01* 0.02 0.11* 0.58 0.01* 0.28 o.ob "072U 0.31* 0.76 0.00 0.21* 0.12 0702 0.02 0.38 0.02 0.00 0.00 0.00 0.02 0.32 0.06 O.OU 0.00 0.00 0.00 0.30 0.00 0.00 0.02 0.00 0.02 0.U2 0.00 0.02 0.00 Generation 2 0.23 "072T 0.20 1.76 O.OU 0.11 0.25 072T 0.18 1.5U 0.03 0.03 0.06 "6715" 0.10 0.76 0.03 o.ou 0.01 "5715" 0.11 1.77 0.03 0.03 o.ou "oToT 0.06 0.U5 0.01 o.oo o.oo o.ou 0.01 0.16 0.01 0.00 0.00 0.01 0.00 o.Ui 0.00 0.00 0.00 IM 2L I U E 20M 10L 11E 0.33 0.39 1.51 0.06 0.06 0.23 0.26 1.75 0.06 0.05 0.18 0715 0.31 0.7U 0.06 0.10 0.23 0713 0.10 1.11 0.01 0.01 0.03 0.01 0.01 0.U3 0.01 0.00 0.01 Dead" Generation 1 "OTHT 0.18 0.66 O.OU 0.12 0.06 0.06 O.OU 0706 0702" IM 2L IUE 20M 10L 11E 0.20 0.18 0.7U 0.08 0.08 0.20 0.10 0.26 0.61* 0.08 0.20 0.30 o.oa 0.20 0.58 0.02 0.30 0.22 0.08 0.72 0.08 0.22 0.22 "075B" 0.12 0 . 6 U 0.10 0.10 0.06 0 . 0 6 0.16 0.U2 0.02 0.06 0.10 "0732-0.12 0.80 0.02 O.UO 0.12 0.06 0.72 O.OU 0.02 0.00 0.02 0.56 O.OU 0.00 0.02 0.08 0.66 0.02 0.00 0.02 0.02 0.68 0.02 0.00 0.02 Generation 2 077J6~ 7J70T 0.09 0.23 0.01 0.00 0.01 1570T 0.01 0.13 0.00 0.00 0.01 o.ou 0.01 0.11 0.00 0.00 0.00 ~o7bT 0.00 0.08 0.01 0.00 0.00 " I F 2L IUE 20M 10L 11E 0.20 0.09 0.15 0.03 0.05 0.09 0.05 0.20 0.00 0.03 0.05 7J709" 0.06 0.10 o.ou o.ou 0.05 "0709 0.09 0.16 0.01 0.03 o.ou •urn* 0.10 o.iU o.ou 0.03 0.01 TJ7HT 0.06 0.13 0.01 0.03 0.00 0.11 0.09 0.11 0.01 0.03 0.00 0.01 0.00 0.10 0.03 0.00 0.00 Key: E = early bud-opened tree, M = medium bud opened tree, L = late bud-opened tree, N = North, S = South, W = West, E = East. 90 Appendix N Table 52. Average numbers of li v i n g and dead insects per needle by level and tree. Generation 1 Generation 2 Level Low Medium High Low Medium High T. No _L "OF 0.36 0.26 0.ij6 0.31 0.21 0.07 0.27 0.07 0.21 0.38 0.36 0.27 0.69 0.17 0.06 0.0k 0.03 0.05 0.06 o.ok 0.21 0.11 o.ok 0.02 D D D "oTIT 0.26 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 _L "072lT 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 0.01 D 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 0.13 0.18 0.19 0.19 0.27 0.12 0.22 0.19 0.03 0.20 0.2k 0.16 0.23 0.72 0.10 0.05 0.03 0.03 0.06 0.07 0.03 0.13 0.10 0.03 0.01 0.20 0.16 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 0.01 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 0^5 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 0.00 o.ok 0.11 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 0.16 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 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 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 Key: L = number of li v i n g insects, D = number of dead insects. 