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UBC Theses and Dissertations

Population dynamics of the Lodgepole Needle Miner, Recurvaria starki Free. (Lepidoptera: Golechiidae)… Stark, Ronald William 1957

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F a c u l t y o f G r a d u a t e S t u d i e s PROGRAMME OF THE FINAL ORAL EXAMINATION F O R T H E D E G R E E O F DOCTOR OF PHILOSOPHY of R O N A L D W I L L I A M S T A R K B.ScF. University of Toronto M . A . University of Toronto I N R O O M 187A, B I O L O G I C A L SCIENCES B U I L D I N G M O N D A Y , F E B R U A R Y 24, 1958 at 2:30 p.m. C O M M I T T E E I N C H A R G E DEAN G . M . SHRUM, Chairman I. M c T . C O W A N W . S. H O A R K. G R A H A M G. J. SPENCER R. D A N I E L L S G . S. A L L E N V . K R A J I N A P. A . L A R K I N External Examiner: Dr. E. M . D U P O R T E McGi l l University A B S T R A C T The lodgepole needle miner, Recurvaria starki Free, has been studied intensively since 1948. Until 1953, this insect was referred to in publica-tions as Recurvaria milleri Busck. The life history and taxonomic position of R. starki are reviewed briefly and an historical review of the research carried on since 1948 is given. A full description is given of the procedure of applying life table techniques to needle miner studies since 1954 and examples are given for selected study areas. Six sampling intervals, one egg, four larval and one pupal are deemed suitable to assess the course of the population of a single generation from the time of oviposition to moth emergence. The life tables and survivorship and death-rate curves show clearly that there are five periods in the two-year life cycle of the lodgepole needle miner during which extensive mortality may occur: (1) between egg formation and oviposition; (2) between oviposition and larval estab-lishment; (3) during the first larval hibernation; (4) during the second larval hibernation; (5) during the spring of moth emergence. Population success is also undoubtedly affected by conditions during the adult life. Population sampling shows that the outbreak has declined since 1948 and defoliation and increment studies show that the period of greatest de-foliation occurred from 1940 to 1944 and that the outbreak probably began in the late 1930's. It is shown that winter temperatures, probably those of the coldest month, were responsible for the decline. It is esti-mated that the needle miner populations in Banff National Park can have a high survival if extreme minima of — 3 0 ° F . to — 4 0 ° F . do not persist long enough to depress the mean monthly temperature close to or below the zero mark. Parasitism was not an important factor in the outbreak decline and it was shown that this was probably due to a greater mortality in parasite populations due to winter temperatures. Other natural control factors are discussed as well as the possible effects of climatic factors on oviposition and fecundity. From a detailed survey of records since 1920 and yearly averages since 1885, it is postulated that the origin of the needle miner outbreak was due to a warming trend in the climate of the region. This began in the late 1930's, reached a peak in the mid-1940's and has declined since that time. The warming trend has been noted by other authors for north-ern latitudes and is substantiated by the weather records of this region. From these data it is further postulated that the climate of this part of Western Canada is generally too severe for an outbreak of the lodgepole needle miner, Recurvaria starki Free, to be prolonged. P U B L I C A T I O N S Stark, R.W. Sequential sampling of the lodgepole needle miner. For. Chron. 28(2): 57-60, 1952. Stark, R.W. Analysis of a population sampling method for the lodge-pole needle miner in Canadian Rocky Mountain Parks. Canad. Ent. 84(10): 316-321. 1952. Stark, R .W. Distribution and life history of the lodgepole needle miner (Recurvia sp.) Lepidoptera: Gelechiidae) in Canadian Rocky Mountain Parks. Canad. Ent. 86(1): 1-12, 1954. Stark, R.W. The effects of defoliation by the lodgepole needle miner (Recurvaria starki Free). In Press. Forest Science. Graham, K. and Stark, R .W. Insect Population Sampling. Ent. Soc. Brit. Col. Proc. 1954. G R A D U A T E S T U D I E S K. Graham J. D. Chapman W . S. Hoar . . .V. J. Krajina Other Studies: Forest Research Methods Biological Methods Forestry Seminar Field of Study: Forest Entomology Forest Insect Ecology Climatology Experimental Zoology Forest Associations POPULATION DYNAMICS OF THE LODGEPOLE NEEDLE MINER,, RECURVARIA STARKI FREE. (LEPIDOPTERA: GELECHIIDAE) IN CANADIAN ROCKY MOUNTAIN PARKS. RONALD WILLIAM STARK B.S 0.F.,(l948), M.A.(l95l) University of Toronto, A THESIS SUBMITTED IN PARTIAL FULFILMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY IN FOREST ENTOMOLOGY i n the Departments of Forestry and Zoology We accept t h i s thesis as conforming to the required standard THE UNIVERSITY OF BRITISH COLUMBIA December, 1957 - i -ABSTRACT The lodgepole needle miner, Recurvaria s t a r k i Free, has been studied intensively since 194.8. U n t i l 1953, this insect was referred to i n publications as Recurvaria m i l l e r l Busck. The l i f e history and taxonomic position of R. s t a r k i are reviewed b r i e f l y and an h i s t o r i c a l review of the research carried on since 194-8 i s given. A f u l l description i s given of the procedure of applying l i f e table techniques to needle miner studies since 1954- and examples are given f o r selected study areas. S i x sampling i n t e r v a l s , one egg, four l a r v a l and one pupal are deemed suitable to assess the course of the population of a single generation from the time of oviposition to moth emergence. The l i f e tables and survivorship and death-rate curves show c l e a r l y that there are f i v e periods i n the two-year l i f e cycle of the lodgepole needle miner during which extensive mortality may occurJ ( l ) between egg formation and ovipositionj (2) between oviposition and l a r v a l establishment? (3) during the f i r s t l a r v a l hibernation? (4.) during the second l a r v a l hibernation; ( 5 ) during the spring of moth emergence. Population success i s also un-doubtedly affected by conditions during the adult l i f e . Population sampling has shown that the outbreak has declined since 194-8. Defoliation and increment studies have shown that the period of greatest d e f o l i a t i o n occurred from 1940 to 1944- and that the outbreak probably began i n the l a t e 1930's. The major cause of - l i -the decline was winter temperatures, probably during the coldest month. From laboratory experiments and population sampling compared with weather records i t is estimated that needle miner populations can have a high survival i f extreme minima of -30°F to -4-0°F do not persist long enough to depress the mean monthly temperature to near 0°F. Parasitism was not a particularly important factor in the outbreak decline probably because of a greater depressant effect on parasite populations by winter temperatures. Other natural control factors are discussed as well as the possible effects of climatic factors on oviposition and fecundity. From a detailed survey of weather records since 1920 and yearly averages since 1885 i t is postulated that release of the needle miner population was due to a warming trend in the climate of the region. This began in the late 193O's, reached a peak in the mid-194.0's and has declined since that time. The warming trend has been noted by other authors for northern latitudes and is substantiated by the weather records of this region. It is further postulated that the climate of this part of western Canada is generally too severe for an outbreak of the lodgepole needle miner, Recurvaria starki Free, to be prolonged. In presenting t h i s thesis i n p a r t i a l f u l f i l m e n t of the requirements f o r an advanced degree at the University of B r i t i s h Columbia, I agree that the l i b r a r y s h a l l make I t f r e e l y available f o r reference and study. I further agree that per-mission f o r extensive copying of t h i s thesis f o r scholarly purposes may be granted by the Head of my Department or by h i s representative. I t i s understood that copying or publication of t h i s thesis f o r f i n a n c i a l gain s h a l l not be allowed without my written permission. Department of Zoology The University of B r i t i s h Columbia, Vancouver 8, Canada. Date December 30. 1957 - I i i -TABLE OF CONTENTS 1. INTRODUCTION 1 1.1. The organism 2 1.1.1. Taxonomy 2 1.1.2. L i f e h i s t o r y 3 1.1*3* Asynchronized population phenomena 4-1.2. The host 6 1.3. Description of the region 7 1.4-. H i s t o r i c a l review 9 2. LIFE TABLES FOR THE LODGEPOLE NEEDLE MINER U 2.1. Review of l i f e tables i n entomology 14-2.2. The preparation of l i f e tables.... 16 2.3. L i f e tables f o r the lodgepole needle miner..... 18 2.A. Examples of needle miner l i f e tables 28 Example 1. Mount Eisenhower, v a l l e y bottom, 29 Example 2. Mount Eisenhower, 5,4-00' a. s . l 38 Example 3. Massive Range, 5,500' a . s . l 4-0 Example 4-. Mount Girouard, 6,000' a. s . l 41 Example 5. Cathedral Mountain, 4->700' a . s . l 4-2 3. DISCUSSION OF NATURAL CONTROL FACTORS 4-3 3.1. Climatic factors... 4-3 3.1.1. Winter mortality 43 (1) Theories of cold resistance and death i n insects 44-(2) Winter temperatures as a c o n t r o l l i n g factor i n insect populations 46 (3) Winter mortality i n needle miner populations 4-8 (4.) Climatic conditions causing winter mortality observed i n lodgepole needle miner populations 55 - i v -3.1.2. Spring mortality 67 3.1.3. Other climatic factors possibly involved in the reduction of populations 67 (1) Effects of weather on eggs and first-instar larvae 67 (2) Effects of climatic factors on larvae during the summer. 70 (3) Effects on pupae 71 (4.) Effects of various factors on moth behaviour in respect to needle miner abundance 72 3.2. Parasitism 74 3.2.1. Description of the parasite complex of R. starki Free. 75 3.2.2. Population dynamics of needle miner parasites 83 3.3. Predation 98 3.3.1. Types of predation 98 3.3.2. Importance of predation in dynamics of needle miner populations .100 3.4. Disease ....101 3.5. Other natural control factors 107 3.5.1. Resination 107 3.5.2. Competition and Toverpopulation' factors 108 EPIDEMIOLOGY 110 4.1. Epidemiology since 1942 110 4.2. The origin of the outbreak . . . . . I l l 4..2.1. The theory of climatic release... .111 4-.2.2. The theory of climatic release applied to the outbreak of the lodgepole needle miner, Recurvaria starki Free.117 (1) Climatic controls of the region 117 (2) Climate and epidemiology of the lodgepole needle miner..119 (3) Geographical limitation of the needle miner outbreak....121 (4.) General discussion of needle miner epidemiology with regard to future studies .12? - V -5. LITERATURE CITED 125 6. APPENDICES 136 1. Daily maximum and minimum temperatures f o r winter months, Banff, Alberta. 1920 - 1954 137 2. Daily maximum and minimum temperatures f o r winter months, Lake Louise, Alberta. 1932 - 1954 171 3. Daily maximum and minimum temperatures f o r winter months, Calgary, Alberta. 1954 - 1956 193 4. Mean monthly temperatures, Banff, Alberta 1920 - 1954.195 5. Mean monthly temperatures, Lake Louise, Alberta. 1932 - 1954 196 6. Comparative records, Calgary, Alberta. Monthly and annual averages and extremes for t o t a l period of observation (1885 - 1955) 197 7. Annual and 5-year running mean temperatures .198 8. Mean monthly temperatures f o r selected stations -winter months 199 9. Air-mass summaries, Bow Valley drainage, winter months 2d LIST OF ILLUSTRATIONS Following page Figure 1. Stages i n the l i f e h i s t o r y of Recurvaria s t a r k i Free 3 n 2. Schematic i l l u s t r a t i o n of l i f e cycle of the lodge-pole needle miner and sampling periods for l i f e - t a b l e studies 3 " 3. Alberta d i s t r i b u t i o n of lodgepole pine, Pinus CQntprta, var. l a t i f o l i a (After Moss,195577777 7 n 4* Map of western Canada showing location of needle miner outbreak ( c i r c a 1948) 7 - v i -Figure 5 . Survivorship and death-rate curves f o r Mount Eisenhower v a l l e y bottom U , 8 0 0 ' ) and 5 ,4-00 ' f o r the 1952-54- generation of needle miner 3 2 n 6 . Survivorship and death-rate curves f o r the 1954--56 generation. Mount Eisenhower, v a l l e y bottom U , 8 0 0 ' ) 33 ff 7 . Survivorship and death-rate curves f o r the 1954--56 generation. Mount Eisenhower, 5,400'.... 3 7 " 8 . Survivorship and death-rate curves, 1 9 5 4 - 5 6 gener-atio n . Massive Range, 5,500' 4-0 B 9 . Survivorship and death-rate curves, 1954--56 gener-atio n . Mount Girouard, 6 , 0 0 0 ' 4 1 n 1 0 . Survivorship and death-rate curves, 1954--56 gener-at i o n . Cathedral Mountain, 4-»700' 4 5 " 1 1 . . Needle miner d i s t r i b u t i o n p r i o r to 1 9 4 9 - 5 0 • 44-tt 1 2 . Needle miner d i s t r i b u t i o n following 1 9 4 9 - 5 0 . 4 4 " 1 3 . Mortality of lodgepole needle miner at d i f f e r e n t elevations i n four selected years 54-B L i * Hygrothermograph tracing showing temperature con-d i t i o n s at v a l l e y bottom ( 4 , 8 0 0 ' ) and 5 f 5 0 0 ' on Mount Eisenhower from January 29 to February 1 , 1 9 5 6 when area was i n polar continental a i r 57 " 1 5 Hygrothermograph tracing showing temperature con-ditions i n the same locations from February 3 to 6 , 1 9 5 6 when the area was i n polar maritime a i r 57 n 1 6 . Hygrothermograph tracing showing humidity and temper-ature conditions on Mount Eisenhower, 5 , 5 0 0 ' , from August 18 to 25, 1954- and 1956 6 9 n 1 7 . Parasites R. nta r k i Free 8 2 B 1 8 . Parasites R. s t a r k i Free 8 2 n 1 9 . Mean monthly temperatures f o r December, January, and February, 1 9 2 0 - 1 9 5 4 . Banff, Alberta 1 1 9 " 2 0 , Mean monthly temperatures f o r December, January, and February, 1 9 3 2 - 1 9 5 3 . Lake Louise, Alberta 1 1 9 " 2 1 . Annual mean temperature and 5-year running mean, 1893 - 1 9 5 5 . Banff, Alberta 1 2 1 " 2 2 . Topographic map of Banff National Park showing the outbreak region md the main sampling areas.. F l y l e a f - v i i -LIST OF TABLES Table I Egg sampling, 1954- 21 " I I Egg sampling, 1956 21 " I I I Comparison of egg sampling and f i r s t l a r v a l establishment, 1956 22 " IV L i f e table f o r the 194.8-50 generation of needle miner. Mount Eisenhower, v a l l e y bottom.... 31 " V L i f e table for the 1952-54- generation of needle miner. Mount Eisenhower, v a l l e y bottom 32 " VT L i f e table for the 1954--56 generation of needle miner. Mount Eisenhower, v a l l e y bottom....." • 33 " VII L i f e table f o r the 194-8-50 generation of needle miner. Mount Eisenhower, 5,400' a . s . l 34 " VIII L i f e table for the 1950-52 generation of needle miner. Mount Eisenhower, 5,400' a . s . l • 35 " IX L i f e table for the 1952-54 generation of needle miner. Mount Eisenhower, 5,400' a . s . l 36 " X L i f e table f o r the 1954-56 generation of needle miner. Mount Eisenhower, 5,400' a . s . l 37 " XI Percentage parasitism of the t o t a l established l a r v a l population 39 " XII L i f e table f o r the 1954-56 generation of needle miner. Massive Range, 5,500' a . s . l • 40 " XIII L i f e table f o r the 1954-56 generation of needle miner Mount Girouard, 6,000' a . s . l 41 " XIV L i f e table f o r the 1954-56 generation of needle miner. Cathedral Mountain, 4*700' a . s . l 42 " XV Percentage winter mortality of established needle miner populations i n four locations, 1948-56 49 w XVI Recorded winter mortality. Mount Eisenhower, Banff National Park 50 n XVII Recorded winter mortality. Massive Range, Banff National Park 51 - v i i i -Table XVIII Recorded winter mortality. Cathedral Mountain, Yoho National Park 51 n XIX Recorded winter mortality. Lake Louise, Banff National Park 52 n XX Recorded winter mortality. Mount Norquay (Edith), Banff National Park 52 n XXI Recorded winter mortality. Bankhead, Banff National Park 52 n XXII Recorded winter mortality. Hawk (Snow) Creek, Kootenay National Park 53 " XXIII Recorded winter mortality. Miscellaneous areas, Banff National Park 53 n XXIV Recorded winter mortality by a l t i t u d e . A l l areas 54-w XXV Percentage winter mortality, 1953-54- 58 " XXVT Daily maximum and minimum temperatures, January 10 - 29, 1954-. Banff and Lake Louise 60 " XXVTI Summary of winter temperature data, years of low mortality. Banff, Alberta 64 w XXVIII Summary of winter temperature data, years of high mortality. Banff, Alberta 65 w XXLX Parasite complex of Recurvaria s t a r k i Free 78 " XXX Percentage parasitism i n moth f l i g h t years. A l l areas 84 n XXXI Percentage parasitism i n moth f l i g h t years f o r four areas now under constant investigation 85 " XXXII Species composition and percentage parasitism i n a l l areas sampled i n 1954 87 n XXXIII Supplement to Table X showing mortality of l a t e ins t a r larvae and pupae. Mount Eisenhower 88 " XXXTV Supplement to Table XII showing mortality of l a t e i n s t a r larvae and pupae. Massive Range 89 n XXXV Supplement to Table X I I I showing mortality of l a t e i n s t a r larvae and pupae. Mount Girouard 90 " XXXVT Supplement to Table XIV showing mortality of l a t e i n s t a r larvae and pupae. Cathedral Mountain 91 - i x -ACKNOWLEDGMENTS Grateful acknowledgments are due the following* K. Graham, University of B r i t i s h Columbia, who, i n 1949 and 1950, guided the f i r s t steps of the project i n population sampling upon which much of t h i s thesis i s based. His guidance was also given during the authors post-graduate study at the University of B r i t i s h Columbia. W.R, Henson, Yale University, kindly permitted the use of portions of h i s air-mass analyses and contributed much to the t h e o r e t i c a l considerations. G.R. Hopping, Officer-in-Charge, Forest Biology Laboratory, Calgary, Alberta, has given the benefit of h i s advice on the project since i t s inception. Other research o f f i c e r s at Calgary, Alberta have contributed to various phases of the work, p a r t i c u l a r l y J.A. Cook, Miss M.E.P. Cumming and R.F. Shepherd. Thanks also are due the Calgary and Edmonton Meteorological Offices of the Department of Transport whose heads gave free access to t h e i r records and data. Thanks are owing to M.L. Prebble, Chief, Forest Biology D i -v i s i o n , f o r permission to use, f o r t h i s t h e s i s , data gathered In the course of work on a D i v i s i o n a l project. T.N. Freeman, Systematics D i v i s i o n , Division of Entomology, described and named t h i s species of needle miner and to him my thanks are due. POPULATION DYNAMICS OF THE LODGEPOLE NEEDLE MINER, RECURVARIA STARKI FREE. (LEPIDOPTERA: GELECHIIDAE) I N CANADIAN ROCKY MOUNTAIN PARKS by R.W. STARK 1. INTRODUCTION The lodgepole needle miner is a defoliating insect which attracted attention because of its increase to outbreak abundance in the Canadian Rockies during the 194-0's. The forests attacked by this insect cover a vast western watershed and are the main forest stands in four National Parks. The stands are adjacent to the extensive lodgepole pine stands of the Eastern Slopes of the Rocky Mountains in Alberta which are a major source of revenue to the forest industry of Alberta. These considerations made i t of paramount importance to analyse the factors responsible for the increase in numbers in the region, to explain zones of abundance within the outbreak and to determine the factors which may limit outbreaks to the region recent-ly affected. Since the mid-1940's needle miner populations have declined until at the present time (1957) outbreak conditions do not exist f This leads to the question whether the outbreak may recur. No evidence exists that any previous outbreaks occurred in this region. The purpose of this thesis therefore, is to analyse the factors responsible for the decline in the outbreak since the mid-194-0's and to formulate an hypothesis for the origin of outbreaks of this insect. 2. 1.1. The organism 1.1.1 Taxonomy. When the outbreak was f i r s t discovered in 194-2, adult specimens were compared with the description of a species with a similar l i f e cycle occurring on lodgepole pine in Yosemite National Park, California, (50,52,93). It was con-cluded that they were the same species, Recurvaria millerl Busck of the family Gelechiidae, Lepidoptera (12). The use of this determination persisted until 1953 when T.N. Freeman of the Systematics Unit, Division of Entomology, Science Service, Department of Agriculture, Ottawa, indicated doubt that they were the same species (Freeman, T.N. pers. comm.). At his suggestion the insect was then referred to as Recurvaria sp. in publications. In 1954- a total of 1,962 needle miner adults from many points within the outbreak areas and from several elevations were sub-mitted to Freeman to assist him in unravelling the complex (125). In 1957 the species occurring in the Canadian outbreak was described by Freeman and named Recurvaria starki (30). His de-scription follows: Recurvaria starki Freeman, new species "Recurvaria milleri auct. (in part) nac. Bsk.j Hopping, 194-5, Proc. Ent. Soc. British Columbia 42: l-2j McLebd, 1951, Canadian Ent. 83: 295-301. Recurvaria sp., Stark, 1954* Canadian Ent. 86: 1-12. Antenna alternately marked with ocherous-white and black bands. Palpus rather short, not tufted in the male. Second joint of palpus whitish inwardly, ocherous-fuscous outwardly; third joint white with ocherous-fuscous base. Face and vertex shining white. Thorax and forewing light grey, the latter with somewhat obscure blackish patches crossing the wing at the basal third, at the outer two-thirds, and near the apexj the post-median band bordered out-wardly with white and appearing to be sharply angled outwardly at its middle. The patch near the apex extending obliquely inward almost to the posterior margin. Apex of wing mostly white, and with an obscure, blackish central area. Hind wing pale smoky. Fringes 3 of a l l the wings shiny, l i g h t , ocherous«-grey. Under surfaces of a l l the wings, d u l l white. Male with a long ocherous hai r -penci l ar is ing from beneath the anal angle at the base of the hind wing. Fore and mid t ibiae and ta rs i alternately banded with black and white scales. Hind t ib ia whit ish, with long hairs above. Each segment of hind tarsus grey, with a white t i p . Wing expanses 12 - 13 mm. Moth in the la t ter half of July . Male geni ta l ia . Uncus roof l ike , Gnathos with three hooklike processes,the median one s l igh t ly the longest. Caudo-lateral pro-jections of tenumen asymmetrical, f l ap l i ke . Claspers asymmetrical, tubular twistedj the r ight clasper much larger than the l e f t . Aedoeagus pistol-shaped. Vinculum produced apical ly into two some-what asymmetrical hooklike s icae. Holotype. - Male, Mt. Eisenhower (near Banff) , Banff National Park, Alberta, July 19, 1954.• Reared from Plnus contorta Dougl. by off icers of the Forest Insect Survey, Forest Biology Div is ion . No. 6298 in the Canadian National Col lec t ion , Ottawa. Paratypes. - Twenty males and 16 females, Mt. Eisenhower, Banff National Park, Alberta, July 17 and 19, 1954-• Eleven males and f ive females, Cascade Mountain (near Banff) , Banff National Park, Alberta, July 18 and 19* 1954-. Two males, Lake Louise, Banff National Park, Alberta, July 20, 1954.. Two females, Mt. Edith Cavel l (near Jasper). Jasper National Park, Alberta,.July'15,1954.. A l l paratypes reared from Pinus contorta. and No. 6298 in the Canadian National Col lect ion , Ottawa. Food plant. - Pinus contorta Dougl. This species is closely a l l i e d to and has been confused with &• M i l l e r ! Bsk. The male genital ia appear to be ident ica l with those of that species and of g. moreonella Heinr . , a species described from a single male reared from Pinus scopulorum (Engelm.) Lemmon at Cheyenne Mountain, Colorado. R. m i l l e r ! is somewhat larger , mainly white with black longitudinal streaks or d is t inc t irregular spots. R. moreonella has a narrow irregular l ine of white scales extending longitudinally through the middle of the wing, from the end of a sub-basal black streak to near the apex, ft. s tark! and R. m i l l e r ! are needle miners that require two years to complete their l i f e cycles in the type l o c a l i t i e s . There is some evidence to suggest that R. miller!< R. moreonella. and R. s t a r k i . as well as some a l l i e d species, do not belong to the genus Recurvaria Haw. The male genital ia of R. nanella Hbn., the genotype of Recurvaria. are b i l a te ra l l y symmetrical. The male genital ia of the group of species under consideration are asy-mmetrical. There is also a difference in the shape of the sugnum in the bursa of the female. Further studies are necessary to e l u c i -date the generic signif icance of these characters." 1.1.2 L i fe history. A photograph of the major l i f e stages is presented in Figure 1 and a schematic i l l us t ra t ion of the l i f e cycle Figure 1. Stages i * the l i f e h i s t o r y of RflCttrYfflla S±a£ki Free. Figure 2. Schematic i l l u s t r a t i o n of l i f e cycle of the lodgepole needle miner and sampling periods f o r l i f e - t a b l e studies. S C H E M A T I C I L L U S T R A T I O N O F L I F E C Y C L E O F T H E L O D G E P O L E N E E D L E M I N E R , R E C U R V A R I A S P . E V E N - N U M B E R E D Y E A R X - 3 i z X - 5 1 X-l E G G m H X - 6 X - 2 1 X - 4 1 2 P U P A N E X T E V E N - N U M B E R E D Y E A R M M A D U L T X = S A M P L E P E R I O D F O R L I F E T A B L E _1_ N in Figure 2. The needle miner in the Canadian Rockies has a two-year l i f e cycle. In the even years the adults emerge in July; eggs are laid in late July and August and hatch in August and September. Each larva immediately enters a needle in which i t spends the f i r s t winter. The following spring the miner commences to feed in late April or May, depending on spring weather and completes mining the f i r s t needle. Transfer to a second needle takes place in mid-summer; climatic conditions affect the time and duration of the transfer period considerably. The larva over-winters in the second needle and the following spring, again an even year, transfers to a third needle. It completes mining by early June, pupates and the moth emerges three to four weeks later. Details of the l i f e history have been published (123). 1.1.3* Unsynchronized population phenomena. There have been two instances of off-year maturation of the needle miner. Discussion of these has deliberately been omitted from the main thesis as i t was felt that i t would only lead to confusion. The f i r s t occurred in the Bow Valley of Banff National Park in 194-9 (32,34) and was restricted to an area from Mount Eisenhower to the town of Banff. Only a small portion of the total population was involved, a maximum of less than 30 per cent in any one location (32). Parasitism of this off-cycle population averaged 20 per cent (34). The second occurred in the Kiokinghorse Pass of Yoho National Park in 1951 and was very restricted in extent; maturing larvae and pupae were found in an area of less than one acre (123), The f i r s t indication that this might ocour was noticed in the winter of 1950. Larvae brought into the laboratory in September continued 5. to feed u n t i l by November 20 these Larvae were three times the size of larvae in the f i e l d . Larvae collected i n November resumed feeding under laboratory conditions and increased i n size (103). Sampling in July showed an average of two to three 'precocious * larvae per t i p , the average of a l l larvae per t i p was about 20. A to t a l of 305 advanced larvae were collected and reared. Parasitism proved to be high; of the 305 larvae collected 300 were parasitized (114-)• There are three possible explanations for this off»cycle occurrence of adults. The f i r s t is that there i s another species involved. Unfortunately no specimens were preserved for i d e n t i f i -cation. However, were another species present we would expect recurrence of adults in the odd-numbered years, progeny of those found. Careful watch has been kept since 194-9 and other than those found in 1951 i n another area, no further occurrence of the phenomenon has been noted. The second is that possibly part of the needle miner population possesses an obligatory diapause factor which automatically interrupts development regardless of environmental conditions. The remainder may lack a diapause factor and become sufficiently abundant i n the two locations found to be noticed (34-)• Although the numbers found were relatively small, there were enough that we would expect to have individuals occur i n the winter sampling which would show this diapause factor. A l l experiments made i n breaking diapause tend to discount this theory. The third theory i s that the small portion of the popu-lation affected were individuals which were i n particularly favorable niches and developed more quickly than their neighbors. They cannot be regarded as 'stronger' than those whioh did not mature in the off-year as i t is difficult to conceive of parasitized larvae being capable of more rapid development than unparasitized larvae in the same location. As no progeny of either example were found in following years, no further work could be done. The fact that no progeny were found makes any of the three explanations difficult to support. We can only regard i t as a chance occurrence where development was accelerated in certain individuals by some unknown factor but that they were incapable of reproduction. The effect of this phenomenon on the epidemiology of the total population was negligible, although had i t occurred later i t would have been listed as a mortality factor, as those individuals were lost to the normal population. In 194-9, this would have averaged 20 per cent, in 1951 about 10 per cent in the areas where the phenomenon occurred. Had offspring of this off-cycle popu-lation become established, we would have had to consider i t as a separate population. 1.2. The host Lodgepole pine is found from lower California to Alaska and the Yukon Territory and from sea level to an elevation of 11,000 feet in the Rocky Mountains. Some botanists recognize coastal and inland forms of lodgepole pine. The former is sometimes designated as Pinus contorta Dougl. whereas the inland form from north to south is known as P^ nus, contorta var, latifpJUfl Engelm.(76). In the outbreak area of Requrvarfta s t a r k i Free., Pinna contorta var. l n t i f o l i a Engelm. ( = P. contorta Dougl. of Freeman) is the sole host. The distribution of lodgepole pine in Alberta is shown in Figure 3 (82). The Alberta distribution given by Moss follows closely that given by Halliday and Brown (39) except for the region between Edmonton and Peace River. 1,3, Description of the region. Needle miner populations were found in four national parks; Banff. Yoho, Kootenay and Jasper. The outbreak dealt with in this thesis was more extensive and severe in Banff Park than in any of the others (Figure 4 and flyleaf). The terrain is mountainous, the mountains being steeper and more rugged on the east side than on the west. The parks are similar in that there is a relatively narrow valley in each with steep, high sides formed by the mountain ranges, often up to 10,000 feet in height. In Banff the direction of the valley is west-north-west; in Kootenay south, in Yoho, west. Timberline varies from about 6,500 feet in some areas to 8,000 feet, being around 7,200 feet in the Banff area (83). Valley bottom varies in altitude from about 4,000 feet at the eastern approaches to almost 5.000 feet at the Continental Divide. The outbreak areas f a l l within Halliday's sub-alpine forest region SA-1 (East Slope Rockies section). His description reads (38): "A characteristically coniferous forest covers the eastern slope of the Rockies from around the 4,000 foot elevation to the tree line. The sub-alpine dominant engelmann spruce forms the bulk of the forest cover, the subclimax lodgepole pine being mixed with this species following fires and often forming large areas of stands of varying degrees of purity. Alpine f i r enters toward the tree line, together with white-bark pine and, in the south, alpine (Lyall's) Larch. Along the lower slopes, in the southern parts, there is some intrusion of Douglas f i r from the Montana forest region and a fringe of aspen occurs where the section comes into contact with the Grassland formation." Figure 3. Alberta d i s t r i b u t i o n of lodgepole pine, Plnus co^forta var. l a t l f o l i a (After Moss, 1955). Figure 4-. Map of western Canada showing location of needle miner outbreak (circa 194-8). Hachured area represents the outbreak. I A L B E R T A E D M O N T O N i . . . . . . . . . . . * \ J A S P E R N. ? \ \ H ^ , . . . . . •••• « V ^ ^ B A N F F N! '?'.; Y O H O N. P.A K O O T E N A Y C A L G A R Y BRITISH C O L U M B I A \ I N T E R N A T I O N A L B O U N D A R Y i y W A S H I N G T O N jIDAHo! i M O N T A N A 8. Horton (59) describes the area as the "subalpine d i v i s i o n " corresponding clo s e l y to Halliday's SA-1 section. He, however, states: "the c h a r a c t e r i s t i c tree species i s engelmann spruce but subclimax stands of lodgepole pine predominate. White spruoe replaces engelmann i n the lower valleys and hybrid forms of the two species are common i n the intermediate zone. Black spruce i s present i n t h i s d i v i s i o n only north of the north Saskatchewan River and then only at lower l e v e l s , presumably as a r e s u l t of intrusion from the f o o t h i l l s along the low Athabasca v a l l e y . Alpine f i r i s a frequent component, p a r t i c u l a r l y i n older stands. Douglas f i r occurs spasmodically i n the three main mountain passes, which d i s t r i b u t i o n suggests intrusion from B r i t i s h Columbia. The poplars a t t a i n f a i r l y good development i n protected v a l l e y s i t e s but seem incapable of competing with the coniferous species on the mountain slopes." Horton divides the Eastern Slopes area, adjacent to the outbreak area, into two d i v i s i o n s , High F o o t h i l l s and Low F o o t h i l l s both of which l i e i n Halliday's B19 or F o o t h i l l s Section of the Boreal Forest Region. The High F o o t h i l l s d i v i s i o n i s "a long narrow s t r i p of country t y p i f i e d by high wooded h i l l s and deep v a l l e y s , usually between the altitudes of 4>000 and 6,000 feet." Lodgepole pine i s again preponderant but white and black spruce are the major species. Alpine f i r i s less prevalent than i n the mountains and poplar i s r e l a t i v e l y unimportant. The Low F o o t h i l l s Division i s comprised of low h i l l s and plateaux from 2,000 to 4,000 feet elevation. The main difference between t h i s and the High F o o t h i l l s D i v i s i o n i s the prevalence of mixedwood types. Aspen and balsam poplar are able to compete with lodgepole pine as po s t - f i r e pioneers. White and black spruce are common but alpine f i r i s rare. Hopping (54) recognizes s i x d i s t i n c t timber types i n the Canadian Rocky Mountains ^ none of^which disagree e s s e n t i a l l y with the descriptions given by Halliday and Horton. The main forest cover i n the outbreak area consists of lodgepole pine stands of varying ages but l a r g e l y mature (80 years or over) with a spruce understory. The extensive stands of lodgepole pine resulted from a series of f i r e s l a t e i n the nineteenth century. Horton (59) suggests that succession from pine to engelmann spruce-alpine f i r i s indicated f o r the sub-alpine region and the High F o o t h i l l s region, with the substitution of white spruce f o r engelmann and the addition of black spruce. Such successions may be completed i n periods ranging from 225 to 325 years. Moss (83) reviewed the l i t e r a t u r e on succession i n lodgepole pine i n Alberta and h i s views regarding the successions! position of lodgepole pine agree with those of Horton. 1.4. H i s t o r i c a l review. The outbreak was f i r s t noticed i n June, 194-2, on an area of approximately 50 square miles i n Banff National Park where i t joins Kootenay National Park at Vermilion Summit ( l l ) . The attack was la r g e l y confined to elevations between 5,000 and 6,500 feet from Vermilion Pass east to Brewster Creek on the south side and only four miles eastward on the north side of the Bow River. I t was p r a c t i c a l l y non-existent at v a l l e y bottom (4.5). A s l i g h t decrease was noticed i n 1943 (47,48,62) but i n 1944- i t had spread i n t o loho and Kootenay Parks for short distances. No marked a l t i t u d i n a l change was observed (63). In 194-5 the area of the outbreak was estimated as 200 square miles i n Banff, Yoho and Kootenay Parks, l a r g e l y i n the f i r s t (64). This estimate was revised to 300 square miles i n 194-6 (65). Surveys indicated that the outbreak had spread eastward almost as f a r as Banff, south into Kootenay Park as f a r as Hawk 10. Creek, north along the Banff-Jasper Highway to Hector Creek and for several miles westward into Yoho Park from Wapta Lake (52). It was also noted that the population had increased in stands below the 5,000 foot l e v e l s , that i s , in val ley bottom stands (5 l ) . In 1947 D.K. Campbell of the Vernon Forest Insect Laboratory spent some time studying the outbreak. He attempted to determine the populations present and experimented with chemical cont ro l . His results were inconclusive (13,14,15). The fourth generation of needle miner under observation (1946-48) increased in distr ibut ion to 400 square miles in the outbreak region (69). A second outbreak, centred in Jasper National Park, was reported. This was ent irely separate from the more southerly outbreak occurring on the slopes below Mount Edith Cavell (70,108), and was thought to be autochthonous (35). The Calgary Laboratory of Forest Zoology (later Biology) was established in May, 1948. One of i t s or ig inal projects was the lodgepole needle miner and the writer was assigned as project leader that year. Main objectives included the determination of the l i f e h is tory , precise sampling methods and possible controls (53). In 1949, K. Graham of the University of Br i t ish Columbia was assigned seasonally to the project with the author. Consider-able data were gathered on l i f e history and sampling procedures (32,34,110,111). The distr ibut ion in 1949 included areas adjacent to the Eastern Slopes forests . In Jasper the infestation was continuous 11. throughout the upper Sunwapta and Athabasca valleys on the forest f loor but confined to the higher alt itudes in the northerly sections of Jasper Park. The outbreak was generally l igh t but two areas were more seriously infested (36,71). Results of experiments conducted by Graham and Stark in 194-9 (35,112) indicated l i t t l e hope of success in chemical cont ro l . During the winter of 194-9-50 there was high mortality of lodgepole needle miner larvae. The region part icular ly affected was from the Great Divide eastward to the eastern l i m i t of the outbreak. Populations were pract ica l ly eliminated in stands at val ley bottom (56,57,113). Additional information on natural controls and l i f e history was obtained (55,103). During 1950-51 further studies were made on the effects of winter temperatures on larvae and diapause was found to be facultat ive (58). Temperature was found to have a profound effect on la rva l feeding behaviour (l29). From material col lected during the moth f l i g h t year in 1950 a clearer picture of the parasite complex began to emerge (33,114-). In 1952 the number of larva l instars was determined (120) and a precise sampling technique was perfected. The average number of larvae per branch t ip (including f ive years' needles) was accepted as the sample uni t ; four branch t ips were taken from two crown levels in each tree and the average number per branch t ip per tree calculated for each crown leve l (115,116). This system was established by sampling 4-0 trees at val ley bottom, 35 at the 750 foot elevation and 30 trees at the 1,500 foot elevation (from val ley bottom) on Cathedral Mountain in Yoho National Park and has since been tested in a l l sampling areas. For pract ica l purposes the lower crown samples could be eliminated and the measure of the outbreak taken as the average number of larvae per t ip in the more v i t a l upper crown. Since this technique was established i t has been modified s l i g h t l y . Samples are now taken from the upper third of the crown with an extension pole pruner (78) from several t rees, u n t i l the accuracy desired is obtained. Conversion of papulation estimates by this method to a per tree basis or per acre basis was made possible by the determination of the relationship of number of branch t ips per tree to tree height (33)* A sampling technique designed for rapid population estimates was derived on the sequential plan. In this system the outbreak was c lass i f i ed as l i g h t , medium-low, medium-high or heavy. Sampling was continued u n t i l the estimate f e l l into one of the four classes (116,121). Ecological studies were made on the effects of solar radiation on temperatures within the needle minesyby Henson and Shepherd (42). Their results showed that orientation and aspect of the needle, extent of the mine, and the presence or absence of wind modified the heating effect of radiat ion. On clear nights the mine temperatures were below a i r temperature, on clouded nights s l igh t ly higher. At the highest radiation leve l mine temperatures exceed a i r temperatures by as much as 6.8 F . ° . In 1953, there was a reduction in populations throughout the outbreak areas. Intensive studies were commenced on the damage done by the lodgepole needle miner in collaboration with J . A . Cook. 13. Curves were prepared from which i t was possible to predict the amount of defoliation expected from a given needle miner popu-lation. Increment and growth studies based on the work of Duff and Nolan (27) were begun by Cook (84.). In 1954- the l i f e history and distribution data were Integ-rated and published (123) and a tentative theory was advanced for periods of high winter mortality observed since 194.8 (4-3). Beginning that year, a systematic approach to population dynamics was begun. The method proposed was that of the l i f e table - a systematic tabulation of a measureable portion of the population throughout i t s l i f e cycle (125). The studies presented i n this thesis cover the period from 1953 to 1957 during which the author was enrolled at the University of British Columbia. Distinct from the population dynamics studies to be presented, however, are two publications resulting from intensive growth studies• The f i r s t summarizes the damage done to lodgepole pine i n the cut-break area (128) and the second deals with c r i t i c a l growth studies of lodgepole pine i n the outbreak region (84.). These studies permit certain deductions to be made concerning past population history. Portions of the authors' summary of the paper on needle miner damage w i l l give some indication of the potential importance of this insect i n forests. "1. It was demonstrated that the lodgepole needle miner caused as much as 80 per cent defoliation of lodge-pole pine in Banff National Park. The defoliation did not, except in a few overmature trees, reach 100 per cent. The period of greatest defoliation probably occurred from 194-0 to 1944. Since 1945 there has been a decrease i n defoliation varying i n location with differences i n needle miner populations, u n t i l at the present time (1956), maximum defoliation i n any part of the outbreak is not above 40 per oent. Had the outbreak continued at levels present from 1940 to 1944, mortality of trees would have occurred, probably by I960. 2. A method is presented whereby i t is possible to estimate percentage defo l ia t ion, expected tree mortality and approximate time to tree mortality from population e s t i -mates obtained by survey sampling. 3. Terminal and la tera l growth were s igni f icant ly reduced by a defol iat ion of 40 per cent with negligible reduction at 10 per cent. 6. Analysis of many (tree r ings) in the oourse of these studies c lear ly showed a lag in effect of defol iat ion on increment between the upper and lower bole," 2. LIFE TABLES FOR THE LODGEPOLE NEEDLE MINER 2.1. Review of l i f e tab les , in entomology The importance of l i f e tables in epidemiological studies has beoome increasingly important since the review by Pearl and Miner (94). Their recognition of the necessity of following a s t a t i s t i c a l l y acceptable cohort of an organism throughout i t s l i f e cycle to achieve this was perhaps the beginning of the intense interest which has since been generated. The extensive review by Deevey (24) focussed ecologists' attention on the transi t ion of demographic l i f e tables to ecological l i f e tables. The l imi t ing factor to their use in ecology has been the development of suitable techniques for measuring natural populations. Prior to Deevey18 review the use of l i f e tables was recognized in few textbooks. Leopold (66) gave a comprehensive review of ecological l i f e tables which he oalled " l i f e equations". Bodenheimer (9) discussed the concept at some length for insects as well as mammals and b i rds . Since Deevey's review at least three major ecological texts have devoted considerable space to discussions of l i f e tables. These 15. are: A l l ee , Emerson, Park, Park and Schmidt (2), Odum (92), and Andrewartha and Birch (3). The use of l i f e tables in forest insect population dynamics has lagged behind that for other organisms. Again, this i s largely due to the lack of suitable sampling techniques. A form of l i f e table for insect populations has been in use in Europe since at least the 1930's. Schwerdtfeger, In h is concise text "Fundamentals of Forest Pathology" presents what i s i n t r i n s i c a l l y a l i f e table: that i s , the graphic presentation of the "Intracyclic change" for a generation of PanoUa flammea Schi f f ((Panoils griseovarlegata (Goeze), pine beauty, of Varley (14.6)). These are l inked for many successive generations to compile "Gradation" or infestat ion curves (l02). Varley, in 1947, published his intensive studies on the Knapweed g a l l - f l y where he followed the reduction of popu-lat ions as a result of natural mortality factors . His sampling periods were at short intervals and he was able to describe the action of natural control factors in some deta i l (145). His methods were essential ly the l i f e table approach. Morris and M i l l e r , in 1954-j presented the f i r s t detailed example in North America of a l i f e table designed spec i f ica l ly for a forest insect , the spruce budworm (81). This excellent paper introduced a series which follows the development of techniques with the objectives of compiling l i f e tables for long-term studies of population dynamics. When completed this series should stand as a model for future workers for a long time to come. 1 6 . 2.2. The preparation of l i f e tables. The l i f e table is a continuous quantitative record of the abundance of an insect throughout i ts l i f e and as such is de-pendent upon suitable techniques for measuring changes in popu-lat ion l e v e l . It is of fundamental importance in the study of population dynamics but only i f records are kept for successive generations and the mortality factors determined. The method used in formulating l i f e tables for the lodgepole needle miner is that used in standard demographic tables but with the inclusion of a column for l i s t i n g and separating mortality factors and the elimination of the column giving " l i f e expect-ation" (81). Andrewartha and Birch (3) define the l i f e table (Ix table) as: "the age-schedule of surv iva l . For any part icular age-group of pivotal age x , lx is the proportion of individuals a l ive at the beginning of the age- interval ." However, l i f e tables as are commonly used also include the i r "dx table", or that giving the age-schedule of mortality. The terms and columns used are best defined by Allee et a l . (2). "Conventionally, a l i f e table is a series of columns eaoh of which describes something about the mortality relations within a population when ages of the components are taken into account. Conventionally a lso , a l i f e table starts with a certain sized group. . . .a t i ts time of b i r th and tabulates the events to whioh that cohort is subjected This tabulation takes the following form: x - Age in appropriate units stated as an in terva l . l x - the number surviving at the beginning of the age interval stated in the x column. dx - the number dying within the age interval stated in the x column. qx - the number dying in the age interval divided by the number of survivors at the beginning of the i n t e r v a l s . The rate of mortality. 17. ex - l i f e expectation. Mean length of l i f e remaining to each organism al ive at the beginning of the age inter -v a l . " In demographic l i f e tables age intervals are generally equal, beginning at age zero or b i r t h . In some cases age at f e r t i l i z -ation is considered (24) but this obviously is often impractical in studies other than demographic. In forest entomology the egg stage is generally considered as age zero. A lso , with insects i t is often impractical to sample at equal intervals because i t may not be possible to sample certain l i f e stages in the same manner as others and sampling may be prohibited by seasonal be-haviour. Therefore age intervals appearing in the x column generally represent major stages in the l i f e cycle or "susceptible stages" These w i l l be described in greater de ta i l in the fol low-ing sect ion. The or igin of the l i f e table conventionally starts with a certain-sized group, usually 100,000 or 1,000 (2 ,24) , In forest entomology, an important objective is to determine how mortality factors are affected by population density changes of the insect from generation to generation. Therefore, as recommended by Morris and Mi l ler (81), Ix is given as an aotual population estimate, the number of individuals per f ive-year branch t i p (117). When popul-ations are low this necessitates the use of fractions so that for convenience the figure ia taken as 100 Ix or the number of larvae per 100 t i p s . Simi lar ly qx is usually expressed as rate per 1,000 popul-at ion or 1,000 qx (24) whereas for our purposes i t i s more Convenient to use 100 qx, i . e . , percentage mortality (81). No I 18 Immediate value for inclusion of the " l i f e expectation 'ex' column" in needle miner work could be seen and so was discarded. The value of a column describing mortality factors respons-ib le for the corresponding mortality (dx) i s obvious (81) and so i s incorporated in needle miner l i f e tables. 