91 Appendix 0 Table 53. The average number of living insects per needle related to 8 factors. T.No . XI X2 x3 XU x5 X6 X7 XB Y 1 5 .5 37 2U 19 18 51.3 2 u 0.25 2 5.U Uo 20 10 11 72 .5 3 u 0.35 3 7.9 UU 20 13 9 79 .5 3 u 0.32 U 8.7 62 26 16 11 82.3 1 3 0.U8 5 9.6 63 25 20 7 88.9 3 3 0.U3 6 10.8 66 29 17 12 81.8 1 8 0.23 7 10 .3 58 28 19 8 86.2 3 8 0.08 8 12.1 59 26 22 12 79.7 1 8 0.30 9 u.u 25 27 10 7 72.0 3 2 0.08 10 12.8 63 22 32 10 8U.1 3 9 0.18 11 11.1 53 28 2U 9 83.0 1 9 0.2U 12 9.7 5U 25 22 9 83.3 1 7 0 .25 13 3 .3 19 25 9 6 68.U 2 5 0.17 LU 3 .3 21 20 11 2 90 .5 1 5 1 .65 15 2.7 17 20 10 u 76 .5 3 6 0.28 16 6.6 61 2U 18 18 U7.5 1 1 0.05 17 5.8 67 2U 1U 38 U3.3 3 1 0.03 18 7 . 5 52 25 17 33 3 6 . 5 2 1 0 .05 19 U.9 U9 2U 11 37 2U.5 1 1 0.07 20 5.2 U5 26 11 17 62.2 2 1 0.07 21 6.8 69 25 1U 3U 50.7 1 1 0.07 22 0.6 2 U 2 00 99.9 1 9 0.U6 23 2.2 8 7 u 1 92.2 3 9 0.30 2U 3 .7 31 22 8 16 U8.U 3 1 0.06 25 2.5 65 2U 1U Ul 36.9 3 1 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 e faced East, 6 = faced East-North, 7 = faced South-East, 8 = faced South-West, 9 = open growth. XI = D.B.H. in inches, X2 = height in feet, X3 = age, XU = crown width in feet, X5 = clear stem in feet, X6 = % of living crown, X7 a bud open-ing time, X8 = location of tree, Y = average number of living insects per needle. 92 Appendix P Table 5U. Percentages of mortality by generation and tree. Generation 2 Tree No. Generation 1 l^ofD "51772-38.2 Uo.o 32 .U U6.2 U2.1 75.0 U6.9 1*2.9 1*8.1 1*1* .1* 37.5 1*6.7 52.3 1*7.1 50.0 33.3 1*0.0 57.1 55.6 20.0 36.8 1*1*.0 50.0 75.0 TToTF "D L + 5 %olB Generation 1+2 D ~07lT 0.13 0.10 0.11 0.18 0.08 0.12 0.15 0.03 0.13 0.12 0.09 O.U* 0.68 0.16 0.03 0.01 0.02 O.OU o.o5 0.01 O.U* 0.11 0.03 0.03 L+D "072TT 0.31* 0.25 0.31* 0.39 0.19 0.16 0.32 0.07 0.27 0.27 0.2U 0.30 1.30 0.3U 0.06 0.03 0.05 0.07 0.09 0.05 0.38 0.25 0.06 o.oU "072TT L ~o~7Hr o.U* 0.17 0.25 0.22 0.12 O.OU 0.13 o.ou O.OU 0.09 0.10 0.01 1.03 0.10 0.02 0.01 0.02 o.ou 0.03 0.03 0.22 0.16 0.03 0.00 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 D IT+D %6fD 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 0711 0.21 0.15 0.23 0.21 0.11 o.ou 0.17 o.oU o.U* o.i5 0.15 0.16 0.62 0.18 0.03 0.02 0.03 0.03 o.oU o.ou 0.2U O.U* 0.03 0.01 "072T 0.19 0.22 0.32 0.27 0.17 0.06 0.17 o.ou 0.06 0.12 0.13 0.02 1.16 0.15 0.03 0.03 0.03 0.06 0.05 o.ou 0.28 0.23 0.05 0.00 "o7lo~ "3371 26.3 22.7 21.9 18.5 29.1* 33.3 23.5 00.0 33.3 25.0 23.1 50.0 11.2 33.3 33.3 66.7 33.3 33.3 UO.O 25.0 21 .U 30.U UO.O 00.0 "2B7J "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 0.20 0.18 0.15 0.18 0.23 0.13 o.u* 0.19 0.03 0.15 0.15 0.12 0.15 0.81 0.21 o.ou 0.03 0.03 0.06 0.07 0.02 0.20 0.18 0.05 0.03 0.15 "oTUT 0.53 0.U7 0.66 0.66 0.36 0.22 0.U9 0.11 0.33 0.39 0.37 0.32 2.U6 O.U9 0.09 0.06 0.08 0.13 o.U* 0.09 0.66 0.U8 0.11 O.OU o.ui 31*.0 31.9 27.3 3U.8 36.1 63.6 38.8 27.3 U5.5 38.5 32 .U U6.9 32.9 U2.9 UU.U 50.0 37.5 U6.2 50.0 22.2 30.3 37.5 1*5.5 75.0 U0.6 Ave. 0.13 0.11 0.13 Key: L = number of li v i n g insects per needle, D' = number of dead insects per needle. 93 BIBLIOGRAPHY 1. Balch, R. E., and G. R. Underwood. 1950. The lif e - h i s t o r y of Pineus pinifolae (Fitch) (Homoptera: Phylloxeridae) and i t s effect on white pine. Can. Ent. 82j 117 - 123. 