2.3 Irtfe tables for the lodgepole needle nftn.gr Graham (31) pointed out the su i tab i l i t y of the needle miner as an Insect for c r i t i c a l population studies. He recognized a number of "crucial t r i a l s " through which any Inseot must pass i f i t i s to survive and by implication "succeed" in the sense of great abundance. For the needle miner he mentions the fol lowing"tr ials"* (a) The obtaining of adequate food reserves for the maturing larvae (b) Successful pupation and oogenesis (o) Successful f e r t i l i z a t i o n in the adult stage (d) Oviposition (e) Eclosion (f) Establishment in the needles (g) Larval growth from establishment in the f a l l of an even year to the spring of the next even year. Each of these t r i a l s includes many environmental factors which may affect the insect at that part icular stage. A discussion of these must wait for a la ter sect ion. However, the re lat ion of the stages l i s t e d above to the sampling stages decided upon merits some discussion here. The sampling stages chosen for the needle miner were based upon pract ical considerations as well as theoret ica l . They were chosen with a view to obtaining as much c r i t i c a l information as 19. possible within the l imits of pract ica l i ty . The development of sampling techniques for the needle miner was not faced with as many d i f f i c u l t i e s as are many other forest insects, l i ke the larch sawfly (142) or the spruce budworm (79). The former presents the d i f f i c u l t y that the pupal stage is spent in a dif ferent sampling universe than the larvae; the l a t t e r , among others, that the larvae are subject to wide-spread d ispersa l . Eggs of the needle miner are deposited in o l d , mined needles (103,123). The larva spends i ts whole existence within three mined needles except for the short time taken for transferring from one needle to another, and pupates within the last mine. Consequently, the only truly mobile or dispersable stage of the needle miner is in the moth stage. The sampling intervals for needle miner l i f e tables are i l lus t ra ted in Figure 2 along with a schematic i l l us t ra t ion of the l i f e cycle . Described in d e t a i l , they are: X - l The egg stage. The distr ibut ion of needle miner eggs in lodgepole pine trees is similar to that of the larvae (126). Oviposition is greatest in the upper crown, least in the lower crown. Upper crown samples give the maximum population estimate for a tree, mid-crown samples a reasonable average (117). V a r i -a b i l i t y between trees arises from differences in crown length and Stand density. The larva upon hatching usually mines a needle in the same t i p i f avai lable, within a few hours of eclosion. For egg and l a r v a l sampling therefore, mid-crown samples are taken where time does aiot permit sampling a l l crown leve ls . Distr ibut ion of the eggs wi th in oviposit ion si tes (empty mined needles) is re la t ive ly oonstant for any one area and the f ive-year branch t ip samples are there-fore convertible to a "per tree" basis and from that to a "per acre" basis i f required (33). The time of sampling of this stage is naturally dependent upon completion of oviposit ion. Generally, this is about mid-August, but as the weather during the summer affects both moth emergence and oviposition the progress of these events must be checked be-fore intensive sampling is done. Hatching usually commences with-in one to three weeks after the completion of oviposit ion, leaving a l imited period to make the sample. The sampling technique is necessarily slow and painstaking. Careful dissection of the needles is required for an accurate egg count. They are usually l a id within a few millimetres of the moth emergence hole (103) (Figure 1) but with disturbance frequently f a l l to lower levels in the needle. The eggs are large enough to be counted by the naked eye or with a low-power hand lens. Sampling is usually continued on a per t i p basis u n t i l a mean number per f ive-year-branch t i p within set error l imits is reached (standard error less than 10 per cent of the mean). Egg sampling was attempted on a large scale for the f i r s t time in 1954* Three areas were sampled, Mount Eisenhower (5,400 fee t ) , Brewster Creek and Baker Creek on four separate days, August 9, 10, 17 and 19th. On the las t date the three samples were taken at the same time of day by three separate workers from the mid-crown of the trees. A tota l of 1500 mined needles was opened and eggs counted. Eggs were la id singly and in bunches up to as many as 23 in one needle. One needle contained 46 eggs in two c lusters . Table 1 presents a summary of these data (124). 21. TABLE 1 EGG SAMPLING - 1954 Mean number needles Elevation containing eggs (in groups of 50; Mean nunH ber eggs per needle Area Mount Eisenhower Brewster Creek Baker Creek 5,400 15.8 5,700 12.2 6,000 U . 8 4.95 3.97 2.79 No s igni f icant difference between means for the u t i l i za t ion of oviposition sites (empty, mined needles) was found. As popul-ations pr ior to moth f l i gh t were comparable, this was to be expected. However, the mean number of eggs l a id at Baker Creek was s i g n i f i -cantly less than on the other two areas. In 1956, egg sampling was carried out in the four areas chosen for continuous l i f e table sampling, Mount Eisenhower, Mount Girouard, Massive Mt. and Cathedral Mt. i n Yoho National Park. Three hundred branch t ips from mid-crown of several trees in each looal i ty were examined. Table II presents a summary of the results of this examin-at ion. TABLE II EGG SAMPLING - 1956 Total No. Number Total Average Average Area Kiev- Mined contain- No.eggs per per ation Needles ing eggs found needle t i p Mt. Eisenhower 5,400 3423 441 1133 2.6 3.8 Girouard 6,000 2287 390 1151 3.0 3.8 Massive 5,500 1380 105 228 2.2 0.8 Cathedral 4,700 953 42 89 2.1 0.3 2 2 . Some interesting comparisons may be made between the res u l t s of t h i s and the 1954- sample but as these have a bearing on natural mortality and epidemiology, discussion w i l l be l e f t to a l a t e r section. The estimates made i n 1956 were on a comparable basis to the f i r s t l a r v a l examination made i n September, 1956. These are compared i n Table I I I . TABLE I I I COMPARISON OF EGG SAMPLING AND FIRST LARVAL ESTABLISHMENT 1956 Average number of eggs Average number of Area per t i p (Table I I ) larvae per t i p Eisenhower 3.8 3.1 Girouard 3.8 4-.0 Massive 0.8 1.9 Cathedral 0.3 0.6 In two areas therefore, established l a r v a l populations were double those expected from the egg sampling. Two explanations bear-ing equal weight are responsible f o r t h i s and, fortunately, both are susceptible to correction i n the future. (a) Time did not permit the usual practice of sampling u n t i l estimates were equally accurate and sampling was therefore l i m i t e d to 300 t i p s per area. At the Massive and Cathedral stations populations are much lower than i n the other two areas and these areas should have been more intensively sampled. (b) During the egg sampling period a tota l of f ive assistants were used. These included one other research o f f i ce r , one laboratory technician and three student assistants. This variety of help was at i ts peak when sampling the two areas, Massive and Yoho. In an attempt to ascertain the source of the error involved an analysis of variance was made comparing the workers' e f f ic iency. It was found that two of the workers were unsuited for the c r i t i c a l work of f inding and counting eggs. The combination of these two sources of error thus led to low estimates for the two areas and must be taken into account prior to future egg sampling. The occurrence of this error does not permit any speculation concerning the progress of the population from oviposition to la rva l establishment. More time and space have been devoted to describing this sampling interval as the results of this work have not yet been published in d e t a i l . X-2. F i r s t l a rva l sample. This sample Is taken in Late September or early October and can be done without dissection of the needles. The new mines are thread-like and careful examination of the tips is necessary for accuracy. Entrance into the f i r s t needle is almost invariably in the d i s t a l quarter of the needle and from the curved surface. The needle epidermis over the fresh mine is a pale green in contrast to the dark green of the unmined portion of the needle. Often the larva is v i s i b l e under the epidermis of 24 the mined portion* Unsuccessful mines are also detectable with-out dissection but these are usually checked. The larva present may be f i r s t or second instar depending on the time of sample; ecdysis usually occurs before winter hibernation. The advantage of sampling at this time is that i t determines the established start ing population and the loss of needle miner larvae between eclosion and establishment. New mines per branch t i p are counted from as many t ips as necessary to achieve the desired accuracy. This procedure is followed for a l l l a rva l and pupal sampling. X-3. Second la rva l sample. This is done after diapause is broken the following spring and fresh feeding is noticeable. Past studies have indicated that winter mortality is one of the major factors in population reduction (4-3,123). If the sample i s l e f t too late the larvae k i l l e d during the winter period dry and shr ive l and are hard to f i n d . A f a i r l y accurate estimate can be made by counting the mines showing fresh feeding and obtaining the mortality estimate by subtraction from the f i r s t la rva l sample (X-2). However, i f time permits, greater accuracy is obtained by actual counts of dead larvae. This permits a precise check of the previous autumn's sampling. A portion of the dead larvae are examined for incidence of disease. This work is now performed by Miss M.E.P. Cumming of the Calgary Laboratory. Disease examinations pr ior to 1954 were made by members of the staf f of the Forest Insect Disease Laboratory, Sault Ste. Marie, Ontario. X - 4 . Third la rva l sample. This sample determines the population which w i l l hibernate during the second winter. Again, i t is possible to do this without dissection of the needles by In-specting and counting the fresh mines. Third-and fourth-instar larvae are present in the autumn of the second year and are large enough to be seen in the mined needle. The population reduction over the summer can be determined by substraction from the spring sample (X-3). Mines in which the needle miner have been k i l l ed during the summer are character is t ic , the pale green re -sul t ing from feeding being absent and previously mined portions drying to a straw color . As long as the larva l i v e s , fresh feed-ing can always be distinguished. Experience has shown that this sample may be omitted without much loss in accuracy. Mortality during the summer of the la rva l years, even though a transfer from the f i r s t to the second needle is effected in mid-summer, has always been extremely low. X-5 . Fourth la rva l sample. This is an extremely important sampling period as i t shows the second winter's mortality and the la rva l parasite complex. Sampling must be done after mid-May as prior to this the parasites have not developed suf f ic ient ly to be v i s i b l e . Mass insectary rearings are necessary to check parasitism estimates and to obtain pertinent information on para-s i te biologies. Parasitism is not generally evident prior to this time except by la rva l sectioning or dissect ion. Again, a portion of the larvae found dead is examined for incidence of disease. 26. X-6. Pupal sampling. After moth emergence, careful examin-ation of pupal cases w i l l y ie ld information on pupal parasitism, mortality from other causes, moth population and the sex ra t io . However, i t is much simpler and more in keeping with the principles of l i f e tables to obtain this information from mass insectary rearings and to check these results with l imited f i e l d samples. It is possible at the time of the last la rva l sample (X-5) to dif ferentiate the status of each miner examined without undue disturbance of the larva. Once recorded for the purpose of that sample, the specimens are then placed in a f i e l d insectary for rearing. F ie ld checks are necessary to avoid any a r t i f i c i a l effect which may arise from insectary rearings. These sampling intervals , one egg, four la rva l and one pupal, are the ones deemed suitable to assess the course of the popul-ation of a single generation from the time of ovipositIon to moth emergence of the needle miner. The emphasis on sampling of the la rva l stage is understandable from the l i f e cycle . Approxi-mately 89 per cent of the l i f e cycle of the needle miner i s spent in the la rva l stage, covering the most adverse cl imatic periods, 4*3 per cent is spent in the pupal stage, 4.3 per cent in the moth stage and s l ight ly more than two per cent in the egg stage. Whenever possible, extra samples are taken, as the oftener the population is sampled the more information on population dynamics is obtained. However, these facts do not detract from the importance of the pupal, moth and egg stages in population ecology. In many, perhaps most instances, one of these may be the determining factor in population r ise and f a l l . To f u l l y understand long-term fluctuations of the population in succeeding generations i t is necessary to l ink together the l i f e tables of eaoh gener-ation through the reproductive stage. The comparison of established la rva l populations w i l l give the trend of the out-break but w i l l not y ie ld any information on reductions which occur in the c r i t i c a l reproductive stage. The adult stage can-not be sampled by any of the means described and other study methods must be used. Two methods have been used in attempts to determine moth fecundity: controlled matings and moth dissect ions. The con-ditions for successful mating of needle miner moths have never been met as attempts to mate moths in capt ivi ty have been singularly unsuccessful. Perhaps f l i gh t is necessary pr ior to copulation. Moth dissections were made in 1950, 1954» and 1956. When the 1956 results of moth dissections were compared with actual numbers of eggs found and larvae established the counts by dissection were found to be too inaccurate to use. Unt i l the problem of mating the moths in capt iv i ty and inducing them to lay their eggs can be surmounted we can only speculate from observations made by dissection and in the f i e l d on oviposit ion behaviour. Indications are that there has been a reduction in fecundity, at least since the 1954- generation. The evidence for t h i s , and implications of i t , w i l l be discussed in the natural controls sect ion. 2.4 Examples of needle miner l i f e tables. L i fe tables are presented for needle miner populations from four loca l i t i es in the outbreak area. These are. (a) Mount Eisenhower, Banff National Park. The sample area is located 22 miles northwest of the town of Banff, Alberta, on the east side of the Bow Val ley . Two alt i tude levels are represented: Valley bottom (4,800 feet a . s . l . ) and 5,400 feet . Par t ia l l i f e tables based on actual samples are presented for three generations for the val ley bottom and four for the higher elevation. Only the tables for the 1954-1956 generation follow the system de-scribed for their formulation. (b) Massive Range, Banff National Park. This area is located about eleven miles northwest of Banff on the west side of the Bow Val ley. The alt i tude leve l sampled is approximately 5|500 feet . Only the l i f e table for the 1954-1956 generation is presented. (c) Mount Girouard, Banff National Park. This area i s located about nine miles northeast of Banff on the south side of Lake Minnewanka para l le l to the open-ing of the Cascade Val ley. The elevation i s approxi-mately 6,000 feet . Only the 1954-1956 generation is presented. (d) The Kickinghorse Pass, Yoho National Park. This area is located on a north-faoing slope of Cathedral Mountain about 250 feet above val ley bottom 29. (elevation 4,700 feet a.s.l.). Only the 1954-1956 generation is tabulated. Example 1. Mount Eisenhower - valley bottom. These are presented in Tables IV to VT. Life tables for the 1948-1950 (Table IV) and the 1952-1954 (Table V) generations are incomplete but the sample periods are so timed as to show the major mortality factors. The l i f e table for the 1954-1956 generation (Table VI) shows that there was no measurable population after the winter of 1955-1956. The number of larvae present per 100 tips in the ix column is based upon actual samples statistically evaluated and indicates estimated numbers within plus or minus 10 per cent of the mean (117). The established larval populations following the 1948-1950 and the 1952-1954 generations agree fairly well with the expected number of eggs. What is demonstrated clearly by these tables is the existence of three periods, when natural control factors may drastically reduce the larval population. These are the two overwintering periods and the spring of the last year of the l i f e cycle when parasitism becomes effective. The winter of 1949-50 accounted for 78 per cent of the population (Table IV). In the winter of 1953-54, between December 16 and February 5, almost 95 per cent of the population was killed (Table V). During the winter of 1955-56 the whole measureable population died (Table VI). The data from valley bottom show that climate (the specific effect will be described in the following section) during a l l three generations represented, was the factor which in effect "controlled" the established population in this area from 1948 to 1956. 30. The l i f e table lends i t s e l f readily to graphic presen-ta t ion . Usually three graphs are derived from the table! the survivorship (lx) curve, the death (dx) curve, and the death-rate (qx) curve (24). As the death curve i s merely the complement of the survivorship curve i t s value i s questionable and in this presentation only the survivorship and death-rate curves are given for each area. The death-rate curve emphasizes the periods of high mortality with respect to the l i f e stage affected. An addition to the conventional death-rate curve i s the bar to the r ight of the curve. The so l id portion represents the total mortality of the start ing population. Survivorship and death-rate curves are presented for the 1952-1954 and 1954.-1956 generations from Mount Eisenhower, val ley bottom in Figures 5 and 6. Figure 5 also presents the curve for the 1952-1954- generation from the 5,400 foot elevation. TABLE IV LIFE TABLE FOR THE 1948-1950 GENERATION OF NEEDLE MINER MOUNT EISENHOWER - VALLEY BOTTOM x Ix dxF dx lOOqx Sept. 1948 - June, I and II instar June 4 , 1949 III and IV instars May 3, 1950 IV and V instare Pupae-no sample, assumed emerged Moths SR 50:50 GENERATION 3013 99.93 Expected no. eggs per 100 tips - 24 Actual not measured. Established larval population per 100 tips - 25 Population trend (larvae) - 0.856 1949 3015 Climate 932 30.91 2083 Climate 1622 77.87 Parasites 4,59 22.03 2081 99T87 2 2 M F 1 1 32. TABLE V LIFE TABLE FOR THE 1952-1954 GENERATION OF NEEDLE MINER MOUNT EISENHOWER - VALLEY BOTTOM lx dxF dx lOOqx From ecloaion to May 12, 505 1953. I and II instara . May 12, 1953. II,III and 479 IV instara. Deo. 16, 1953. I l l and IV 458 instars . Feb. 5, 1954. H I and IV 24 instars. June 2, 1954. V instar 5 and pupae. Moths SR 50:50 M 2.5 Climate Climate Climate Climate F 2,5 GENERATION 26 21 434 19 5.2 4.3 94.69 79.17 500 99.01 Expected eggs per 100 t ips - 60 Actual not measured Established la rva l population s 98 Population trend (larvae) - 19.40* Figure 5. Survivorship and death-rate curves for Mount Eisenhower, val ley bottom U , 8 0 0 ' ) and 5,4-00' for the 1952-54 gener-ation of needle miner. ^ 15 A S O N D J F M A M S O N D J F M A M J J 1952 1953 1954 TABLE VI LIFE TABLE FOR THE 1954-1956 GENERATION OF NEEDLE MINER MOUNT EISENHOWER - VALLEY BOTTOM Ix dxF dx lOOqx Eclosion to Dec. 13, 1954 I and II instars 98 Dec. 13, 1954. II instar 15 Summer, 1955 8 June 26, 1956 0 GENERATION  Climate 83 84.23 Climate 7 46.67 Climate 8 100.00 98 100% Expected number of eggs - none Figure 6» Survivorship and death-rate curves, 1954.-56 generation. Mount Eisenhower, valley bottom (4,,800') A S O N D J F M A M S O N D J F M A M J 1954 1955 1956 TABLE VII LIFE TABLE FOR THE 1948-1950 GENERATION OF NEEDLE MINER MOUNT EISENHOWER - 5,400 FOOT ELEVATION dxF dx lOOqx Eclosion to 1949. I , I I 3990 and I I I i n s t a r s . Climate 927 23.23 June 1, 1949. I l l , IV 3063 and V ins tars Climate 2199 Parasites 450 2649 71.80 t*68 48 8o\ Pupae Moths SR 50:50 414 M F 207 207 GENERATION .1576 89,62 Expected no. eggs = 4968 Actual not measured Established l a r v a l population - 2500 Population trend (larvae) - 62.75? Figure 7. Survivorship and death-rate curves, 1954-56 generation. Mount Eisenhower, 5,400'. A S O N D J F M A S O N D J F M A M J J 1954 1955 1956 TABLE VIII LIFE TABLE FOR THE 1950-1952 GENERATION OF NEEDLE MINER MOUNT EISENHOWER - 5,400 FOOT EIEVATION lx dxF dx lOOqx Eclosion to summer, 1951 2500 Climate 500 '0.00 I, II and III instars. Summer 1951 to spring 2000 Climate 315 15.77 1952. I l l , IV and V Parasites 85 A.23 ins tars 400 20.00 Spring, 1952 1600 Pupae, no sample, assumed emerged. Moths SR 50:50 M F 800 800 GENERATION 900 36.00 Expected number of eggs - 19,200 Actual not measured Established larva l population - 2709 Population trend (larvae) -. 108.0$ 36, TABLE H LIFE TABLE FOR THE 1952-1954 GENERATION OF NEEDLE MINER MOUNT EISENHOWER - 5,400 FOOT ELEVATION dxF dx lOOqx Eclosion to summer, 1953 2709 I, II and III instars June 9, 1953. II, III 2557 and IV instars Dec. 16, 1953. I l l and 2338 IV ins tars Feb. 2, 1954. H I and TV 1734 instars . May 25, 1954 Pupae not sampled, assumed emerged. Moths SR 50:50 GENERATION _ 1250 M 625 F 625 Climate Climate Climate Climate Parasites 152 219 604 248 236 484 5.6 8.55 25.81 14.30 13.60 27.90 1359 50,17 Expected number of eggs = 15,000 Actual number of eggs 4,700 Established larva l population - 1114 Population trend (larvae) a 41.1% 37. TABLE X LIFE TABLE FOR THE 1954-1956 GENERATION OF NEEDLE MINER MOUNT EISENHOWER - 5,400 FOOT LEVEL X l x dxF dx lOOqx X - l -eggs 4700 Needle drop 3586 & unknown 76.30 X-2 Instars I and I I 1114 Climate 409 36.68 Extra- Deo. 14, 1954 I I i n s t a r 705 Climate (winter (spring) 219 JZP_ 289 31.06 40.97 X-3 July 1, 1955 X-4. I l l and TV i n stars 416 Climate 143 Parasitism 142 Unknown 8 293 34.37 34.09 1.98 70.44 X-5 IV and V instars 123 Parasites 18 14.55 X-6 Pupae 105 Unknown 26 Parasites less than X 26 + 24.76 O.A«i 25.21 Emerged 79 Moths SR ^8:52 M F 38 41 GENERATION 4621 98.32 Expected number of eggs = unknown Actual number of eggs = 378 Established l a r v a l population = 308 Population trend (egj *s) = 8.04$ 38. Example 2. Mount Eisenhower, elevation 5.4.00 feet. This example is more complete than the first, as, following the severe winter of 194-9-50 when it appeared that the higher elevations offered "refuge" areas for the needle miner (4-3) sampling was continued more intensively there than at the lower level. The life tables for this location are presented in Tables VII to X for the 1948-1950, 1950^ 1952, 1952-1954 and 1954-1956 generations. Survivorship and death-rate curves for the two later generations are presented in Figures 5 and 7. The first three tables also demonstrate the importance of the two winter periods and of parasitism in the final year of the life oycle. The egg sampling, begun in 1954- and repeated in 1956 (Tables LX and X)indicates possibly two additional periods in the life cycle where natural control factors may effect drastic re-ductions in the egg stage. These are: between egg formation and actual oviposition and/or between oviposition and establishment of the larvae in the needles. Again, climate is the predominant controlling factor, para-sitism remaining low throughout the four generations indicating that the parasites are equally or slightly more susceptible to adverse climatic factors. Table XI presents the percentage para-sitism based on the established larval population for the four generations (Tables VII - X). The heaviest larval k i l l was during the winter of 194-9-50 and from the above this would appear to be equally true for parasites. Since that time however, where a residual larval population has remained, parasitism has increased. This will be discussed in greater detail In a later section. 39. TABLE XI PERCENTAGE PARASITISM OF THE TOTAL ESTABLISHED LARVAL POPULATION Year Percentage Parasitism 1948-50 10.13 1950-52 3.40 1952-54 8.71 1954-56 U.36 Example 3. Massive Range. Banff National Park A l i f e table is presented for the 1954-56 generation only (Table XII). The survivorship and death-rate curves are presented in Figure 8. Example A. Mount Girouard. Banff National Park A l i f e table is presented for the 1954-56 generation only (Table XIII). Survivorship and death-rate curves are presented in Figure 9. AC-TABLE X I I LIFE TABLE FOR THE 1954-1956 GENERATION OF NEEDLE MINER MASSIVE RANGE - 5500 FEET X l x dxF dx lOOqx X - l Not measured X-2 F a l l , 1954 1257 Climate I and I I instar (winter) 737 58.60 (spring) _55_ ,4T38. 792 62.98 X-3 Summer, 1955 465 Climate X-4 I I I and IV i n s t a r (winter) 94 20.22 (spring) 2 0.43 Parasitism 123 26.45 Bird predation 124. 26.67 343 73.77 X-5 Spring, 1956 IV and V i n s t a r 122 X-6 Pupae 122 Climate 30 24.84 Moths SR 45:55 92 M F 41 51 GENERATION 1165 92.68 Expected number of eggs — unknown Actual measured number - 76 Established l a r v a l population - 190 Population trend (larvae) s 15.11* Figure 8. Survivorship and death-rate curves, 1954.-56 generation. Massive Range, 5,500'. 1500 MASSIVE RANGE 5500 FEET A S O N D J F M A - S O N D J F M A M J J 1954 1955 1956 TABLE X I I I LIFE TABLE FOR THE 1954.-1956 GENERATION OF NEEDLE MINER MOUNT GIROUARD - 6000 FEET l x dxF dx lOOqx X - l Not measured X-2 F a l l , 1954 2633 I and I I ins t a r X-3 Summer, 1955 896 X-4, I I I and IV instars X-5 June 4-, 1956 V and pupae X-6 Pupae Moths SR 47:53 GENERATION 270 263 205 M F 96 109 Climate (winter) (spring) Climate (winter) (spring) Parasitism Bird Predation Unknown Larval paras-i t i s m Climate Parasitism 1593 144 1737 60.50 .5*4$, 65.96 213 15 281 76 626" 23.77 1.67 31.39 8 .50 4*60. 69.87 56 2 58 2.59 21.29 7r60 28.89 2A28 92.21 Expected number of eggs - unknown Actual measured number - 385 Established l a r v a l population - 395 Population trend (larvae) - 14.6055 Figure 9. Survivorship and death-rate curves, 1954.-56 generation. Mount Girouard, 6,000'. 42. TABLE XIV LIFE TABLE FOR THE 1954-1956 GENERATION OF NEEDLE MINER CATHEDRAL MOUNTAIN - £700 FEET  lx dxF dx lOOqx X - l Not measured X-2 F a l l , 1954 924 I and II instar X-3 Summer, 1955 181 X-4 III and IV instar X-5 May 27, 1956 V instar X-6 Pupae Moths SR 51*49 GENERATION M 21 53 42 42 F 21 Climate (winter) (spring) Climate (winter) (spring) Parasitism Parasitism 687 743 111 6 11 128 74.4 6.02 80.42 61.33 3.31 6.08 70.72 11 20.75 882 95.45 Expected number of eggs — unknown Actual measured number = 30 Established la rva l population - 60 Population trend (larvae) 6.49# Figure 10. Survivorship and death-rate curves. 1954-56 generation. Cathedral Mountain, 4 ,700 f . 1000 A S O N D J F M A - S O N D J F M A M J J 43. Example 5. Cathedral Mountain. Yoho National Park. As i n the two preceding examples only the 1954-56 generation i s presented. The data are given i n Table XIV and Figure 10. The l i f e tables given above present a tabulated record of mortality and survivorship from four l o c a l i t i e s and two a l t i t u d e l e v e l s . Additional data are available from other areas and al t i t u d e levels but these were taken i n a manner unsuited to presentation i n l i f e tables. However, they w i l l be referred to i n the discussion of natural control factors. The l i f e tables show c l e a r l y that there are four and possibly f i v e periods i n the two-year l i f e cycle of the lodgepole needle miner during which extensive mortality probably occurs: (1) between egg formation and oviposition. (2) between oviposition and l a r v a l establishment. (3) during the f i r s t l a r v a l hibernation. (4) during the second l a r v a l hibernation, and ( 5 ) during the spring of moth emergence. 3. DISCUSSION OF NATURAL CONTROL FACTORS 3.1 Climatic factors 3.1.1 Winter mortality Larval mortality caused by extreme winter conditions i s with-out a doubt the primary cause of the decline i n populations of the lodgepole needle miner since 1944* As was stated e a r l i e r , the outbreak was very heavy i n 1942 and continued to spread and generally increase (45-50). The f i r s t major check was observed i n 1946 (51) but populations were s t i l l high u n t i l the winter of 44. 1949-50 (53,113). Following this year the populations have re-treated to the area where they were f i r s t noted, at the inter-mediate levels on the mountain slopes, and at much lower levels of abundance. This will be discussed in greater detail in a later section but mention here serves to introduce the discussion of the factor which caused this to come about. (See Figures 11 and 12). (l) Theories of cold resistance and death in insects. Our knowledge of the process of insect death by cold temper-ature is far from complete but certain fairly well substantiated conclusions may be made. Uvarov (144) summarized the early work and from his review came to several conclusions: (a) There is a seasonal variability in cold resistance, the greatest resistance being found in winter and least in summer. (b) Cold-hardiness of an insect is affected by desiccation, starvation, type of food, developmental stage and sex. (c) Degree of cold-resistance depends on the balance of easily freezable water and fat present in the body and is not therefore constant even within individuals of a species. (d) Two problems are involved, designated as "quantity factor" (prolonged moderate cold) and "quality factor" (degree of cold). (e) The lethal temperature is apparently dependent on velocity of cooling, stage of insect development and sex, physiological state of insect, repetition of cooling, Figure 11. Needle miner distr ibut ion pr ior to 1949-50. Horizontal l ines - heavy Vert ica l " - medium Crossed " - l i gh t Figure 12. Needle miner d ie i r ibuUc^- fc l lowing 194.9-50 I A5. time of exposure and the constitution of the body f l u i d s . More detailed works since Uvarov's review have ver i f ied the basic truth of the above conclusions and have added information regarding them. It i s now believed "that a very small port ion.of the insect fauna can withstand actual freezing of t issues. (2,28,99). However, the type of freezing, whether in t ra - or extra-ce l lu lar has an important bearing on the survival of most insects at low temperatures. Generally insects which hibernate in an active developmental stage can survive a high degree of extra-ce l lu lar freezing but in t ra -ce l lu la r freezing results in damage to organs and/or death (4.,98,99,100,10l). Various protective mechanisms against in t ra -ce l lu la r freezing are known. These i n -cludes dehydration and contraction of t issue ce l l s j hydrophilic col lo ids in condensed blood layers forming a mechanical barr ier against penetration of tissues by ice crysta ls : and the formation of gels r ich in hydrophilic col loids which depress the freezing point U»4l)» The temperatures above which such insects can survive are not constant for any one species, stage, or sex. Many factors , extr insic and i n t r i n s i c , have a modifying influence. The lowest le tha l l im i t that i t may be safe to generalize about l i e s between -AO°F to -50°F, or i t s equivalent in time x temperature (2,3). Limited tests of cold-hardiness made on lodgepole needle miner larvae in the laboratory indicate they are extremely cold-hardy, even in the immature stages. Cold temperature tests were made on f i r s t - i n s t a r needle miner in the f a l l of 1950 before establishment. It was determined that they could withstand temperatures of 21°F for periods up to 2A hours. Temperatures of 10 F caused about 25 per cent mortality of larvae in 24 hours. Temperatures of 0°F caused no mortality in 1 hour but almost 100 per cent mortality in 24 hours (113)• Further tests were conducted on larvae removed from the f i e l d in November, 1951. Examination of the gut disclosed no food material suggesting that feeding had ceased and the larva hibernates with no food in i t s ' gut. No s igni f icant mortality was observed at temperatures down to -8°F of 24 hours duration (125). Comparison of the mortality which has occurred in the. f i e l d with the low winter temperatures associated with periods of mortality gives a better conception of the a b i l i t y of the needle miner to res is t cold temperatures (see sections (3) and (4 ) )« (2) Winter temperatures as a control l ing factor in populations. Uvarov takes the extreme view that "the mortality of insects caused by winter cold is probably the main factor in control l ing the abundance of most insects in the temperate latitudes" and ci tes many examples to support his statement (144). This may be taken as the opposite end of the scale to the various "biot ic" theories (90). More r e a l i s t i c are the recent "comprehensive" theories (102,106) which allow that under certain circumstances any environ-mental factor may act as a check on insect abundance. It is the intention in this section to show, by example, that reduction of Insect abundance by winter temperatures is not un-common. A l l overwintering larvae of the brown-tail moth are reported o to have been destroyed by temperatures approximating -25 F in the 47. northeastern United States. Low winter temperatures are apparently a barrier to this insect 's northward spread (74-). Similar con-clusions have been reached for the European pine shoot moth in Michigan (5). Percentage mortality approached 100 per cent with exposure for 14 days to temperatures of ~4°F and increased pro-gressively with time of exposure. At -13°F mortality approached 100 per cent in 72 hours; at -22°F over hal f succumbed after an hour's exposure and a l l were dead after eight hours. Nolte (9l) reviewed the theories then current for control of the rape f lea beetle in Germany. It had been commonly accepted that extreme cold winters were the major control l ing factor . Nolte found that this was not always so. that i t was not always winter temper-atures. Low temperatures from mid-August on. and in March, were equally successful in preventing outbreaks of this agricultural pest . The insect population studied overwintered in every developmental stage which would contribute to different population effects from severe environmental factors . The Cal i fornia oak moth, Phrvganidea ca l i forn ica Pack., over-winters in an act ively feeding larva l stage. Exhaustion of available food supply Is the most common biot ic cause of mass mortality in oak moth populations; natural enemies, parasites and predators, are not usually the cause of crash declines in populations. Crash declines of Phrvganidea populations in central Ca l i forn ia usually are due to factors associated with severity of winter weather. Populations fluctuate only moderately in southern Cal i forn ia where the physical environment i s essential ly "benign". They are more violent in Central Cal i fornia and in northern Cal i forn ia the climate i s too severe 48. to allow more than temporary summer colonies to survive (40). This i s an example of an insect with apparently very narrow l imi ts of cold tolerance. Hutchinson (60) as well as several other authors (23.25,60) have shown that populations of some scale insects are dependent upon winter temperatures for their increase (or lack of i t ) . Thus the Cal i fornia Red Scale numbers remain low in areas of low temperature but increase in areas of mild winter temperatures result ing in large spring populations. Hutchinson found that population density of the Cal i fornia Red Scale was inversely correlated with the number of winter nights when temperatures dropped below freezing. In summary, i t appears that the temperature environment of insects , l i ke that of most organisms may be divided into several "zones". These are termed: "the zone of le thal high temperature; zone of favorable temperature; and zone of le thal low temperature." The zone of le thal low temperature may be further divided into two sub-zones: "the zone of freezing temperatures", ch ie f ly important in those species that spend some time in dormancy and "the zone of non-freezing le thal low temperatures" (3). The needle miner i s part icular ly subject to the zone of freezing temperatures as the larva l stage spends two winters of approximately f ive months duration when freezing conditions pers is t . (3) Winter mortality i n lodgepole needle miner populations. The importance of winter mortality as a control factor i s c lear ly shown in the l i f e tables presented ear l ier (Tables IV -XIV). Table XV shows the percentage winter mortality of the tota l 4 9 . population i n the two-year cycle. TABLE XV PERCENTAGE WINTER MORTALITY OF ESTABLISHED NEEDLE MINER POPULATIONS IN FOUR LOCATIONS Per Cent Mortality of t o t a l established population Area Generation 1948-50 1950-52 1952-54 1954-56 Mount Eisenhower 85 - 99 100 Valley bottom Mount Eisenhower 78 33 45 70 5,400' Girouard, 4,700' - 68 Cathedral, 6,000' - - 86 These figures of percentage mortality f o r the whole gener-ation based on the established l a r v a l population are more s i g n i f i c a n t , I f less s t r i k i n g , than those based on yearly estimates of the population present before the eff e c t took place. This i s another advantage of the l i f e table that such a comparison i s ea s i l y obtained. To determine the winters which have had the greatest e f f e c t yearly estimates are necessary. These are present-ed f o r a l l areas and estimates made from the beginning of the study i n Tables XVI to XXIII. The r e l i a b i l i t y of the estimates v a r i e s , those beginning with the 1948-49 estimates being s t a t i s t i c a l l y acceptable. The comments a r i s i n g from perusal of these tables are best dealt with together. TABLE XVI RECORDED WINTER MORTALITY MOUNT EISENHOWER, B 0v Valley, Banff National Park Year Alti t u d e 4.800 5300 5800 6300 194-3-44 9 1944-45 -no estlmate-1945-46 50 1946-47 1 1947-48 20 1948-49 31.65 27.58 20.04 23.92 1949-50 99.9 78.8 64.8 61.9 1950-51 low 25.0 1951-52 low 15.8 1952-53 5.2 5.6 1953-54 98.4 17.7 6.3 1.5 1954-55 91.84 48.1 67.6 1955-56 100.0 34.4 TABLE XVII RECORDED WINTER MORTALITY MASSIVE MOUNTAIN, Bow Valley, Banff National Park Altitude Y e a r 4600 5100 5600 6100 1948- 49 9.8 16.6 11.8 28.3 1949- 50 100.0 91.0 92.6 87.0 1950- 51 no estimate 1951- 52 " " 1952- 53 " " 1953- 54- 79.8 6.7 0.5 2.9 1954- 55 51.0 58.6 1955- 56 20.2 TABLE XVIII RECORDED WINTER MORTALITY CATHEDRAL MOUNTAIN, Kickinghorse Canyon, Yoho National Park . Alt i t u d e e a r 4500 5000 5500 1948- 49 14.0 14.3 44.6 1949- 50 24.6 43.4 30.8 1950- 51 21.4 43.2 12.6 1951- 52 2.4 1952- 53 5.3 1953- 54 11.2 7.0 3.7 1954- 55 74.4 1955- 56 61.3 TABLE X K RECORDED WINTER MORTALITY LAKE LOUISE, Bow Vall e y , Banff National Park Year Alti t u d e 5050 5550 6050 6550 1948-49 23.0 18.5 16.1 8.9 1949-50 100 61.1 41.3 63.4 1950-51 no estimate 1951-52 II n 1952-53 6.1 1953-54 99.5 15.8 17.4 6.8 1954-55 85.2 (90.3) 1955-56 no estimate TABLE XX RECORDED WINTER MORTALITY MOUNT NORQUAY (EDITH), Bow Vall e y , Banff National Park Year Alti t u d e 4700 5200 5700 6200 1948-49 34.8 17.7 13.4 17.8 1949-50 97.8 93.5 88.0 82.4 TABLE XXI RECORDED WINTER MORTALITY BANKHEAD, Bow Valley, Banff National Park Year Al t i t u d e 4800 5300 5800 6300 1948-49 23.9 16.9 7.0 38.9 1949-50 t 91.0 89.5 87.8 84.0 53. TABLE XXII RECORDED WINTER MORTALITY HAWK CREEK (SNOW CREEK) Kootenay National Park Altitude Year 4400 4900 5400 5800 1948-49 23.2 13.9 15.5 9.9 1949-50 56,9 75.8 81.2 60.7 1953-54 23.8 TABLE XXIII RECORDED WINTER MORTALITY MISCELLANEOUS AREAS - Banff National Park Year Location Alt i t u d e Mortality 1951-52 Baker Creek 6000 15.8 Stoney Creek 5500 6.9 Upper Cascade 5750 9.4 1953-54 Baker Creek 6000 1.7 Stoney Creek 5500 11.9 Brewster Creek 5200 8.9 Saskatchewan Crossing 4700 11.2 Eisenhower Junction 4676 91.3 1954-55 Baker Creek 6000 77.7 Stoney Creek 5500 73.7 Cascade Valley mouth 5500 73.4 Mount Coleman 5000 87.6 Brewster Creek 5700 83.2 Brewster Creek 5200 64.1 Mount Girouard 6000 60.5 1955-56 Mount Girouard 6000 23.73 The conclusion drawn from the above tables i s that f i v e winters since 1943 were s i g n i f i c a n t i n t h e i r effects on l a r v a l populations. These are: 1945-46; 1949-50; 1953-54; 1954-55; and 1955-56. 54. The regular occurrence of comparable results at s i m i l a r elevations led to the formulation of Table XXIV where mortality estimates of comparable elevations were combined. TABLE XXIV RECORDED WINTER MORTALITY BY ALTITUDE - ALL AREAS Year Alti t u d e up to 5000 5000 5500 6000 % Mort. N % Mort. N % Mort. N % Mort. N 1943-44 - 9.0 1 •» -1944-45 - SB -1945-46 50.0 1 «. m 1946-47 - 1.0 1 -1947-48 - 20.0 1 -1948-49 24.6 5 19.4 5 13.7 5 23.6 5 1949-50 97.7 5 82.8 5 74.9 5 75.7 5 1950-51 - 25.0 1 m mm 1951-52 - 11.3 2 12.6 2 -1952-53 5.6 2 5.6 1 - •at 1953-54 92.2 4 12.2 5 6.5 4 3.7 3 1954-55 89.7 2 65.9 6 73.0 6 -1955-56 100 1 27.3 2 23.7 1 -N = Number of mortality estimates at each a l t i t u d e . The data f o r four of the s i g n i f i c a n t years are plotted i n Figure 13. Averaged i n th i s way, the decreased mortality with increase i n elevation i s less s i g n i f i c a n t but i n years when heavy mortality does Figure 13. Mortality of lodgepole needle miner at d i f ferent elevations in four selected years. 100 55. not occur (1948-49 particularly) the decrease in mortality is slight and that at the highest elevation is greater than at valley bottom. (4) Climatic conditions causing winter mortality observed in  lodgepole needle miner populations. In 1954, i t was postulated that the weather of the coldest month of the winter was the major factor causing winter mortality of the lodgepole needle miner (43). This work compared the winters of 1948-49 and 1949-50 in relation to mean monthly temper-atures (November to March) and demonstrated that although mortality was far greater in 1949-50 than in 1948-49 (See tables XVI - XXII and Figs. 11-12) the mean monthly temperatures were higher in 1949-50 than I948-49 with one exception. January. The lower mortality observed at the Cathedral sampling area did not conform to the rest of the area and i t was pointed out that more complex relationships between air masses occur on the western side of the Great Divide separating the areas. The general conclusions of this study were: n l . In the main part of the Bow Valley, the vertical d i s t r i -bution of temperature is a function of the predominant type of general circulation. Thus during a month characterized by frequent, but rapid, invasions of polar continental air, the upper slopes are more often colder than the valley floor. On the other hand, stagnating air of any type produces extremes of cold on the valley floor, and these are much more severe when the air is of polar continental origin. (Invading maritime air warms the upper slopes so that effectively, the valley bottom is colder than the upper slopes at such times. Thus effects are produced that are similar to, but more moderate than, the effects of stagnating polar conti-nental air ) . 2. In the Bow Valley, winter mortality seems to be distributed in the same way as the zones of extreme cold that occur during the coldest winter month. Because the dominant type of circulation 56. during this month may be d i f f e r e n t i n d i f f e r e n t years, the zone of most extreme cold may occur either at v a l l e y bottom or at the tops of the slopes. Consetfiently, greatest mortality should occur i n either of these locations, although i t should occasionally be exceptionally great at v a l l e y bottom. Best s u r v i v a l should occur most consistently along the middle of the slopes. 3. This middle zone then, should constitute the most persistent reservoir f o r r e - i n f e s t a t i o n of the other zones and, therefore, the most serious attempts to control the insect should be concentrated i n i t . 4.. In or near the major passes that enter the Bow Valley from the west, the a i r flow i s more complicated and, therefore, the v e r t i c a l d i s t r i b u t i o n of mortality i s much less predictable than i t i s i n the rest of the v a l l e y . " Two major postulates were susceptible to t e s t i n g : that pronounced inversions i n the lower a i r were produced by stagnating cold a i r masses i n the Bow V a l l e y , and that the bulk of l a r v a l mortality occurred during the coldest month. During January, 1956, Fuess hygrothermographs i n Stevenson screens were set out at v a l l e y bottom and v a l l e y bottom + 750 feet on Mount Eisenhower and attended from January 2 to February 1A. Air-mass analyses f o r t h i s period were made by W.R. Henson from charts supplied by the Dominion Meteorological O f f i c e , Calgary (Appendix 9). Even i n t h i s short period temperature inversions were common. On January 7, a f t e r a day and a ha l f of cold polar continental a i r there was an inversion from 0700 to 1100 MST with a peak difference of 12 F°. This was due to a weak invasion of maritime polar a i r whose warming effect was f e l t f i r s t on the upper slopes, f i v e hours e a r l i e r than i n v a l l e y bottom. This i s reported to be a common phenomenon noticed by s k i i e r s i n this area. Leaving the town of Banff i n sub-zero weather they f i n d i t near the freezing point on the s k i slopes. The next inversion occurred on January 13th. Polar cont i -nental a i r had invaded the Bow Valley late on January 11th but as c i rculat ion was weak, the a i r mass could not displace the warm maritime a i r completely. This inversion undoubtedly resulted from the cold cP a i r displacing the warm mP a i r at val ley bottom f i r s t . Another short inversion occurred on January 18th when mP a i r entered the valley replacing cP a i r and in the same day i t s e l f was replaced by cP a i r . The cP a i r remained in the val ley from January 24th to February 2nd, becoming increasingly colder. The conditions approached those postulated by Henson et a l . (43) for 1949-50 but to a lesser degree. The nightly inversion began on January 28th and was repeated each night u n t i l February 1st . The effects on val ley bottom and slope temperatures are i l lus t ra ted in Figure 14 which is a tracing of the actual thermograph chart. Figure 15 shows a four-day period following the period i l lus t ra ted in Figure 14 after polar maritime a i r had f i n a l l y supplanted the colder polar continental a i r . The to ta l temperature difference from 0100 to 1100 on the three successive days showed val ley bottom to be 59,53, and 45 F ° colder than the upper slopes. Not only was the temperature more severe at val ley bottom, but It was much more var iable . The diurnal ranges for the three days at val ley bottom were: 9°F to -25°F ; 11°F to -23°F j 13°F to -18°F compared to: 5°F to -19°F j 8°F to -19°F j and 10°F to -13°F on the upper slopes. Conditions such as these were responsible for the d i f fe rent ia l mortality observed in 1949-50 and 1954-55 and possibly also for the mortal i -ty being restr icted to val ley bottom in 1953-54 and 1955-56 (See Table XVI and others). Figure 14. Hygrothermograph tracing showing temperature conditions at valley bottom (4,800') and 5,550' on Mount Eisenhower from January 29 to February 1, 1956, when area was in polar continental air. Figure 15. Hygrothermograph tracing showing temperature conditions in the same locations from February 3 to 6, 1956, when area was in polar maritime air. CONDITIONS UNDER POLAR MARITIME AIR FEB. 3 FEB. 4 FEB. 5 FEB. 6 — mP - * m P 24 12 24 12 24 12 24 12 24 4800' 5550' The second postulate was tested i n 1953-54. Winter sampling was carried out on three of the sample areas, Mount Eisenhower, Cathedral Mountain and Lake Louise (See l i f e tables V and IX). The f i r s t samples were collected December 10 and 16 and examined from the 15 to the 20th.; the second samples were collected February 2 to 5 and examined from the 3rd to the ninth. The usual spring sampling was done i n 1954, the results of which are presented i n the mortality tables above. The summarized results of the three samples are presented i n Table XXV. TABLE XXV PERCENTAGE WINTER MORTALITY - 1953-54 Location Elev. Dec. 1953 Feb. 1954 Spring High Low High Low 1954 Mount 4600 8.6 4.3 95.0 94.6 98.4 Eisenhower 5300 2.6 2.4 19.4 18.1 17.7 5800 13.7 6.2 17.0 15.6 6.3 6300 4.3 - 9.9 9.1 1.5 Cathedral 4500 7.3 17.4 18.4 11.2 Mountain 5000 7.1 - 29.2 31.3 7.0 5500 9.0 3.8 6.8 7.5 3.7 Lake 5050 10.0 5.4 94.1 91.7 99.6 Louise 5550 69.9 43.5 12.5 11.3 15.8 6050 17.0 13.0 39.0 20.6 17.4 6550 19.7 13.9 8.7 6.8 There i s considerable discrepancy between the mortality estimates i n the winter months ( p a r t i c u l a r l y February) and the f i n a l spring estimate. This i s largely due to the fa c t that the larvae brought i n i n February were less e a s i l y roused from dormancy (58) and when they f a i l e d to respond i n any way they were classed as dead. The great differences between December and February at Mount Eisenhower and Lake Louise and the close agreement between the February and spring samples for the low elevation at least leaves l i t t l e doubt that the bulk of the mortality found occurred between December 16 and February 5th. The fact that the high mortality found was restricted to valley bottom indicates a weather phenome-non similar to that described above. Examination of weather records of two stations, Banff and Lake Louise, both on the valley bottom and approximately equal distances in opposite directions from the sampling area should give some inkling of the mortality-producing agent. The only period of prolonged cold between the sampling dates was from January 10 to 29th. The maximum and minimum temperatures for this period are presented in Table XXVI (See appendices 1 , and 2). It is reasonable to assume that temperatures at valley bottom Mount Eisenhower would l i e somewhere between the temperatures given at the two stations. If we assume that the inversions which occurred so commonly in January, 1956, also occurred during this period, then i t is likely that temperatures at the sampling stations above valley bottom were considerably warmer. As a comparable mortality pattern was observed in 1955-56 i t is likely that the causes were the same. The air mass summaries (Appendix 9) show that during the winter of 1953-54-> the area was in polar conti-nental air for 22 days. This is only three days less than in 194.9-50 but the important difference was that in 1950 there were only two polar continental fronts in January with a total number of 60. 3 fronts of a l l types whereas i n 1953-54 there were eight cP fronts with a t o t a l of 20 of a l l types. This means that there was less stagnation of polar continental a i r during the winter of 1953-54 and temperatures throughout the whole v a l l e y had less chance of becoming as extreme as i n 1949-50. TABLE XXVI DAILY MAXIMUM AND MINIMUM TEMPERATURES, JANUARY 10-29. 