2 . Balch, R. E. 1952. Studies of the Balsam woolly aphid, Adelges piceae (Ratz) and i t s effects 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 eghajlat-tan alapjai. Unpublished, University of Sopron. k. Cameron, A. E. 1936. Adelges cooleyi Gillette (Hemiptera, Adelgidae) of the Douglas f i r i n Britain Completion of i t s l i f e cycle. Ann. Appl. B i o l . 23: 585 - 6 0 5 . 5. Cameron, A. E. 1936. The present of status of the Douglas f i r woolly aphids (Adelges cooleyi Gillette) i n Britain. Forestry 10: 133 - 3l*2. 6 . Chystal, R. N. 1916. The l i f e history of Chermes cooleyi Gillette i n Stanley Park, Vancouver, B.C. Ent. Soc. Ont. k 6 t h Ann. Rept.: 123 - 130. 7. Chrystal, R. N. 1922. The Douglas f i r chermes (Chermes cooleyi). For. Comm. Bull. 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. 1956. The Adelginae of Alberta, identification and investigation of l i f e histories. I. Adelges cooleyi ( G i l l . ) . Unpublished report. For. Bi o l . Lab. Calgary, Alta. 1 0 . Cumming, M. E. P. 1959. The Biology of Adelges cooleyi ( G i l l . ) (Ho-moptera: Phylloxeridae). Can. Ent. 91: 601 - 617. 11 . Forest Club. U.B.C. 1959. Forestry Handbook for British Columbia. 2nd Ed. The Forest Club, U.B.C., Vancouver 8 , B.C. Canada, pp. 800. 1 2 . Forestry Commission. 19k6. Adelges attacking Spruce and other conifers. Leaflet No. 7. 1 3 . Forestry Commission. 19U7. Adelges cooleyi an insect pest of Douglas f i r and Sitka spruce. Leaflet No. 2 . Ik. Francke - Grosmann, H. 1950. Die Douglasienlaus G i l l e t t e e l l a cooleyi ( G i l l . ) C.B. als Schadling der Sitkafichte. Forst-Wissenschaftliches Centralblatt. 69 ( 8 ) . k83 - U93. 15. Geiger, R. 1950. The Climate near the ground (Tanslated from the second German edition of Das Klima der Bodennahen Luftschicht). Harvard University 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 of Briti s h Columbia Research Forest as related to climate and s o i l . U.B.C, For. Bull. 2. 17. Gyorfi, J. 1957. Erdeszeti rovarton. Akademiai Kiado, Budapest. pp. 670. 18. Herrick, G. W., and T. Tanaka. 1926. The Spruce Gall-aphid. Cornell University, Agr. Exp. Sta. Bull. k5k. 19. Imms, A. D. 1925. A General Textbook of Entomology. 9th Ed. Methuen and Co. Ltd., London, pp. 886. 20. Jeffers, J. N. R. i960. Experimental design and analysis i n forest 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 Territory. Edmond Cloutier, C.M.G., O.A., D.S.P., Ottawa, pp. 222. 22. Laing, F. 1929. The Douglas f i r Adelges. Scot. For. J. k3: k - 10. 23. Smith, D. N. 1956. Studies on Adelges cooleyi G i l l , i n relation to two-year Douglas f i r seedlings. Unpublished report. For. B i o l . Lab. Victoria, B.C. 2k. Snedecor, G. W. 1956. S t a t i s t i c a l methods. 5th Ed. Iowa State College Press, Ames, Iowa. pp. 53k. 25. Steel, R. G. D., and J. H. Torrie. I960. Principles and procedures of Statistics with special reference to the biologi-cal sciences. McGraw-Hill Book Company, Inc., New York, Toronto, London, pp. k8l . 26. Teucher, G. 195k. Die Douglasienwollaus. ( G i l l e t t e e l l a cooleyi G i l l . ) . Institut fur 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 Silver f i r s . For. Comm. Bull. 26. A p p e n d i x L C r o w n - s h a p e s o f t h r e e d i f f e r e n t l y l o c a t e d t r e e s Interior growth tree (No- 25) Fig-15 Relationship between the number of living insects per needle and height on the living crown (A = interior growth tree, B = marginal growth tree) a> T3 co CD z 0-4r CD CL % 0-31-CD co _c 2 0-2 o 6 CD > < 01 0-5 5-10 10-15 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 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 F i g - 1 3 R e l a t i o n s h i p b e t w e e n n u m b e r o f l i v i n g i n s e c t s p e r n e e d l e a n d d i s t a n c e f r o m t h e s t e m a t t h e l ow Distance from the Stem — Feet F i g - II R e l a t i o n s h i p b e t w e e n n u m b e r o f l i v i n g i n s e c t s p e r n e e d l e a n d h e i g h t o n t h e l i v i n g c r o w n J u n e 5 32-5 375 42-5 Height from the Ground — Feet F i g - 1 0 R e l a t i o n s h i p b e t w e e n t w i g l e n g t h s a n d n u m b e r o f l i v i n g i n s e c t s p e r n e e d l e Log Sampling Period 01 02 0-3 0-4 0-5 0-6 07 0-8 0-9 10 Log- Sampling Period 0 1 0-2 0-3 0-4 0-5 0-6 0-7 0 8 0-9 10 F i g 9 T r a n s f o r m e d c u r v e s o f f i g - 7 ( B ) a n d f i g - 8 ( A ) 1 ° July 9 i6 23 30 A u p 13 20 27 Sept-3 10 17 24 Sampling Period F i g - 8 N u m b e r 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 2 f o r d i f f e r e n t p e r i o d s a t the low l e v e l o f t h e l i v i n g c r o w n 27 \ \ \ \ \ \ \ \ I I I L Number of living insects Number o* dead \rv July 9 16 23 30 Aug 6 13 20 27 Sept 3 10 17 24 Sampling Period 7 N u m b e r 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 i n g e n e r a t i o n 2 f o r d i f f e r e n t p e r i o d s 2$ June 4 il 18 25 July 2 Sampling Period F i g - 6 N u m b e r 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 I f o r d i f f e r e n t p e r i o d s a t t h e l o w l e v e l o f t h e l i v i n g c r o w n June 4 II 18 25 July 2 Sampling Period F i g - 5 N u m b e r 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 I f o r d i f f e r e n t p e r i o d s 10 F i g - 4 A , S i s t e n s n y m p h , f i r s t i n s t a r B , P r o g r e d i e n s a d u l t ( A f t e r C h r y s t a l ) F i g - 2 A , G a l l i c o l a m i g r a n s w i t h w i n g s c l o s e d l a y i n g e g g s o n f i r n e e d l e ( X 2 0 ) B , E g g - m a s s p r o d u c e d b y G a l l i c o l a o r p r o g r e d i e n s ( X 3 0 ) F i g - 1 T h e l i f e c y c l e o f A d e l g e s c o o l e y i 

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