1954 BANFF AND LAKE LOUISE Temperature of Banff Lake Louise Date Max. Min. Max. Min. Jan. 10 17 - 1 18 - 26 11 21 8 26 - 8 12 17 - 5 15 - 23 13 18 9 16 7 14 5 - 15 - 8 - 16 15 - 22 - 26 - 21 - 35 16 1 - 39 - 3 - 52 17 4 - 13 - 2 - 8 18 6 - 19 8 - 31 -.9 - 9 - 18 - 15 - 32 20 - 11 - 33 - 12 - 47 21 - 18 - 26 - 14 - 23 22 - 16 - 23 - 6 - 25 23 - 15 - 25 - 14 - 22 24 - 12 - 21 - 12 - 19 25 - 1 - 20 - 7 - 20 26 1 - 14 15 - 12 27 17 - 24 14 - 21 28 27 10 19 7 29 18 - 3 17 - 21 Comparison of 1953-54 winter temperatures with those for 1948-49 shows a more severe and prolonged cold period i n 1953-54. In 1953-54 the maximum was below zero eight days and the minimum 18. In 1948-49 the maximum was below zero on only fouri'days'and the minimum 17. Also, i n 1953-54 the maximum was below zero for seven consecutive days, the minimum for 14. consecutive days. In 1948-4.9 the maximum was below zero for only two consecutive days, the minimum for only eight consecutive days. Thus, the comparison of these three winters shows that the winter of 1953-54 was intermediate; conditions were not severe enough to cause the ex-treme mortality at a l l altitudes as in 1949-50, but were severe enough to cause high mortality at valley bottom. The general heavy mortality of 1954-55 was comparable to that of 1949-50 although slightly less severe on the upper slopes. The air-mass summaries (appendix 9) do not show the large number of days In which the area was under polar continental a i r as i n 1949-50 but frontal activity was reduced and stagnation over the 12-day period could have occurred. Calgary weather records show that, generally, 1954-55 was a mild winter (l) so this appears to be a slightly different case. The air-mass summary shows that during March the area was under the influence of cP a i r for 16 days. This i s a high number for March, being exceeded only three times since 1920 and equalled twice. It i s possible, therefore, that March was the 'coldest month' of 1954-55 and winter mortality occurred at that time. I f temperatures in Banff corresponded to those observed at Calgary the extremes of temperature to which the needle miners were exposed could have been the mortality factor involved. Daily maximum and minimum records are not yet available for Banff and Lake Louise for the winter of 1955-56. However, the records of Calgary, Alberta (84 miles East of Banff and out of the Rocky Mountains), for this interval show several periods when this mortality could have occurred ( l ) (see appendix 3). However Calgary records are not always applicable to the outbreak area, <*s the eastern ramparts of the Rooky Mountains protect that portion of the Bow Valley in Banff National Park from many i n -vasions of cold continental arct ic a i r from the east and north-east. As w i l l be seen la ter this keeps the Eastern Slopes of the Rockies colder, on the average, than the outbreak area. We cannot, therefore, with any degree of confidence, pinpoint the period when mortality occurred In 1955-56 u n t i l figures are available for Banff and Lake Louise, During the winter of 1945-46 mortality was apparently high, although the degree of mortality is not certa in . Examination of the dai ly temperatures for this winter at Banff and Lake Louise (Appendices 1 and 2) shows that in general i t was a mild winter but the mild periods were separated by sharp cold periods. Thus a minimum of -35°P at Lake Louise and -26°F at Banff was recorded on November 8, whereas on November 3 the maximum was 51°F and the minimum 31°F. Similar cases, though less extreme, occurred through-out the winter. The air-mass summary (Appendix 9) shows that the area was in cP a i r for 12 days in November, This amount has been exceeded only once (1927-28) and equalled once (1935-36)., The only safe conclusion is that the mortality was perhaps overestimated although the comments made above s t i l l may obtain. As samples made were small and local ized i t would be unwise to generalize. The f ive winters during which heavy mortality occurred have been discussed In de ta i l and when mortality is compared with that of other years i t should be possible to come to some general con-clusions as to what temperature l imits the needle miner can with-63. stand. For this ve use pr incipal ly Table XIV. Thus the years 1943-44, 1946-47, 1947-48, 1948-49, 1950-51, 1951-52 and 1952-53, were years of low mortality ( i . e . highest 31.6) and the years discussed above were years when mortality was extreme in some locations. Air-mass summaries are available for only three of the low-mortality winters 1946-47, 1948-49 and 1952-53. These show no long-term Invasions of cold polar continental a i r except for 1948-49, but the high frontal ac t iv i ty pointed out ear l ie r (43) reduced the opportunity for stagnation of a i r . Table XXVII summarizes the low temperature information for the winter months of s ix of the 'low mortality' winters. Extreme minima alone s igni fy l i t t l e , time and variat ion of temperature may be equally important (See section (1)). Mean maxima and minima and mean monthly temperature give some indication of extent and variat ion of temperature. In par-t icu lar cases however, a careful scrutiny of da i ly temperatures would probably be required. Table XXVIII presents comparable information for the years when mortality was high; 1945-46 (?), 1949-50 and 1953-54. Comparison of these two tables year by year reveals certain differences which could contribute to differences in mortality. We may, therefore, generalize that the lodgepole needle miner popul-ations in the outbreak region in Banff National Park can have a o o high survival i f extreme minima of -30F to -40 F do not persist long enough to depress the mean monthly temperature close to or below the zero mark. This generalization must be considered with respect to the month in which low temperatures occur and the TABLE XXVII SUMMARY OF WINTER TEMPERATURE DATA - YEARS OF LOW MORTALITY BANFF, ALBERTA. Extreme man man "1 Monthly — Year Month Minimum Maximum Minimum Mean' 1943-44 Nov. 4 38*5 19.5 29.0 Deo. - 8 28.0 10.7 19.4 Jan. - 8 29.1 11.0 20.0 Feb. - 13 28.8 5.9 17.4 Mar. - 15 35.8 10.4 23.1 1946-47 Nov. - 27 27.2 9.0 18.1 Dec. - 31 25.6 7.2 16.4 Jan. - A l 24.0 5.8 14.9 Feb. - 34 30.3 5.2 17.8 Mar. - 31 39.8 8.5 21.6 1947-48 Nov. - 9 31.5 14.3 22.9 Dec . * - 15 26.4 5.0 15.7 Jan. - 21 30.8 11.1 21.0 Feb . * - 29 23.8 - 6.0 8.9 Mar. - 23 33.6 7.5 20.6 1948-49 Nov. 0 33.9 18.6 26.2 Dec. - 28 16.9 - 1.8 7.6 Jan. - 44 15.3 - 7.4 4.0 Feb. - 39 21.7 - 2.9 9.4 Mar. - 19 38.8 15.5 27.2 1950-51 Nov. - 28 28.4 11.0 19.7 Dec. - 20 28.5 14.3 21.4 Jan. - 41 16.2 - 4.2 6.0 Feb. - 23 26.7 1.3 14.0 Mar. - u 28.1 5.4 16.8 1951-52 Nov. - 13 33.3 17.5 25.4 Dec. - 39 14.5 - 1.6 6.4 Jan. - 34 17.8 0.1 9.0 Feb. - 15 29.3 9.5 19.4 Mar. - 13 35.0 10.3 22.6 6 Lake Louise - Banff not complete. 65. TABLE XXVIII SUMMARY OF WINTER TEMPERATURE DATA - YEARS OF HIGH MORTALITY BANFF, ALBERTA. Extreme Mean Mean Montnoy Year Month Minimum Maximum Minimum Mean 1945-46 Nov. - 26 27.4 8.8 18.1 Dec. - 15 22.8 7.6 15.2 Jan. - 13 28.7 12.7 20.7 Feb. - 15 31.8 11.2 21.5 Mar. 3 40.3 20.6 30.4 1949-50 Nov. 11 45.1 28.6 36.8 Dec. - 18 17.0 0.9 9.0 Jan. - 51 - 5.2 - 26.7 - 16.0 Feb. - 18 31.9 12.1 22.0 Mar. - 29 31.2 11.9 21.6 1953-54 Nov. 6 38.4 24.9 31.6 Dec. 0 29.6 17.2 23.4 Jan. - 39 11.1 - 4.1 3.5 Feb. - 6 35.9 21.7 28.8 Mar. - 22 33.1 9.1 21.1 duration of cold cP or cA a i r in the region. Thus, in November or March a higher minimum may be l e t h a l , part icular ly i f alternated with warm periods. It is unlikely that extremes of temperature of short duration are a major cause of winter mortal ity. It would be unwise to generalize on the mortality in populations in a region such as was encompassed by the past out-break from a few samples at one alt i tude or from the weather records of a 8ingle station such as Banff, or even from the reasonably comprehensive sample coverage given to populations in Banff National Park. The comments made above do not obtain to mortality conditions found in Yoho or Kootenay National Parks f at least not to the degree found in Banff Park. 6 6 . The lack of temperature data from Yoho and Kootenay Parks and, as yet , a lack of understanding of the complex relationships of a i r masses in the trans-Divide area do not permit more than speculation. That the populations in these areas have subsided at an equivalent rate to those in Banff Park is certain but the causes for this are less cer ta in . That i t is attributable to weather factors we are reasonably cer ta in , as neither parasites nor disease have ever figured prominently in the control complex. In the Yoho area climatic conditions are much more var iable , i f less extreme, than in the Banff area and while these conditions have not resulted in excessive mortality except in later years (Table XVIII) i t is l i ke ly that they have affected the population in other ways, through development, f e r t i l i t y and fecundity, a l -though there is no material evidence for this hypothesis. In the Kootenay area, mortality differences are not as great (Table XXII) and the remarks pertaining to Banff Park probably are pertinent but to a lesser extent. Alt itude and orographic differences probably are in f luent ia l in causing the variations noted. With increased use and refinement of air-mass and frontal analyt ica l techniques and consideration of loca l fluctuations in olimate in the Bow Valley such as discussed above i t is now possible to develop a predictive system for use in mortality and population studies. Prom the above studies we now know that the heaviest persistent infestations are l i k e l y to be centered on the mid-slopes in the Bow Valley of Banff National Park because although popul-ations may increase above and below this favorable zone during a few favorable years they w i l l eventually suffer 'catastrophic' mortality 67. during the types of winter weather noted above (152). 3.1.2. Spring mortality. A small percentage of larvae k i l l e d after feeding in the spring, has been found since detailed sampling for l i f e tables studies began in 1954- (see l i f e tables) . This mortality never exceeded six per cent of the la rva l population entering hibernation. That spring mortality can be an important factor has long been recognized, part icular ly in open-feeding insects (1A4-). The tent caterp i l la r has suffered 'catastrophic' mortality from spring temperature extremes in Ontario (8) and spring frosts shortly after hatching were credited with a high degree of control in one out-break in Minnesota (UU). Early studies on the needle miner indicate that commencement of feeding in the spring i s largely dependent on spring temper-atures. Generally, i t appears that feeding ceases in the f a l l when maximum temperatures f a l l below A5°F and the minimum temperature i s commonly below freezing. These conditions in reverse are associated with commencement of feeding in the spring (110,128). Spring f ros ts , common during this period, may be severe enough to cause the mortality observed. Lack of knowledge of the precise time of occurrence and the low incidence of the mortality precludes intensive discussion of the temperature fluctuations for this period. 3.1.3. Other climatic factors possibly involved in the reduction of populations. ( l ) . Effects of weather on egg and f i r s t instar larvae. 68. No mortality of eggs in the f i e l d has been observed. They are apparently able to endure any f i e l d conditions which occurred in the two years that egg sampling was carried out. However, that large reductions in populations between oviposition and larva l establishment may occur per iodical ly i s evidenced by 1954 sampling when a count of 4700 eggs per 100 t ips was made and larva l establishment was only 1,114 lsvrvae per 100 t i p s . No such loss was observed in 1956. That this loss i s not due to a factor acting d i rect ly on the eggs i s reasonably cer ta in . A l l eggs kept for rear-ing and experimentation (without extensive handling) hatched and no dead eggs have ever been found in the f i e l d . In 1954 l imited tests were conducted on the effects of humidity and temperature on egg development. While these exper-iments were not precise enough nor extensive enough to report in deta i l they did indicate that eggs of the needle miner are capable of withstanding extremes of humidity and temperature not found in the f i e l d for any length of time. However, development may be delayed by adverse conditions which may have an effect on successful establishment of the larvae (109). Two sources of loss have been observed in the f i e l d but techniques have not been designed by which they can be evaluated except by subtraction from present sampling stages. One cause of loss i s the drop of mined needles containing eggs. The other cause i s the prevention of la rva l establishment by adverse weather factors . That drop of mined needles containing eggs does occur has been substantiated by examination of the needle l i t t e r at the base of t rees. No estimate of loss has yet been made but i t i s reasonably 69. certain that such eggs would be l o s t to the population. Morgan (77) estimated t h i s loss to be 18 per cent i n the C a l i f o r n i a needle miner. As loss of eggs by needle casting did not occur i n the Bow Valley i n 1956. i t i s probably only an occasional phenomenon, perhaps r e s u l t i n g from strong gusts of wind during the time of egg development. No measurement has been made of loss of hatching larvae but i t i s recognized that i t could be an important factor In popul-ation reduction. Experiments by Shepherd (103) indicated that the threshold of a c t i v i t y of newly emerged larvae i s rather high (59 - 65°F) and the l i m i t e d experiments on eggs noted above indicated that high humidities (usually accompanied by cool weather) r e s t r i c t f i r s t - i n s t a r l a r v a l a c t i v i t y . Examination of hygrothermograph records maintained on Mount Eisenhower i n 1954 Indicates a c o l d , wet period from August 18 to August 24th which corresponds c l o s e l y to the beginning of the hatching period. Temperature and humidity conditions f o r t h i s period are shown i n Figure 16. Again, no measurement of loss i s possible except by interpolation from the l i f e table but i t i s reasonably certain that the combination of low temperature and high humidity i s responsible for the loss noted i n 1954 between ovipo-s i t i o n and l a r v a l establishment. Humidities were above 90 per cent f o r most of t h i s period, except for a short time on the 20th, 21st, and 22nd. The temperature, except for the mid-day period on the 20th (6l°F) went to 55°F only once and was about 50°F or less f o r the remainder of the period. As t h i s loss between oviposition and l a r v a l establishment did not occur i n 1956, comparison of conditions f o r the same period should be markedly d i f f e r e n t i f the above suppositions are Figure 16. Hygrothermograph tracing shoving humidity (upper) and temperature (lower) conditions on Mount Eisenhower, 5,550' from August 18 to 25, 1954- and 1956. Sol id l ine - 1954 Broken l i n e - 1956 HYGROTHERMOGRAPH TRACING - 2400 AUGUST 18 - 1200 AUGUST 25 (MST). MOUNT EISENHOWER - 5,550' AUG.18AUG.19 AUG.20 AUG.21 AUG.22 AUG.23 AUG.24 AUG.25 24 12 24 12 24 24 24 12 24 12 24 . 12 70. correct . The dotted l ine in Figure 1 6 shows the hygrothermograph tracing for the same period but from a thermograph maintained at Elsenhower F ie ld Station a few miles away. Conditions in 1 9 5 6 were more favorable for la rva l establishment than in 1 9 5 4 . . The lower night temperatures in 1 9 5 6 were not severe enough to cause mortal i ty. ( 2 ) Effects of climatic factors on larvae during the summer. Morgan (77) has suggested that larvae of R. m i l l e r l Busck may be forced from their mines by excessive heat and that this may be an important factor in contro l . I t has been established that temper-atures of mines may be increased by as much as 1 1 ° F under optimum radiation conditions (42) but the behaviour suggested by Morgan has never been observed in the Canadian needle miner. Although actual mortality does not occur during summer when only larvae are present, adverse cl imatic factors during this period may have a long-term effect on the population by affect ing la rva l development. The only occasion observed when this may have happened to a marked degree was in Yoho National Park in the summer of 1 9 5 1 . This summer was cool and wet and the la rva l transfer from f i r s t to second needle lacked any semblance of order. Transferring began about June 20th and on August 23rd some larvae were s t i l l trans-fe r r ing . Approximately 90 per cent had completed transfer by the 30th of July but this process i s usually completed in two to three weeks in July (123). This could be a contributory factor to reductions in populations by long-term effects on winter survival and possibly on fecundity. As this phenomenon was observed only in the Cathedral Mountain sampling area i t i s also possible that t h i s , rather than excessive winter mortality was one of the factors which caused 71. reduction of Yoho populations. (3) Effects on punaa. As shown in the l i f e tables, in a l l areas except Mount Cathedral in Yoho Park mortality of pupae was estimated at from 20 to 25 per cent in 1956. No other factor i s evident for this mortality except cl imate. As there was no apparent mortality of pupae in 1954 comparison of weather records during pupation should show a d i f f e r -ence. Examination of hygrothermograph charts from Eisenhower F ie ld Station for June in both years gives l i t t l e basis for speculation. The mean maximum temperatures differed by less than 1°F, the mean minima were almost iden t ica l . The considerably higher re la t ive humidity in 1956 than in 1954 eliminates the poss ib i l i t y of desic-cation as an explanation. There occurred in June, 1956, a four and a s ix day rainy period, separated by only one day, which were charac-terized by temperatures averaging 42°F. On f ive of these ten days the minimum was below the freezing point. There was an equivalent number of days in June, 1954» with minima below the freezing point but in 1954 these were not usually associated with low maximums. Thus on f ive consecutive days in June, 1954, when the minima f e l l below freezing, the maxima were a l l 60°F or above, with an average dai ly temperature of 45*4°^ for the f ive day period. It i s unl ikely that high temperatures are involved although i t has been demonstrated by the work of Henson and Shepherd (42) that needle mine temperatures may be increased by as much as 11°F in the most intense radiant conditions (see also 150). If this were the factor involved we should have found equal or greater mortality in 1954 as the month of June, 1954, was generally warmer and sunnier than 72. June, 1956. As pupal development was retarded and this pupal mortality did occur, i t is therefore postulated that in 1956, developmental con-ditions were sub-optimal and the period described above (June 5-8; 11 - 16) may have been a contributory factor to the mortality. U) E f f e c t s of various factors pn moth b e h a v i o u r In respect to n,eed,le miner abundance. AU moth activities relating to insect abundance, flight, copulation, and oviposition are dealt with together, as well as fecundity and fe r t i l i t y , for the same factors have some effect on them a l l . Detailed information on this extremely important stage is scarce and we have data from only two generations when the population from 'potential' to actual has been assessed, viz; 1954 and 1956. The estimate of potential population is extremely crude and only indicative. Dissections by several workers have shown that egg potential in the needle miner approaches 100 eggs per female (35,125, unpublished data). The number of apparently mature eggs in captive females was found to be 16 to 30. Whether this number is ever exceeded in nature is not known. Field samples in 1954 and 1956 indicated a reduction in egg potential or in ability to lay eggs . The average number of eggs laid per female (from l i f e tables) was nine and eight respectively. It is therefore possible that we have been overestimating the potential egg capacity of the needle miner and part of the egg losses discussed above does not actually occur. The apparently low fecundity of the females in 1954 and 1956 may be the effect of adverse climatic factors acting on developmental 73. stages (see Section 3 .1 .1 . ) . Conditions during the 30-day pupal I! period could have a drastic influence on oogenesis of females. The low temperatures during pupal development may have interrupted development enough to be detrimental to eventual egg development (102,144). Pradhan (97) found that development of Earias fabia S t a l l was quicker under variable than constant low temperatures but slower under variable than under constant high temperatures. I f t h i s were true of the needle miner, a month characterized by high maxima (70°F) and low minima (about freezing), or short periods of cold weather followed by warm to hot weather could have a delaying e f f e c t on pupal development and a resultant adverse ef f e c t on moth fecundity. l e t another factor which may ef f e c t fecundity i s the presence of non-fatal disease i n the moth population (86). There are many factors which may a f f e c t successful o v i p o s i t i o n . Possibly greater i n h i b i t i o n of oviposition i n 1956 than i n 1954 was indicated by egg samples. Fewer large egg masses were found i n 1956 than i n 1954 and the number of single eggs found i n proportion to the t o t a l number was f a r greater. Greatest f l i g h t a c t i v i t y occurs at sunset. This i s believed to be due to diminishing l i g h t . Diminution of l i g h t (or radiation) such as occurs p r i o r to heavy clouds passing over, caused increased a c t i v i t y during the day. At sunset, i f there i s no wind or r a i n , a c t i v i t y i s at a peak, the moths f l y i n g upward to the tops of the tree crowns, which gives r i s e to the d i s t r i b u t i o n of eggs and larvae described e a r l i e r . I f the temperature i s not too low t h i s f l i g h t occurs even when the sky i s overcast but Is less than when the sky i s c l e a r . The moths are quiescent at winds above 5 m.p.h. and during r a i n storms. On calm 74. days there i s general exc i tab i l i ty at high humidities but as these were usually associated with cloudiness the precise factor involved i s not certain (35). Fluctuations in barometric pressure cause increased f l i g h t act iv i ty in some insects (131,148). However, as great f l i gh t a c t i v i -ty of the needle miner occurs at sunset on calm, reasonably warm days, changes in pressure cannot be part icular ly s ign i f icant . Pressure could be a minor factor during daylight hours when associated with changing weather conditions ( l6) . On hot, sunny days, when the humidity i s low, needle miner moths seek shaded locations and display a minimum of ac t iv i ty . On cloudy days act iv i ty occurs in spurts, associated with passing clouds. Cold, windy, or rainy weather during the moth f l igh t could have a profound effect on the success of oviposition and resul t in a wide discrepancy between the potential number of eggs and the actual number l a i d . 3.2. ParasiUaro Parasites have long been upheld as the major influence in so-cal led "biotic control" and many instances have been ci ted where economic control of insect pests has been ef fect ive , largely through the introduction of new parasites. Sweetman (132) subjected many claimed examples to intense scrutiny and came to the conclusion that many of the 'controls ' claimed to be due to b io logical factors , parasites and predators, were in fact , due to other factors.. How-ever, he concluded that parasites can be a control l ing influence on insect populations. Andrewartha and Birch (3) quote Elton who states "It i s 75. becoming increasingly understood by population ecologists that the control of populations i . e . the ultimate upper and lover l i m i t set to increase, i s brought about by density-dependent factors , either within the species or between species. The chief density-dependent factors are in t ra -speci f ic for resources, space or prest ige, and in ter -speci f ic competition, predators or parasi tes." They, however, take strong exception to this statement and claim there i s l i t t l e evidence to support i t . Indeed, they go so far as to c a l l i t downa rather than a conclusion (p.19). Milne (75) claims that there has been no proven case of control of an insect population by i t ' s natural enemies but abundant evidence exists that they are unable to do so. However, i t i s not the intention here to enter into this philosophical argument but merely to point out that one does not need to apologize for the fa i lure of a parasite complex to control the host; there are as many people decrying the inef f ic iency of b io logical control as support i t . However, the tendency towards control by parasites i s recognized and for the proper understanding of the dyna-mics of an insect population one must analyze and explain the reasons for fa i lure of b io logical contro l . As has been par t ia l l y shown in the l i f e tables presented above and w i l l be shown in greater deta i l below, the parasite complex of the lodgepole needle miner, Recurvaria starki played no major ro le in the str ik ing declines of population already described. 3.2.1. Description of the parasite complex of R. a tark i . Table XXIX Is a l i s t of the known parasites of the lodgepole needle miner. The l i s t i s compiled from the Annual Reports of the 76. author, the Calgary laboratory of Forest Biology, the Biological Control Laboratory, B e l l e v i l l e , Ontario, and numerous short pub l i -cations (48,57,72,114,118,122). The determinations were made by the Systematics Unit , Division of Entomology, Ottawa. I t i s im-possible to indicate some of the parasites as larva l or pupal as they were obtained from mass rearing and are extremely rare. The order of l i s t i n g i s taken from Miesebeck e t . a l . , ( 8 5 ) . Three of the species l i s t e d comprise 90 per cent or more of the total number of parasites present in f i e l d populations. These are Cqpidosoma n.sp. and the two Apanteles species. Most of the parasites are rare and l i t t l e information i s available on these but i t may be of value in discussing their abundance and dynamics to summarize what i s known about them. These w i l l be dealt with in the order given in Table XXIX and not in the order of their importance. 1.2. Ananteles cal i fornicus Hies, and Ananteles sp. As there i s no apparent difference in the behaviour or l i f e cycle of these parasites they are discussed together. Muesebeck states that a l l species of this subfamily seem to be internal parasites of lepidopterous larvae ( 8 5 ) . A. cal i fornicus i s described from Oregon and Cal i forn ia and Recurvaria w ^ l p - H Bsk. i s given as i t ' s host. The l i f e cycles of the parasites follow that of the host a l -though there i s a greater v a r i a b i l i t y in emergence dates. Emergence records for 1954. show that the parasites emerged about the same time as the needle miner, in early July . In 1956, parasite emergence was markedly ear l ier and there was a tendency towards two emergence peaks, 1 the 23rd and 28th of June. This may be due to a species difference but i t i s thought in this instance to be due to climatic inf luences. 77. After the f i r s t peak of emergence,cold, rainy weather set in and did not l e t up u n t i l June 27th, the day before the second emergence peak. Emergence was high up to the end of the f i r s t week In Ju ly . MtsLeod (72) records emergence of Anantelea in late July and August but th is was not the case in 1954 or 1956. It i s not known for certain which stage of the needle miner the Apanteles species attack but from the emergence dates and the length of l i f e of the parasites (approximately three weeks) i t i s presumed to be the egg stage. Morgan (77) states that this i s the case with Apanteles in Ca l i forn ia whose l i f e span i s f i f teen days, but h is be l ie f i s also based on circumstantial evidence. The parasite i s internal and does not affect feeding behaviour u n t i l the fourth or f i f t h ins tar . Upon the death of the needle miner larva the parasite emerges from the skin of i t s host and spins a white, opaque, si lken cocoon within and near the base of the mined needle. Emergence i s effected by cutting of f a c i rcu lar cap from the cocoon and crawling out of the exit hole prepared by the needle miner la rva . The parasites are not v i s i b l e u n t i l they leave their hosts, in mid-May (See Figure 17). At least two species are hyperparasitic on Apantelaa one of which i s l i k e l y Alegfaifl. p in l fo l l aa (C U sh . ) , the other i s not known. The hyperparasites are either polyembryonic or multiple parasites since Apanteles cocoons have been found with up to four emergence holes. The effect of the hyperparasites on the numbers of Apantales has been s l ight to date. 3. Eubadizon f r a o l l a (Prov.) . Very l i t t l e i s known of the subfamily to which this parasite belongs. E. graci le i s l i s t e d from Canada TABLE XXLX PARASITE COMPLEX OF RECURVARIA STARKI FREE 78. Superfamily Family ICHNEUMDNOIDEA Braconidae Ichneumonidae CHALCIDOIDEA Eulophidae Encyntidae Pteromalidae Chalcididae Species 1. Apan.te3.es c a l i f ernjcus, Maes. 2. Apanteles sp. 3. Eubq,dJ.zon PT&clle Prov. 4.. Meteorus n.sp. 5. Alesdna, p i n i f o l l n a (Cush.) 6. Gelis tenellus (Say). 7. ItoplQQti^ Qfregua Cush. 8. PhaedrQctonus sp. near epinotiae Cush. 9. Phaepgmgs sp. near flpiflotjae, Cush. 10. Diclad,pcqrus sp. 11. Derostenus sp. (?) 12. Etiderus sp. 13. Neo4erQ8lfflffla n.sp. H- Sympj-gs^s sp. 15. Terras Uchus, sp. 16. ZagraropjasQma amerftcana G i r . 17. One unknown. 18. Copido9oma n.sp. 19. Amblvmerus sp. 20. Habrocvtus sp. 21. Pachvneuron sp. 22. S p i l o c h a l c i s sp. prob. albjfrons (Walsh). 79. (no speci f ic loca l i t y ) and Maine. The only host given in Muesebeck i s Recurvaria piceael la Kearf (85). Prebble (pers. comm.) found i t on the black-headed budworm in Br i t ish Columbia. L i t t l e i s known of i t s behaviour. The parasite i s internal and leaves the host carcass in late May and spins a tough, brown, translucent cocoon midway between the base of the mine and the exit hole within the needle. The needle miner i s unaffected u n t i l early May. Emergence has occurred as early as June 18th and as late as July 10th. Specimens were too few to postulate a peak emergence period. Specimens in capt iv i ty l i ved only a few weeks. A Epbadizon sp. i s l i s t e d from R a m i r v « - H « n r f n ^ B a V . (77). (See Figure 17). 4. Meteorug sp. Nothing i s known of i t s l i f e cycle or behaviour as i t was obtained from mass rearing and i s rare . A l l species are described as internal parasites (85). 5. Aleoinw n ln i fo l iae (Cush.). This and the next species Gelia  tenellus (Say) are of the same subfamily. The only other host r e -corded for A. p i n i f o l i a e i s an eastern needle miner Eyoteleia p i n i f o l i e l l f l (Chamb.). This i s one of the suspected hyperparasites of Apanteles species. Members of the same tr ibe are described as occasional or habitual secondary parasites. Many hosts are recorded for G. tenellus but of part icular interest are three species of Apantales and four of Meteorus (85). Neither species i s recorded from Cal i forn ia (77). 7. I toclect is obesua Cush. Members of the tr ibe to which this belongs are parasites of a great variety of pupae.and prepupae of Lepidoptera. 60. There i s l i t t l e host spec i f i c i t y and adults have a corresponding v a r i a b i l i t y In size. The species of Itonlectis may also be secondary parasites (85). It i s not recorded from California (77). 8. Phaedroctonus sp. The tribe includes many important parasites of economic pests. Most species attaok lepidopterous larvae (85). A Phaedroctonus sp. occurs on £. m i l l e r l Bsk. (77). 9. Phaftpgenes sp. The members of the subfamily to which PhBeogenea belongs are described as internal parasites of Lepidoptera. They oviposit into either the host larva or pupa, but always emerge from the pupa (85). The only note made on this relatively rare species i s that I t was observed emerging from the needle miner pupa on August Ath and was extremely active. Depending on the length of adult l i f e i t could either parasitize needle miner eggs or f i r s t Instar larvae. There i s some evidence to suggest that i t may have a one-year cycle. 10. Dlcladocerus sp. This internal larval parasite was not found i n 1951 by McLeod and was one of a number of species of parasites l i b e r -ated in the Banff area. The presence of i t in later rearings may i n d i -cate establishment of this species. It i s very rare, however. Morgan l i s t s i t along with three other Eulophids, SympAesjLa sp., Derostttma sp. and Tetrastichus sp. as occurring in third-and fourth-instar larvae of R. miller! i n the lar v a l year (77). Such a phenomenon has never been recorded i n the Canadian outbreak. 11. Deroatenua sp. Other members of the same subfamily are Euderaa sp., and Neoderoatonus n.sp. Nothing i s known of the f i r s t two except that they are very rare. Neoderostenus n.sp. was recorded as 81 an important parasite in Alberta by McLeod in 1951 (72) but sampling since this time indicates that It i s re la t ive ly rare . I t oompletes i t s development in the host pupa and may parasit ize the larva in the same year carrying over on alternate hosts. I t forms the naked black pupa typica l of Eulophids. There are usually four to s ix parasites per pupa but whether this i s due to polyembryony or multiple parasit izat ion i s not cer ta in . It i s also recorded as a secondary parasite of Copidosoma sp. (72). H • Svnmlesls sp . This i s believed to be an external la rva l parasite and was re la t ive ly abundant in Alberta (72). This i s not the situation now. Parasites believed to be Svmpiesis sp. have been recovered from needle miner larvae and pupae and also from the important parasi te . Copidosoma n.sp. Morgan records i t as an external parasite on la te instar larvae of R. m i l l e r i (77). 16. Zagrammasoma amerlcana G i r . This was one of the ear l iest recorded parasites from Alberta. It was found emerging from host larvae and also from cocoons of Cfipldnanttiw, n.sp. Since that time i t has regu-l a r l y appeared in mass rearings but in small numbers. It also Is l i s t e d as a parasite of R. m i l l a r i ( 77 ) . 18. Copidosoma n.sp. This Chaleid parasite was f i r s t thought to be taxonomically near C. nanellae S i l v . However, in 1954* Mi l ler (pers. comm.) stated: "A further study of the male geni ta l ia of specimens from (Alberta, Br i t ish Columbia, Idaho and Cal i fornia) has made i t possible for me to state that there i s but one species of Copidosoma attacking lodgepole needle miner in Western North America." This includes Recurvaria m i l l e r i , R. starki and possibly several other 8 2 . R e c u r v a r i a , species. It i s a polyembryonic species of which as many as 14 adults emerge from a single host la rva . The average in R. starki i s about seven per host. Emergence occurs over a re la t ive ly short per iod. In 1956 this period extended from July 11 to July 30 with the peak occurring about July 19. This was s l igh t ly ear l ier than in 1954 when emergence began about July 15 and extended into the f i r s t week in August. This corresponds with the oviposition period of the needle miner adults. Recently emerged adults were placed in a v i a l containing a mined needle containing needle miner eggs. The parasites seemed capable of determining where the eggs were in the needle by tapping the needle with their antennae. Once found, the adult investigated the egg cluster repeatedly by insert ing her head into the exit hole . If the egg cluster was too far from the hole for the parasite to reach with her antennae or ovipositor the female would either leave the needle or investigate the surface of the needle around the egg c lus ter . As they were found oviposit ing through cracks in the dried needles i t i s possible that this condition i s what they were searching f o r . Many unsuccessful attempts at oviposition were observed, the ovipositor repeatedly sl ipping off the egg surface. When several adults were placed in the same v i a l with only one egg d u s t e r , they were seen to parasit ize the same eggs. Individual females were seen oviposit ing into the same egg several times. Larval sectioning has established that th is parasite has a l i f e cycle corresponding in length to that of Recurvaria starki Free. (See Figure 18). Figure 17. Parasites of R. atarki Free. (a) Apanteles spp. cocoon. (b) Cocoon of Eubadizon graci le Prov. (c) Unidentif ied pupal parasi te . (d) Spi lochalcia sp . poss. alMfrQn,s (Walsh). Figure 18. Parasites of R. s tark i Free. ( l ) Copidosoma n.sp. oviposit ing in needle miner egg c lus ter . (?) Multiple parasitism by Copidosoma n .sp . (3) Needle miner larvae parasit ized by Crmidosoma n.sp. Parasites almost mature. 83. 3.2.2. Population dynamics of needle miner parasites. One of the advantages of the l i f e table method i s that i t i s possible to isolate any period or stage in the l i f e cycle covered by the l i f e table i t s e l f and examine i t in greater deta i l (73). Inclusion of such deta i l in the or ig inal l i f e table would make i t unwieldy. The spring of the moth f l i gh t year when parasitism becomes evident, Is of part icular interest . Information gained from a more detailed examination w i l l be of greater value i f we can reconstruct what has happened to the numbers of the parasite complex since the beginning of the investigative period. The f i r s t estimate of parasitism, based on a l imited number of samples, was in 1944- (4-0). Scanty estimates are also available for 194-6 (5 l ) , and reasonably precise estimates were made in 194-8, 1950 and 1952 (112,113,122). More precise estimates with better under-standing of the species complex were made in 1954- and 1956. The proportionate abundance of parasitized hosts i s presented in Table XXX for a l l areas. The estimates in these tables are based upon the populations present Just pr ior to the winter before the moth f l i g h t year. The percentage parasitism i s thus only the percentage of parasites which reached maturity. Except for the small fraction which die after they become evident there i s no way of determining the mortality of parasites except by sectioning of dead needle miner larvae. Praotical considerations do not permit such sampling at present. Two values are shown for the spring of 1950. Twelve per cent Is the degree of parasitism evident in the spring, two per cent i s the actual per cent parasitism which survived the winter of 1949-50. 84 TABLE XXX PERCENTAGE PARASITISM IN MOTH FLIGHT YEARS - ALL AREAS -Year Parasitism per cent Most important parasite species lin numbers; 1944 11 P.yttMosom n.sp. 1946 10 1948 10 1950 12 to 1.7 cOPi?loaoma n.sp. 1952 10 fa&id.Qaoma n.sp. 1954 18 P.OTidQSCm n.sp. Ananteles californicus Muea. APftntales sp. 1956 30 Ayasteles ColifomiflUS Hies. Apantel^s sp. PvPlflosoma n.sp. 85. P a r t i a l records are available f o r several areas which demonstrate the v a r i a b i l i t y of parasitism within the general out-break area. These are summarized i n Table XXXI for areas selected to conform with the four areas now being continuously studied. Two estimates are given fo r the 1950 samples i n t h i s table a l s o . TABLE XXXI PERCENTAGE PARASITISM IN MOTH FLIGHT YEARS FOR FOUR AREAS NOW UNDER CONSTANT INVESTIGATION Area 1950 1952 1954- 1956 Mount Elsenhower 19 - 3.7 4 + 20 39 Bankhead 11 - 1.9 17 16 Girouard 32 Massive 1 3 - 0 - 10 26 Cathedral 0 22 14 12 86. The population data for a l l areas other than Mount Eisenhower were not complete enough for the formulation of l i f e tables. , To make the data comparable therefore, supplementary tables were not prepared for any other than the 1954-56 generations. However a comparison of the species composition of parasites in the spring of 1954 i s of interest (Table XXXII). These data c lear ly show the importance of three of the parasite species. Although i t i s not wholly consistent throughout the whole area of the outbreak Apanteles often appear to be more numerous In the val ley bottoms than on the slopes. This may suggest that they are more cold-hardy than Copidosoma n.sp. or that there i s a fundamental difference in conditions required for oviposit ion. Supplementary tables for the c r i t i c a l spring period of the moth f l i gh t year for the four areas on a continuous study basis are presented in Tables XXXIII to XXXV which relate to Tables X , XII , XIII and XIV. It i s clear from the l i f e tables presented ear l ie r and their supplements above that parasites have not played the major role in the population reductions observed since 1948. I t i s generally accepted that population growth follows a l o g i s t i c curve where density increases u n t i l a point is reached at which the trend w i l l be reversed and the rate of increase w i l l begin to decl ine. F ina l ly a more or less constant density i s reached. It is also generally assumed that this phenomenon i s brought about in two ways: through new and more severely unfavourable processes coming into play at successively higher levels of density and by an increase in the intensity of action of various individual TABLE XXXII 87. SPECIES COMPOSITION AND PERCENTAGE PARASITISM IN ALL AREAS SAMPLED IN 1954 Per cent By species-per cent of total Location Elev. Parasit-ism parasitism Copidosoma Apantel-es spp. Others Mount Eisenhower 4800 - -5300 11.0 39.0 59.0 2.0 5800 18.7 86.0 13.5 0.5 6300 20.3 83.0 16.5 0.5 Massive 4800 1.5 - 100.0 5300 7.8 47.4 52.6 -5800 12.2 48.4 51.6 6300 9.4 21.0 79.0 _ Cathedral 4500 13.6 6.2 92.2 1.6 5000 10.7 53.9 45.1 1.0 5500 28.5 40.4 58.6 1.0 Lake Louise 5000 - -5500 8.6 81.0 18.5 0.5 6000 12.9 95.3 4.2 0.5 6500 6.4 92.6 3.6 3.8 Brewster Creek 5200 69.5 37.9 61.9 0.2 Baker Creek 6000 61.2 30.8 69.2 0 Cascade Valley 5500 16.2 60.0 39.7 0.3 Saskatchewan Crossing 4700 1.6 11.3 88.7 -Hawk Creek 4000 14.9 83.4 16.6 • 88. TABLE XXXIII SUPPLEMENT TO TABLE X FOR MOUNT EISENHOWER SHOWING MORTALITY OF LATE INSTAR LARVAE AND PUPAE lx dxF dx lOOqx Instars HI - IV (1955 - 1956) Instars IV - V May, 1956 Pupae 416 Climate-winter mortality 143 273 105 Parasitism, Copidosoma n.sp. 86 (Apanteles cal i fornious Mues. (Ap. anteles sp. Eubadizon gracile (Prov.) Undetermined species Unknown Unknown-poss. climate Parasitism Phfleoeranas sp. and others less than 68 5 160 168 26 34.37 31.50 24.90 1.83 0.38 58.61 3-9? 61.54 24.76 On45 26 + 25.21 89. TABLE XXXIV SUPPLEMENT TO TABLE XII FOR MASSIVE MOUNTAIN SHOWING MORTALITY OF LATE INSTAR LARVAE AND PUPAE X lx dxF dx lOOqx I I I - IV instars (1955 - 1956) 465 Climate - winter mortality spring mortality 94 ~%~ 20.22 O.A3 20.65 IV - V instars May, June, 1956 369 Predation by birds Parasitism 124 33.60 (Apanteles c a l i f o r n i c u s Maes. ( 66 (Apanteles sp. 17.89 Copidosoma n.sp. 52 14.09 EubadlSpn, ffractte (Prov.) 3 0.81 Undetermined species 123 247 66.94 Pupae June, 1956 122 Climate - desiccation? 30 24.84 Emerged 92 90. TABLE XXXV SUPPLEMENT TO TABLE XIII FOR MOUNT GIROUARD SHOWING MORTALITY OF LATE INSTAR LARVAE AND PUPAE X l x dxF dx lOOqx III - IV instara 896 Climate - winter (1955 - 1 9 5 6 ) mortality 213 2 3 . 7 7 spring mortality 15 1.67 . 228 25.44 IV - V instars 668 Predation by birds 76 11 .38 May - June ,1956 Parasittem, Upantelss cal^ fornAeu .s Mues. ( 228 3 4 . 1 3 Upanteles sp. Copidosom n.sp. 50 7 . 4 8 Undetermined snecieslQ 1*50 288 43 .11 Unknown Z l „ „ 6 J A 4 0 5 6 0 . 6 3 Pupae June, 1 9 5 6 263 Climate - desiccation? 56 21 .29 Parasitism - un-ident i f ied species 2 7t60 58 28.89 Emerged 205 91. TABLE XXXVI SUPPLEMENT TO TABLE XTV FOR CATHEDRAL MOUNTAIN SHOWING MORTALITY OF LATE INSTAR LARVAE AND PUPAE X l x dxF dx lOOqx I I I - IV instars 181 Climate - winter 1955 - 1956 mortality 111 61.33 spring mortality 6 117 64.64 IV - V in s t a r 64 Parasitism May - June, 1956 (Ananteles c a l i f o r n i c u s Maes. 17 25.56 (Ananteles sp. Copidosoma n.sp. L 6.25 Undetermined species 1 1.56 22 24.37 Pupae June, 1956 42 0 Emerged 42 92. density-dependent factors or processes (106). Klomp (61) in reviewing the major theories of host-parasite interaction is of the opinion that the work of Tinbergen supports the theories of Nicholson and the l imit ing or 'damping' of population osci l la t ions is brought about in three ways. The f i r s t is based on Nicholson's theoretical model where the density of the other than the regulated host is independent of the ac t iv i ty of a given parasite and remains constant. This is not realized in nature but i t must be considered that the fluctuations of the numbers are not caused by the parasite, the influence of the parasite being compensated by the regulating mechanism of the host. However at high densities of the parasite due to large numbers of the regulated host i t is unlikely that the regulating mechanism of the alternate host is able to compensate for the high mortality. This does not appear to obtain in populations of the needle miner. F i r s t , there is known no alternate host of suf f ic ient numbers to create the problem, and second the numbers of the parasite have never reached "high densit ies" in spite of the large numbers of the host, or in other words the host has never been regulated in the true sense of the word. The second way in which population osci l la t ions are l imited is by a density-dependent reproduction of the host. Examples are given by Klomp where fecundity of the host is reduced at high levels of abundance which permitted the parasite population to overtake the host population in numbers. This also does not appear to apply because the parasite population has not to date come near 'over-taking' the needle miner populations, and although i t has been shown 93 in the section on climatic control that there is a strong i n d i -cation of a reduced fecundity in the needle miner population this is obviously not a result of density-dependence in the sense used by Klomp since i t has occurred at low populations. The third damping mechanism is a density-dependent mortality of the host. In theory, a density-dependent mortality, l ike reproduction, would have a damping ef fect , provided the factor played a considerable part in mortality at intermediate stages. No such factor is operative in the needle miner populations. The factor which has caused the decline by means of catastrophic mortality showed no par t ia l i t y . From Tables XXX to XXXVI i t can be seen that the parasites of the needle miner have never taken much advantage of their 'environ-ment' , certainly never in the manner assumed in the theoretical models br ie f ly reviewed. The proportion of parasites to the host population has remained more or less constant since 1944-. Only the las t two generations, 1952-54- and 1954,-56 have shown a s i g n i f i -cant increase. On the basis of this we may postulate a similar sequence of events pr ior to 1944 which kept the parasite popu-lations at a low l e v e l . The climatic analyses presented in an ear l ier section apply equally well to the parasite population. Indeed, to explain the lack of success of the parasites we must postulate an effect which limited their population growth more than their host and kept i t more or less constant. The f i r s t effect considered was that there was a d i f fe rent ia l mortality of parasites from severe winter weather. That i s , the parasites suffered heavier mortality in the severe winters and/or 94. that t h e i r s u r v i v a l rates from 'normal' cold winters i s lower than the needle miner. Unfortunately we have no concrete evidence to support t h i s claim. The only measurement of parasite mortality ( a l l species) was made i n 1956 and was not s i g n i f i c a n t l y d i f f e r e n t from needle miner mortality. There i s considerable evidence i n the l i t e r a t u r e to support t h i s hypothesis however. Clausen (19) recognized, p a r t i c u l a r l y with introduced parasites, that d i f f e r e n t i a l mortality of parasites can be a l i m i t i n g factor to parasite success. The introduced species may not be able to withstand as low temperatures as i t s host. Dowden's work (25) i s an outstanding example of t h i s e f f e c t . Control of the oyster-shell scale by Aphytis mvtllaspidis i s usually e f f e c t i v e i n the mild-wintered Annapolis v a l l e y i n Nova Scotia but i s inneffective i n New Brunswick where winter temperatures f a l l to - 20°F or lower (68). Success of parasites of scale insects i n C a l i f o r n i a was l i m i t e d by low temperatures and extreme temperature fluctuations dn the winter (20,23). Uvarov (144) reviews several studies which show a d i f f e r e n t i a l mortality of parasites at low temperatures. Some in t e r n a l parasites are more sensitive to low temperatures than the host i n which they e x i s t . However, he also gives examples which show that low temper-atures can favor parasite populations where the parasitized hosts are less susceptible to temperature extremes than non-parasitized. B l a i s et a l . (8) found t h i s to be true f o r parasites of the forest tent c a t e r p i l l a r . Unseasonably warm weather i n May followed by several days of freezing temperatures caused high mortality of larvae but i t s p r i n c i p a l parasite was unaffected. 95. In the needle miner population the major parasites are internal and not evident u n t i l spring of the larva l year. In sections of dead larvae made in 1949 and la ter years the per-centage parasitism was in some cases higher than that observed in late instar larvae. However, we have the observation of parasitism made in 1956 to offset this evidence. It appears then that d i f ferent ia l mortality may be a factor which aided in keeping the parasite population of the lodgepole needle miner at a low leve l but the evidence does not permit i t s unqualified acceptance. A second l imitat ion to parasite success i s thought to be the d i f ferent ia l effect of weather on developmental rates. Uvarov ilUU) gives ample evidence to show that host and parasite are very often unequally adjusted to normal cl imatic conditions and w i l l react to disturbances in different ways. Sl ight differences in rate of development may have a profound effect on the success of a parasite part icular ly when the stage attacked i s short- l ived and the period of oviposition of the parasite i s equally short. Thus DeBach, et a l . (23) found that winter temperatures affected the rates of development of the parasite of a scale insect which disrupted the synchroniz-ation of parasite emergence and the stage attacked result ing in markedly reduced parasitism.. It has been shown above that the emergence dates of two of the major parasite species of the needle miner were markedly different in two different generations. I f the assumption i s correct that these species are parasites of the egg stage, then the numbers of these species may be expected to be less in 1958. Another possible factor , based solely on example and specu-96. l a t i o n , is that cold temperatures may reduce the fecundity and f e r t i l i t y of the parasite to a greater degree than i ts host. There is ample evidence to indicate that fecundity and f e r t i l i t y of insects are affected by external factors. In this thesis we have dealt largely with winter extremes but these effects may occur at any time during the l i f e cycle (144)» DeBach et a l . found that the fecundity of a parasite of the Cal i forn ia Red Scale was markedly reduced by detrimental effects of winter weather (23)• Thalenhorst (134) states that In nature, parasites and predators rarely attain their theoretical maximum eff ic iency and gives as a major reason for this that the hosts are rarely dispersed uniformly over an area, even under outbreak conditions. Differences in mobility and the 'searching a b i l i t y ' of the parasite may place the parasite at a d is t inct disadvantage. Environmental conditions during adult parasite ac t iv i ty may seriously l imi t their a b i l i t y to search out the habitat of the stage attacked by them. So many variables are involved in determining the success of parasitism in comparison to that of the host that i t is not surprising that parasites may be ineffective in control l ing populations of the host Insect. We have reached the general conclusion that the parasite complex of the lodgepole needle miner, R. s tark i has been kept at low levels of abundance during the past outbreak through a combination of factors which acted in a d i f fe rent ia l manner to the detriment of parasite success. The conclusions of Bodenheimer and Schi f fer (10) seem so pertinent to this discussion that a portion of them w i l l be quoted in f u l l . 97. "The fact that no accumulative summation of the p a r a s i t i c e f f e c t i n successive generations^- i s observed i n nature, means i n other words that the parasite i s , as a whole, always more sensitive to decimating factors than the host. This rather mystical state-ment i s , however, accessible to quantitative analysis. This analysis shows that t h i s higher s e n s i t i v i t y i s j u s t what had to be expected? a) . In addition to the hosts and parasites k i l l e d d i r e c t l y by the catastrophe, there survive always a number of parasites within hosts which have been k i l l e d by the catastrophe. In endoparasites which have not yet finished t h e i r growth, t h i s must lead to an almost 100 per cent mortality of these survivors. Ecto-parasites and predacious larvae w i l l be found to leave the dead host and to go i n search for another suitable prey. I t may be safely assumed that never 100 per cent of these migrants w i l l reach t h e i r goal, but that usually - especially at the low host density which prevails just after the catastrophe, a very important percentage of them w i l l die of starvation.^ b) . In many cases, the parasitized hosts w i l l c e r t a i n l y be more sensitive towards the catastrophic f a c t o r , increas-ing thus the host mortality as well as the type of mortality just described. c) . In a considerable number of cases the d i r e c t mortality of the parasite must actually be larger than that of the host. This has a p r i o r i to be expected for a good number of cases i f the r e l a t i v e s e n s i t i v i t y of hosts and parasites are distributed at random i t i s probably correct that the r e l a t i o n of host- and parasite- s e n s i t i v i t y towards the catastrophic factor (s) i s not at random, but that i n the majority of cases i t actually works to the unfavor of the parasites. From these reasons which are f u l l y confirmed by the mathematical analysis, we come to the conclusion that either the n a t a l i t y of the parasites must be very high or i t s mortality rather low as compared to that of the h o s t ; ; In order to overcome th i s unfavor-able e f f e c t . I f we assume an equal and at random s e n s i t i v i t y f o r host and parasites, the l a t t e r w i l l , by reasons a) and b) always be decidedly more reduced than the host. The main operative modus i s an ecological one independent of the higher physiological s e n s i t i v i t y under c ) . This analysis seems to give s u f f i c i e n t reasons to explain the absence of an accumulative effect of para-s i t i s m or - i n other words - why the parasite i s always more "sensitive" than the host." 98. 1. Referring to the Thompson theory that i f certain premises are f u l f i l l e d the parasite w i l l cause the extinction of the host by accumulative increase from generation to generation. Thompson recognizes that such does not occur i n nature but considers the hypothetical case of a r t i f i c i a l introduction of a parasite i n t o an hitherto unparasitized population. 2. This i s apparently what happened i n the spring of 1949-50. The two mortality estimates given i n Tables XXX and XXXI indicate that the parasites comprising the higher estimate must have completed t h e i r development to a stage where they were v i s i b l e i n hosts which were k i l l e d by the severe temper-atures i n January, 1950, before themselves succumbing. 3.3 Predfltfan 3.3.1 Types of predation. Predation was not an important mortality factor i n the out-break u n t i l 1956. No s i g n i f i c a n t loss i n eggs or larvae could ever be attributed to predation of any kind. Repeated f i e l d observations have never led to suspicion of predation of eggs but i n the detailed sampling carried on since 194-9 during the l a r v a l stage an in s i g n i f i c a n t number of shredded needles have been found, which were suspected of having been shredded by bird s . In 1956, i n two areas, Mount Girouard and Massive Mountain, there was found a large number of shredded needles such as could have been done only by flock i n g b i r d s . Tables XXXIV and XXXV show that the extent of the predation was 11.38 per cent at Mount Girouard and 33.60 at Massive. No d e f i n i t e information i s available but from a l i m i t e d knowledge 99. of the bird fauna the suspect predators were narrowed to three. These are: (1) The Black-capped chickadee, Penth$stes a t r icap i l lus  septentr ional ls. or possibly Gambol*8 chickadee, P. gambeli. Although no actual feeding has been observed chickadees are very common in the Bow Valley at times and have been seen f locking in mid-winter around Banff and Mount Eisenhower. Their habits and the high percentage of insect food they are accredited with needing (133) made them a log ica l suspect. (2) Canada Jay or Rocky Mountain Jay, Perlsoreus canadensis  can i ta l i s . This species is also very common in the outbreak area, as many as eight at one time congregating around the f i e l d stat ion. They have actual ly been observed pecking at mined needles and shredded needles have been recovered from their perching trees. As most of the establishments throughout the National Parks are closed during the winter and as the Canada Jay is supposed to shun larger centres of habitation (133) i t is possible that insects may be one of their winter staples. The Jay is omnivorous (133) so is a less l i ke ly suspect for mass predation than the chickadee. (3) Less l i k e l y to be involved in needle miner predation than the other two is the Junco, probably the Slate-colored Junco given the name Junco hvemalis oonnectens by Taverner (133). Taverner's description f i t s the species observed by the author but no specimens have been taken for ident i f ica t ion . Although feeding primarily on weed seeds Juncos have been known to eat insects. This has been included as a doubtful suspect because i t has been seen in f a i r l y large numbers in Banff National Park in early spring and late f a l l . 100. 3.3.2. Importance of predation in dynamics of needle miner popul- ations . This i s very d i f f i c u l t to assess. As has been noted i t has been immeasureable un t i l 1956 and then was very loca l i zed . The extent and importance of such predation over the whole outbreak area, or as a factor in causing population reduction i s l i k e l y very small . I t could however, assume an important role where needle miner numbers are low and restr icted to f a i r l y d is t inc t "refuge areas" such as was discussed in the climatic controls sect ion. The importance of bird predation either during an outbreak or during endemic periods must, however, be treated with caution. Unless the birds actually discriminate between parasit ized and unparasitized larvae their effect i s l i ab le to be more harmful than benef ic ia l . With the needle miner the poss ib i l i t y of discrimination is very remote as the important parasite species are internal and are not v i s i b l e as parasites (to external human observation at least) u n t i l late in the needle miner development. The long-range effect of reducing the parasite population in equal proportion to that of the needle miner could conceivably cause an increase in from needle miner numbers as i t removesAthe control complex a more p r o l i f i c control agent. That birds are continuous predators in many insect communities i s well understood (2) but i t i s also admitted that most bird predators are generally indiscriminate in their choice of prey species. It i s unl ike ly , therefore, that the needle miner i s the sole food at any time for any of the birds mentioned above or the v i s i b l e evidence of predation would be more common. It has been shown that 101. in an outbreak of a forest insect where insectivorous birds are present the birds w i l l eat a substantial and s igni f icant number but no estimate was made of the effects of indiscriminate preda-t ion of parasite and host al ike (26). Such predation as has been observed in the needle miner outbreaks would have a greater depressant effect on parasite populations than the host i . e . in the manner discussed by Bodenheimer and Schi f fer above (10). No estimate of predation of adult needle miner has yet been made. 3.4 Disease Disease has not been an important factor in the needle miner outbreak. An early report of "disease" larvae between 1945 and 1946 (51) is now believed to have been winter mortal i ty, a view in which the originator of the f i r s t report concurs (Hopping, G.R.-pers. comm.). Samples submitted to the laboratory of Insect Pathology at Sault Ste. Marie, Ontario, prior to 1952 showed no incidence of any disease. In 1952, workers at the Insect Pathology Laboratory succeeded In iso lat ing a virus disease from the Cal i fornia needle miner, R. m i l l e r i Bsk. At the suggestion of J.M. Cameron, Officer-in-Charge of that laboratory, an attempt was made to introduce the virus disease into the Canadian infestat ion (117). The supply of the capsufe virus in suspension was very l imi ted. Three concentrations were used: a) f u l l strength, with an appropriate amount of a wetting agent (methocel). b) di luted 1 in 10 of 1 per cent methocel. 102 o) diluted 1 in 100 of 1 per cent methocel. d) 1 per cent methocel was used as a control . The location chosen for the experiment was in the Cathedral Mountain sampling area. Five trees were selected, a l l of which were heavily infested. Twenty-five t ips on each of three trees and 25 in the remaining two trees were chosen on the basis of the number of available egg-laying s i tes i . e . abandoned, mined needles. A large number of these were desirable to guarantee a sat isfactory experimental population. Later, needles containing eggs were picked from surrounding trees and interlaced among the treated t i p s . Every t ip to be treated on each tree was stripped of needles for several inches back of the 1950 or 1951 fol iage l imi t ing the larvae to the desired part of the t i p . The stripped portion was painted an identifying color for each treatment. The capsule material was then applied by brushing It onto the green needles with a camels-hair brush. Only the top ha l f and the convex surface of the needles were painted to conserve virus material , since the f i r s t instar larvae almost invariably enter into the needle in this portion. A further precaution taken was to do the control f i r s t , to avoid contamination, and working up the scale of concentration, r insing the brush after each treatment. This was done for two reasons, to avoid alterat ion of the concentration and to perfect the technique so that when the f u l l strength was appl ied, waste would be at a minimum. Two samples of the treated material were submitted to the Laboratory of Insect Pathology at Sault Ste. Marie, one taken in 103. December, 1952, and the other taken in A p r i l , 1953. In addition to the experiment described above samples of l i v e and dead needle miner were submitted to the insect Pathology Laboratory in 1953 and 1954.. Since 1954 Hiss M.E.P. Cumming has kindly examined a portion of the needle miner population for disease each year. The f i r s t application was begun July 31 and completed August 9th. Larvae began to emerge in the f i e l d within a few days of completion. During the nine days of application there were a few l igh t showers and on the night of August 13th a very heavy r a i n f a l l . As eclosion and larva l establishment were not yet complete this may have affected the resu l ts . Results of the f i r s t sample of the treated t ips were,en-couraging. The t ips treated with the highest concentration showed the greatest number of larvae with 'many' capsules present. This was the only concentration however which showed a s igni f icant i n -crease. At the time of this sample,specimens from other locations in the outbreak area were also submitted. Capsule virus was found there for the f i r s t time (Cameron, J . M . - pers. comm.). The second sample from the treated area in A p r i l , 1953, was less encouraging. There appeared to be a s l ight tendency for increase in incidence in the f u l l strength treatment but an almost equally high incidence was found in the control and in samples taken upwind and downwind from the treated area. There was a noticeable decrease in the occurrence of the capsule virus from the f i r s t sample. However, there were indications that incidence of the virus in the experiment was in fact due to the introduction • 104. of the virus. The incidence of disease elsewhere was much lower. It must be pointed out that although the incidence of the disease was rather high in the treated area i t did not increase the actual mortality. The "dead" larvae examined by the Insect Pathology Laboratory were likely killed by dehydration of the needles or other artifacts owing to the relatively long period be-tween submission and examination at Sault Ste. Marie (April to July). The actual mortality in this locality at that time was about 12 per cent, which is largely attributable to winter k i l l . Examination of material collected in December, 1953, and January, 1954, indicated a high occurrence of capsule virus but again mortality was felt to be due to winter conditions rather than the disease. Results of investigations since this time indicate low occurrence throughout the outbreak area and probably very l i t t l e mortality, i f any, can be attributed to i t . There is l i t t l e evidence to suggest that the capsule virus disease isolated from the California needle miner is actually lethal to the larvae infected (Struble G.R.-pers. comm. (77)). Steinhaus (130) reviewed in considerable detail the knowledge of insect diseases. He outlines nine categories of relationships between micro-organisms and the main body of the treatise deals with the field we are interested in: "9. Insect as a definite host of microbial agents to which  i t is susceptible. In other words, the microbe-insect relationship may be that in which the microbe is a pathogen whose activity causes disease and frequently death in the insect host. The principal groups of microbial agents responsible for infectious diseases in insects are: viruses, bacteria, fungi, protozoa and nematodes. Examples of diseases caused by these 105. agents abound....The regular or periodic occurrence of disease among insect populations i t s e l f constitutes an ecological factor of great importance from the pract ica l standpoint as well as from that of insect ecology generally. The insect ecologist , in part icular should not lose sight of the fact that infectious disease i s a manifestation of parasitism. It represents the reaction of the insect to invasion of the animal's tissues by a micro-parasite. It i s simply a form of the struggle of l i v i n g beings for food, shelter and propagation as expressed in the host-parasite re lat ionship." It has been assumed since a virus disease was isolated from the Cal i forn ia needle miner and la ter from populations of R. starki that we are dealing with such a re lat ionship. The evidence does not support this assumption but on the other hand does not ent irely refute i t . In the re la t ive ly long history of the outbreaks of Recurvaria mi l le r i B s k. in Cal i forn ia (since 1911 at least) (93) there has been no indication that the disease caused any of the several population decl ines. These were thought to be rather the effect of overpopulation result ing in "ghost forests" (Struble, G.R.-pers. comm.). Yet the incidence of the virus in Cal i forn ia needle miner populations i s high enough to extract re la t ive ly large amounts of i t . Thus i t i s possible that the occurrence of the virus i s actually one of Steinhaus' other types of relationship not neces-sa r i l y detrimental to the host. However, the genera! concensus of opinion i s that the viruses are usually disease-causing and may be-come l e t h a l . Even i f we continue to accept the assumption there i s some doubt that the virus present in the needle miner population could become an effective control agent. A conclusion reached by Steinhaus wast " i f the proportion of result ing disease in the total population i s not great and satisfactory adaptation occurs, the mutant may thrive in the susceptible population and gradually the insect may develop a high degree of tolerance to the pathogen that i n i t i a l l y exploited i t . " 106 Yet another consideration which makes the success of virus disease in needle miner populations unlikely is the insects' be-haviour. It is d i f f i c u l t to conceive an e f f ic ient method of dissemination for an insect which spends most of i ts l i f e cycle (certainly the most susceptible stages) inside a protective needle. There is l i t t l e gregarious contact between needle miners, even in the same branch t i p . The only times when 'bodily contact' could serve as a means of dissemination would be immediately after eclosion be-fore the larvae had separated from the egg cluster to f ind their respective neediest during the second year larva l transfer when contact would be rare except in the case of extremely high popul-ations; and during the f i n a l la rva l transfer in the spring of the moth f l igh t year. No examination of adult needle miner for disease incidence has yet been made. A protozoan and polyhedral disease have been isolated from the abdomen of spruce budworm moths (86) and i t is possible that occurrence of disease in adults may cause reduced fecundity in females and lower f e r t i l i t y in males. As It appears certain that disease outbreaks are an "overpopul-ation phenomena" (130) and that diseases have a "threshold" of host density before they become effective i t is further possible that the populations of Canadian needle miner never reached a high enough degree of population density for the virus disease to reach outbreak leve ls . The virus disease present in R. starki. populations has apparently remained at the "enzootic" leve l throughout the course of the present outbreak. 107. 3.5. Other natural control factors. 3.5.1. Reslnatlon. Behaviour of f i r s t instar larvae of Recurvaria stark! Free, and R. m i l i a r ! Busck i s almost iden t i ca l . Upon eclosion they seek a green needle and begin to mine, almost invariably on the convex or outer surface of the needle (77,93,123). The few that attempt to enter the needle from the concave or Inner surface are usually k i l l e d . The abortive mine frequently has a small bubble of resin above i t with the carcass of the larva in i t . No measurement of la rva l loss in the Canadian population has been included in the mortality tables because mortality from this factor i s not s ign i f icant . In the Cal i fornia needle miner Morgan (77) states that "99 per cent of a l l f i r s t - i n s t a r larvae that entered needles on their concave ( f la t ) surfaces were k i l l e d by an excessive flew of r e s i n . " However, less than 0.1 per cent of the larva l population was affected. The only other reference to this phenomenon found was on the pine needle miner, Exotalala nlnifnllftll* (Chamb.) which attacks Jack pine, lodgepole and seven other species of thick-needled pines (6). Jack and pitch pine are i t s favored hosts in eastern North America but lodgepole pine i s severely attacked when planted in the insect 's range. Internal structure of the needle and suscept ib i l i ty to injury by the pine needle miner were correlated. Number, posit ion and siae of the resin canals and possibly tree vigor were the main factors involved. Jack pine and lodgepole pine contain only two moderately sized resin canals, situated one in each corner of the needle. In these two hosts, larvae of E. p i n i f n i l a l l * were able to feed with l i t t l e , i f any, interference from resin flow" (6). This does not 108. agree with observations made on the two Recurvaria species on lodgepole pine i n C a l i f o r n i a and western Canada. One possible explanation, other than differences i n tree vigor, l i e s i n d i f f e r -ences In mining behaviour. The Recurvaria species s t a r t t h e i r mine (whether on the concave or convex surface) i n the d i s t a l t h i r d of the needle whereas f i r s t i n s t a r larvae of E. n i n i f o l i e l l a enter the needles commonly near the base of the needle In the mid-line. The Recurvaria species also usually begin t h e i r mine i n the middle of the needle but are less select; mines are commonly found near the edges of the needles. We can only assume that the majority of those Recurvaria entering the f l a t surface do so i n too close proximity to the re s i n canals. In any event, as noted above, the mortality from t h i s cause i s n e g l i g i b l e . 3.5.2. Competition and ' Overpopulation' fac t o r s . Competition, as discussed here, w i l l only be concerned with the i n t r a - s p e c i f i c l e v e l . In the h i s t o r y of t h i s outbreak there has been no evidence to suggest that any other organism competes with the lodgepole needle miner for i t s food. 'Overpopulation factors' (106) have been credited by various authors with causing f l u c t u -ations i n forest insect pests. Among the phenomena credited to overpopulation are reduced fecundity, increased s u s c e p t i b i l i t y to disease, food shortage and increase of natural enemies. Disease s u s c e p t i b i l i t y and increase of enemies were not important factors i n the decline of the outbreak. One could assume, without r e a l evidence, that the population declined due to reduced fecundity caused by overpopulation (l06) or to a genetic collapse (29). How-ever, the cl o s e l y a l l i e d Californian needle miner, Recurvaria i r d l l e r i 109. Busck has repeatedly reached the saturation point of i ts environ-ment, causing large areas of forests to be k i l l ed without noticeable reduction in population prior to their own starvation and death. Survivors from those populations which destroyed their own environ-ment have caused outbreaks in new locations (77,93). Food shortage could conceivably be an important factor in popul-ation reduction but again in the Cal i fornian needle miner outbreak this has become a c r i t i c a l factor only upon the death of a large number of trees and populations continued to increase around these "ghost forests" (77). In the Canadian outbreak the population never quite reached the levels which would cause a c r i t i c a l food shortage. From intensive defol iat ion analyses begun in 1949 i t was apparent that populations did not persist at high levels long enough to cause complete defo l ia t ion. However, populations from 194-0 to 194-8 were high enough to cause about 80 per cent defol iat ion in some loca l i t i es and had these populations persisted, food shortage would have occurred in local ized areas (128). It is extremely unlikely that competition for food has ever been c r i t i c a l enough in the Canadian needle miner outbreak to have caused any fundamental population phenomenon to occur. However, a reduction in fecundity from a change in food quality as a resul t of intense defol iat ion may have occurred. There is l i t t l e evidence to suggest any change in fecundity prior to the investigative period, 194-8-1956, and only the evidence from inaccurate estimates based on dissection of female moths during this period. As stated above, the only apparent change was in 1954- and 1956. In this outbreak therefore competition is discounted as a s igni f icant population reduction factor . 4. EPIDEMIOLOGY 4.1. Epidemiology since 1942. In the above sections we have discussed the population density and natural mortality of succeeding generations since 1942. It has been established that the population density was high when the infestation was f i r s t noted but was restr icted to elevations between 5,000 and 6,000 feet . Following this year the outbreak spread to a l l al t i tude levels where lodgepole pine grows but did not increase appreciably in population density. In 1949-50 the popul-ation suffered a •catastrophic 1 mortality which reduced the popul-ations to low density (see F igs . 11,12). Most s igni f icant was the comparatively high populations which remained at elevations between 5,000 and 6,000 feet . The population was thus restr icted to the alt i tude levels where i t was f i r s t found. This indicates that the outbreak was probably 'young' in 1942, although numbers per t i p were considerably higher then. It has also been shown from studies begun in 1948 that the reduction in numbers in 1949-50 was largely due to adverse cl imatic factors , ch ie f ly winter temperatures. Other natural control factors , chief ly parasitism, could not have caused the declines noted. Subsequent population decline since 1949-50, while largely due to la rva l mortality during winter months, may have been accelerated in part by reduced fecundity in adult females, reduced f e r t i l i t y in males and/or other factors l imit ing ovipo-s i t i o n . These considerations led to the speculation that the origin of the outbreak was due to climatic conditions less severe than those which occurred during the period under investigation o r , the "theory of climatic release." 111. 4..2. Origin of the outbreak 4.2.1. The theory of climatic release The importance of climate in the epidemiology of insect out-breaks has been a subject of controversy for many years. Early "biot ic" theories placed the emphasis on biot ic factors but some authors recognized that weather factors may cause an "unbalance" which may lead to outbreaks (106). Later theories were more comprehensive (88,102,107,138). Nicholson (88,89,90) has long held the view that populations are in a state of balance and the main control l ing factors (of numbers) are "density-dependent" which include direct competition for resources or space, parasites, predators, and pathogens. Climate is "density-independent" and can never control populations. Andrewartha and Birch (3) hold that the factors of environ-ments control l ing numbers are numerous but that climatic factors are of major importance. They conclude that a l l factors are "density-dependent" and attach no specia l importance to the b io t ic factors which are affected by host density. Thompson (137,138,139) believes that natural control results from an organism l i v i n g in a continuously f luctuating environment. Under favorable conditions, numbers increase; under unfavorable condit ions, numbers decrease. Never do numbers increase indef in i te ly and, rarely ever, decrease to ext inct ion. Var iab i l i ty in population abundance tends to be inversely correlated with the complexity of the "ecosystem" (137) a view held by many authors (2,3,54.). Milne (75) reviewed the theories mentioned above and proposes his own which he describes as a "modification of Thompson's." He 112. objects to the Nicholson theory on the grounds that competing species, parasites, predators, and pathogens can not control because they are imperfectly density-dependent and to Thompson's theory that i t underestimates the importance of density-dependence. The Andrewartha and Birch theory also suffers from their treatment of density-dependence. Milne's own theory is that competition between individuals of a single species is the only perfectly density- dependent factor in nature. This factor is seldom evoked and there-fore the control of' increase is the combined action of factors , density-independent and imperfectly density-dependent. The control of decrease of numbers is brought about by density-independent factors . Ul lyet t (1A3) has cal led climate a "catastrophic" factor and thinks i t can be a contributory cause to insect outbreaks. This i s based on the assumption that "density-dependent" (biotic) factors are more adversely affected by such catastrophes than the insect in question. In the absence of these control l ing factors the insect may reach destructive densities when the catastrophe is spent. Thalenhorst (135) has presented several European examples of insect outbreaks attributed to weather conditions. From obser-vational evidence, he shoira that weather may influence an insect population in many ways. These are: acting d i rect ly on the popu-la t ion; by i ts effect on some other factor which is fundamental to population growth (or lack of growth); by acting on the other factor and the population simultaneously with a reciprocal ef fect between population and factor; or by a maze of interactions involving s o i l , 113. host plant, population and i ts enemies simultaneously, with interactions between the factors affected. He summarizes his paper by paraphrasing Wellington (153): "So far i t can be generally seen that weather factors (whether acting d i rect ly or indirect ly) may play a decisive role in the origin of mass outbreaks, part icu-l a r l y (and possibly even only) when certain meteor-ological phenomenon are repeated in successive years," The development of thought concerning weather in re lat ion to insect outbreaks has slowly given more importance to weather as a causal e f fect . However Wellington (152) has pointed out that a l -though the l i terature is studded with papers dealing with the effects of various meteorological factors on many phases of insect development and behaviour, only a few deal with those effects in terms of large scale weather processes and a very few follow through to the log ica l conclusion: prediction of the b io logica l phenomenon with the aid of modern methods of weather analysis forecasting. Wellington has developed probably the f i r s t inclusive theory re-la t ing insect abundance and weather (152), "Weather and climate are often considered simply as the broad framework within which the complicated b iot ic interactions take place. This viewpoint hastens the process by which numerous instances of the direct effect of meteorological factors are re le -gated to the limbo of density-independence so that the b io log ica l heart of the problem may be pursued without further d is t ract ion . Predictive systems lose a number of potential ly valuable facts in this way. More important however, this viewpoint leads to to ta l disregard of the indirect effects of meteorological factors on the equilibrium of a population by their action on i ts habitat, i ts parasites, i ts diseases, and the supply and quality of i t s food. To assess cl imatic influences correctly i t is necessary to examine cl imatic variations during the period immediately pre-ceding or coinciding with the beginning of an outbreak of an insect that exhibits violent fluctuations in numbers instead of studying the climate while the outbreak ex is ts . This follows from the concept of climatic release of a small indigenous population. That i s , in a region where a species exists in small numbers, and in which b iot ic conditions already favor population growth no i n i t i a l increase may occur u n t i l seasonal cl imatic control is relaxed. The important point to keep in mind, however, is that favorable weather may have to recur several years in succession before a major increase in population can develop. Once the enormous potential for increase that such a species possesses is rea l ized, the population grows so rapidly that no combination of adverse physical or biot ic factors can halt i t immediately. Since i t is usually during this period that the outbreak is studied, i t is not surprising that effects of the various or ig inal govern-ing factors are often obscured." The application of the theory is best dealt with from concrete examples. There are three reasonably detailed examples in Canada. Thus Wellington et a l . (154-) after distinguishing between those weather types favorable and unfavorable to the spruce budworm, related past outbreaks of the spruce budworm in central and eastern Canada to cl imatic changes. It Was shown that outbreaks were preceded by reductions, during three or four consecutive years, in the annual number of cyclonic centers passing through the affected areas and by reductions in June precipi tat ion. By la ter more refined weather analyses Wellington (151,152) has been able to show defini te short-term la t i tud ina l shi f ts in the movements of pressure centers over North America. Pressure centers are closely associated with known air-mass source regions and may be differentiated into groups depending on where they originate. When the tracks for each center are traced and studied independently, shi f ts in the pr incipal courses from one period to the next frequently show up. A southward d i s -placement of the tracks of those centers originating in the central States (Colorado lows) is associated with a corresponding southward displacement of the tracks of pressure centers originating over the polar region. Humid t ropical a i r masses w i l l be mostly barred from the Great Lakes region when such a southward sh i f t of the c i rcu la t ion 115. pattern occurs in the central part of the continent. The majority of a i r masses that pass over the region w i l l then be of polar or ig in . Thus Wellington concluded that in Northern Ontario the required physical conditions for spruce budworm increase tend to occur when the annual number of cyclonic passages in the late spring and summer is below average, and the majority of the a i r masses involved in these passages during these seasons are dry. They are of polar continental or polar maritime o r i g i n , because a southward sh i f t of the c i rculat ion pattern holds invasions of more southern a i r masses to a minimum. In the same study Wellington found that the required physical conditions for forest tent ca terp i l l a r population increase begin to occur with increasing frequency as the annual number of passing cyclones r ises to a maximum. During this period of increase in cyclones, the number of passages during spring and summer is above average, and the majority of a i r masses involved are of south-western or ig in because there is a northward sh i f t of the c i rculat ion pattern which moves the more northern a i r masses to higher lat i tudes. His f i n a l conclusion from these studies is that his findings place the problem of forecasting population increases of the spruce budworm and the forest tent caterp i l lar to possible outbreak levels on a meteorological basis that should f i t into the techniques for long-range weather forecasting that may be developed in the near future. The third example also concerns the spruce budworm in New Brunswick. Greenbank (37) considered outbreaks of the spruce bud-116. worm which occurred in 1912 and 194.9 In relat ion to the theory of climatic release proposed by Wellington. His results were confirmatory. While considering these works i t must be pointed out that the basis for the theory was not in the realm of abstract synoptic meteorology but from laboratory and f i e l d studies on the effects of meteorological factors on the behaviour of the adult , including mating and fecundity, on la rva l development and behaviour and i t s relationship to eventual fecundity, stand conditions, the effects of cl imatic factors on the flowering of balsam f i r and others, eventually related back to the causal effect of the meteorological factors in operation - the over-a l l macro-climate of the region. In summary, the theory of cl imatic release explains the time and place of outbreaks and i t s worth may be measured by i t s a b i l i t y to pre-d ic t outbreaks. It i s not a theory to explain "regulation" of an insect population at levels of abundance comparable to comprehensive theories. The purpose of the theory i s not to postulate regulation of population by climate. Studies on population dynamics in forest entomology during endemic periods are rare, although i t i s apparent that fluctuations in numbers without loss of balance are common and outbreaks the exception. Within the endemic period increase in population from one year to the next can resul t from physical conditions becoming favorable to the insect . Readjustment of the population after this increase may come through density-related processes although these may not be ent irely effective u n t i l physical conditions become favorable again. However, years with unfavorable weather conditions cannot always be expected to ' follow years with favorable conditions and eventually the favorable weather conditions recur several years in succession. During such a period, as the climatic theory postulates, the endemic population may 117. be released from the control l ing influences of both physical and b io t ic factors (153). 4.2.2. The theory of cl imatic release applied to the outbreak of the lodgepole needle miner, Recurvaria s tark i Free. (1) Climatic controls of the region. Weather for any part icular location is described as the sum tota l of i ts atmospheric conditions (temperature, pressure, winds, moisture and precipitat ion) for a short period of time or "the momentary state of the atmosphere" (140). Climate, on the other hand is a compilation of day-to-day conditions related to a part icular place or region with consider-ation given to variations of the various climatic elements (17,14-0). The weather of particular months related to particular mortality phenomena has been described. This weather, however, is related to speci f ic cl imatic conditions peculiar to that region. Dai ly , monthly, or yearly weather in any region is determined largely by type and c i rculat ion of a i r masses. In preceding sections we have described cl imatic elements with reference to the air-mass predomi-nant at the time. In Appendix 9 a summary of air-mass types is given for a considerable number of years for the outbreak region. When a mass of a i r remains stationary for some time over a region i t acquires properties of temperature and water vapor that are characterist ic for the area. In winter an a i r mass that remains for some time over the cold frozen area of north central Canada would become cold and re lat ive ly dry. When extensive differences in a i r pressure develop the a i r mass would begin to move from a high to a low pressure area. A i r masses can be c lass i f i ed and the system used most extensively is the Bergeron c l a s s i f i c a t i o n . In this system, 118. four pr incipal source regions are recognized: polar , a r c t i c , t ropical and equatorial . These are known by capi ta l letters P,A, T , and E. Lower case letters before the source le t ter further identify the a i r mass as originating over land or water and others may be used to describe a property such as temperature (14,0). Discrepancies in air-mass terminology arise from differences in opinion concerning source regions. Five main a i r masses have been used by most authors in describing the climate of western North America. This was the system used by Henson in his air-mass typing (4.3,14-1). There has been, however, a recent trend to consider fewer a i r masses as affect ing the weather of western North America. Penner (95) has summarized the work of his colleagues in the Meteorological Service of Canada whereby for pract ica l purposes, based on air-mass properties, four major a i r masses are recognized as affecting North America in the winter and three in the summer. The c lass i f i ca t ion was based on thermal properties at various pressure levels within the a i r mass. The four a i r masses involved are t ropical maritime (mT)j polar maritime (mP)j arc t ic maritime (mA) and arct ic continental (cA). In summer, the arc t ic continental is modified to arc t ic maritime. The only way that adoption of this system affects us is that the polar continental a i r repeatedly re -ferred to in the above discussions and in Appendix 9 must be considered equivalent to arc t ic continental . As we are largely dependent upon the Meteorological Service for our information and Climatic analyses and as this new c lass i f i ca t ion is becoming wide-spread in the various Meteorological off ices i t would seem ad-visable to adopt this system in our future work, i f we are to 119. understand climatic fluctuations and their relat ion to insect abundance synoptic concepts are essential whether we work from a i r -masses or air-mass properties. The chief advantage of a synoptic approach is that i t w i l l eventually be possible to place i t cn a long-range forecasting basis when more is understood of cl imatic var iat ions. The climate of the outbreak region is controlled by four main air-masses (mT,mP,mA,cA) In the winter and three (mT,mP,mA) in the summer. The main c i rculat ion is from the north and west which results in the predominance of mP and cA (mA) a i r . If we assume that the occurrence of the various air-masses is no more stable than weather and their variations are not necessarily random fluctuations about a mean It makes the assessment of climatic effects on insect populations simpler. The data given in previous sections in addition to that following give a reasonably c lear picture of the general climate of the outbreak region. (2) Climate and epidemiology of the,lodgepole needle miner. The effects of various winters on needle miner populations has been given in a previous sect ion. Figure 19 presents the mean month-ly temperatures for three winter months for Banff from 1920-21 to 1952-53. Mean monthly temperatures for the three months are shown for Lake Louise from 1932-33 to 1952-53 (Figure 20). It can be seen from these figures that conditions which caused heavy mortality in needle miner populations occurred with re la t ive ly high frequency. The largest gap between severe winters occurred from 1937 to 1950. As outlined above, the peak of the current outbreak was postulated to be from 194-0 to 1944. The outbreak was found in 1942, confined to the Figure 19. Mean monthly temperatures for December, January and February, 1920 - 1954. Banff, Alberta. MEAN MONTHLY TEMPERATURES - 1920 - 21 TO 1952 • 53 FOR BANFF, ALBERTA YEARS Figure 20. Mean monthly temperatures, December, January and February, 1932 - 1953. Lake Louise, Alberta. MEAN MONTHLY T E M P E R A T U R E S . 1932 - 1952 - 53 . L A K E LOUISE, A L B E R T A - DECEMBER h F E B R U A R Y • (Banff) 120. middle a l t i tudes. The fact that large populations vere found there indicates that the infestation was 'young 1j in the process of building up. Empirical calculations show that build-up of the needle miner population to numbers far in excess of those found could occur in three generations (six years) with a few reasonable assumptions. If we assume one f e r t i l i z e d female per branch t ip in the f i r s t year with an egg-laying capacity of 15 eggs and a series of mild winters where tota l mortality (including parasitism) did not exceed 20 per cent in any one year, the population in the s ixth year would be greater than 100 per t i p . Thus i t is possible, beginning with the gener-ation in 1938 that such a population growth could occur owing to the series of mild winters following 1936-37. The concentration of larvae per t i p did not occur at the intermediate levels probably because of the dispersal throughout uninfested stands at other al t i tude levels and val ley bottom (149). The series of winters described above, ch ief ly 1949-50, reduced the populations to levels which may have been present In 1938 or even ear l i e r . Such a series of mild winters should have an effect even on so gross a measurement as the yearly average temperature. The yearly average temperature for Banff was plotted with a calculated f i v e -year running average. (Figure 21, after Longley (67)). The running average shows a def ini te warming period from about 1925 to 194-8. As we have already seen however, there were several years pr ior to 1936-37 where the winters were comparable to 1949-50 which supports the assumption of the outbreak beginning about 1938. Evidence of a rea l cl imatic change not attributable to random fluctuations has been compiled and i t was shown that the climate of northern regions of the Figure 21. Annual mean temperature and five-^rear running mean, 1893 - 1955. Banff, Alberta. world did become warmer about 194-0 (152). From the weather records presented i n the text and Appendices i t i s u n l i k e l y that an out-break of comparable magnitude was able to occur p r i o r to that time. Tree r i n g studies do not show any evidence of a previous outbreak (84.,128). I t would follow from these observations that i n t h i s region the 'normal 1 climate i s too severe to permit outbreak popu-lations of Recurvaria s t a r k i Free- to occur f o r any length of time. (3) Geographical l i m i t a t i o n of the needle miner outbreak. The needle miner outbreak was r e s t r i c t e d to the valleys i n Banff National Park and adjacent areas i n Yoho and Kootenay Parks. The r e s t r i c t i o n of spread east or west may also be explained from the severity of climate (22), but t h i s i s less c e r t a i n i n r e s t r i c t i o n of westward extension. In Appendix 8 the mean monthly temperatures from 1939 to 1953 (winter months) f o r selected stations i s presented. The stations reported include three from the eastern slopes of the Rockies i n Alberta and one from the Columbia Valley i n B r i t i s h Columbia. The winter means show that the eastern slopes of the Rockies are generally colder than the outbreak area. This i s also demonstrated i n the Climatological Atlas for Canada (136). This i s due mainly to the f a c t that the outbreak area receives most of i t s cold a i r from the north while the eastern ramparts of the Rockies protect the valleys from cold a i r ori g i n a t i n g i n the east and north-east. Although the same c i r c u l a t i o n a f f e c t i n g the main outbreak areas i n Banff Park affects Kootenay and Yoho Parks, winter extremes are often not so severe but fluctuations i n climate are often more v i o l e n t . C i r c u l a t i o n i s much more complicated west 122. of the Great Divide (43)• The cause for res t r ic t ion on vest -ward extension i s probably violent fluctuations in climate af fec-t ing stages other than the larvae rather than severity of winter climate as in the east. U) General discussion of needle miner epidemiology with respect to future studies. In the previous sections i t has been shown that climate, pr inc ipa l ly winter extremes, has been the primary cause in the elimination of outbreak conditions of the lodgepole needle miner. It was further postulated, on the basis of cl imatological evidence that the outbreak developed during a warming period in the climate of the region. In studying the population the l i f e - t ab le approach -a quantitative one - has been used. However, no attempt has been made to apply any of the various theories of population growth or natural control "formulae" theories to the data. Natural or f i e l d demonstration of the various mathemetical theories or population "osci l la t ions" has proved very d i f f i c u l t (90,106). To attempt to apply such theories on the basis of one short outbreak would be spurious, only continued investigation at the low endemic levels w i l l lead to the formulation of sound theory. It has been cogently pointed out by various authors that very l i t t l e in the f i e l d of population dynamics i s beyond the hypothetical stage and & p r i o r i acceptance of mathematical concepts to describe short-term popu-lat ion studies, part icular ly when these are made at high levels of 123 abundance only, w i l l only lead to misuse and misinterpretation of valuable hypotheses (96,104,105). In a symposium discussing the merits of mathematical models used in population studies, Neyman et a l . (87) states "The focal objective of population ecology i s the understanding of those processes responsible for census-trends of species-popul-ations l i v i n g in natural habitats. Owing to the inherent complex-i t y of the processes this i s an objective far easier to state than f u l f i l l . Superimposed on this i s another complication. The environment i s character is t ica l ly variable and varying. The popul-ations respond s l i g h t l y , or markedly to this variat ion and, in so doing, may modify i t s environment. Such variat ion i s not under the control of the investigator although he may systematically record i t by i n i t i a t i n g in the f i e l d a sustained program of census, and environmental measurement - a program leading to an impressive accumu-lat ion of physical and b iot ic information. But the information i s l i k e l y to be d i f f i c u l t to analyze and even more d i f f i c u l t to generalize conceptually. Despite such handicaps however, this approach must remain the central one in the study of natural popul-ations for the self-evident reason that i t , of a l l others, d i rec t ly comes to grips with conditions and responses as they occur in nature. No implication i s intended that the direct approach lacks power: or indeed that i t s power cannot be increased. More Judicious col lect ion of relevant data, greater u t i l i za t ion of multivariate analysis, and in some cases, actual experimentation in the f i e l d , are valuable extensions that must f ind further perfection and adoption." Morris (80) has pointed out that many of the d i f f i c u l t i e s found in assessing population trends are due to inadequate interpretation of mortality data. Mortality and variations in individual mortal-i t i e s must be studied from the aspect of population trend rather than by generation. He shows that variations at high levels of mortality are potent ial ly more important than variations at low l e v e l s , although variable low mortalit ies may be more important than constant high mortal i t ies. Thus a variable low mortality contemporaneous with a re la t ive ly constant high mortality may be the determining factor in population trend. From the study of the past needle miner outbreak i t i s f e l t that 124. we do not have suf f ic ient information at the present time to merit an attempt at ' f i t t i n g 1 the data to any mathematical theories of population fluctuations now extant. Continued sampling of the now endemic population from the l i f e table approach is f e l t to be neces sary before this can be done. There is much to be learned yet about the relat ive effects of di f ferent environmental factors on the regulation of needle miner populations. This can be achieved by determining the population density as frequently as possible ( 7 ) . Continued sampling studies compared with continuous l i f e history and behaviour studies are therefore equally as essential as determining only fluctuations in population density (21). Perhaps the most im-portant consideration is that c r i t i c a l research must be carried out on the needle miner populations in advance of , or between outbreaks (18). Studies at low populations densities are of fundamental importance in determining the or ig in of needle miner, or any other insect , outbreaks. 125. 5. LITERATURE CITED 1. Anonymous Annual meteorological summaries. Can. Dept. Trans-port, Meteorological Division, Aviation Forecast Office, Calgary, Alberta. 2. Allee, W.C., A.E. Emerson, 0. Park, T. Park, and K.L. Schmidt. Principles of Animal Ecology. 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The ecology of lodgepole pine in Alberta and i ts role in forest succession. Canad. Dept. Northern Affairs and Nat. Res. , For . Res. Div. Technical Note No. 45. 1956. 60. Hutchinson, R.N. Influence of winter night temperatures on the Cal i forn ia Red Scale. J . Econ. Ent. 40(6): 921-922. 1947. 61. Klomp, H. On the theories of host-parasite interact ion. Proc. Twelfth Congress, IUFRO 56/24/10. 1956. 62. Leech, H.B. In, Annual Report of the Forest Insect Survey -Br i t i sh Columbia and western Alberta. Canad. Dept. Agr . , Div. En t . , Ottawa. 1943. 63. Ibid. 1944. 64. _ I b i d . 1945. 65. Ibid. 1946. 66. Leopold, A. Game Management. Scr ibner 's , New York. 1933. 67. Longley, R.W. Mean annual temperatures and running mean temperatures for selected Canadian stat ions. Canad. Dept. Transport, Met. D i v . , C i r c . 2481. Tec. 186. 1954. , 68. Lord, F .T . and A.W. McPhee. The influence of spray programs on the fauna of apple orchards in Nova Scot ia . VI. Low temperatures and the natural control of the Oystorshell sca le , Lepidosophes ulmi (L.) (Homoptera: CoccidaeT. Canad. Ent. 8 5 ( 8 ) : 282 - 291 1953. 69. Mc Guff In, W.C. In Annual Report of the Forest Insect Survey. Canad. Dept. A g r i c , Div. E n t . , Ottawa. 1948. 70* Forest Insect survey. Canad. Dept. A g r i c , Div. En t . , Bi-Monthly Progress Report 5(2) 3-4. 1949. 71. In Annual Report of the Forest Insect Survey, Canad. Dept. A g r i c , Div. En t . , Ottawa, 1949. 130. 72. McLeod, J.H. Notes on the lodgepole needle miner, Recurvaria m i l l e r l Busck. (Lepidoptera:Gelechiidae), and i t s parasites i n western North America. Canad. Ent. 83(11): 295-301. 1951. 73. M i l l e r , C.A. A technique for assessing spruce budworm l a r v a l mortality caused by the parasites. Canad. J . Zool. 33(1): 5 - 17. 1955. 7A. M i l l s , H.B. Weather and climate. In U.S.D.A. Yearbook, Insects. 1952. 75. Milne, A, The natural control of insect populations. Canad. Ent. 89(5): 193-213. 1957. 76. Mirov, N.T. Composition of turpentine of lodgepole and jack-pine hybrids. Canad. J . Bot. 34(4): 443-457. 1956. 77. Morgan, F.D. Factors influencing the abundance of Recurvaria m i l l e r l Busck (Lepidoptera:Gelechiidae). Ph.D. Thesis. University of C a l i f o r n i a , Berkeley, C a l i f o r n i a . 78. Morris, R.F. Technique f o r population sampling on standing trees. Canad. Dept. A g r i c , Div. Ent. Bi-Monthly Progress Report 6(6): 1-2. 1950. 79• The development of sampling techniques for forest Insect d e f o l i a t o r s , with p a r t i c u l a r reference to the spruce budworm. Canad. J . Zool. 33(4): 225-294. 1955. 80. _____ The interpretation of mortality data i n studies on population dynamics. Canad. Ent. 89(2): 49-69. 1957. 81* and C.A. M i l l e r . The development of l i f e tables f o r the spruce budworm. Canad. J. Zool. 32(4): 283-301. 1954. 82. Moss, E.H. Natural pine hybrids i n Alberta. Canad. J . Res. 27: 218-229. 1949. 83. The vegetation of Alberta. Botanical Review 21(9): 493-567. 1955. 84. Mott, D.G., L.D. Nairn and J.A. Coofci Radial growth i n forest trees and effects of insect d e f o l i a t i o n . Forest Science 3(3)* 286-304, September, 1957. 131. 85. Muesebeck, C.F.W., K.V. Krombein, H.K. Townes and othera. Hymenoptera of America north of Mexico. Synoptic Catalogue. U.S. D.A. Agriculture Monograph No.2, Washington, D.C. 1951. 86. Neilson, M.M. The measurement of spruce budworm mortality caused by disease. Ann. Tech. Rept. 1954-7 (Fredericton, N.B.). Canad. Dept. Agric. Science Service, For. B i o l . Div. Ottawa. 1955. 87. Neyman,J, T. Park and E.L. Scott. Struggle f o r existence. The Tribolium model: b i o l o g i c a l and s t a t i s t i c a l aspects. Proc. Third Berkeley Symposium on Mathematical S t a t i s t i c s and Probability V o l . XV. p. 41-79. 1955. 88. Nicholson, A.J. The balance of animal populations. J . Anim, Ecol., 2: 132-178. 1933. 89. ___________ Fluctuation of animal populations. Rep. Aust. N.Z, Ass. Advance S c i . , 26: 134-148. 1947 90. A n outline of the dynamics of animal populations. Austr. J . Zool. 2(1): 9-65. 1955. n 91. Nolte, H.W. Beitrage zur Epidemiologic und Prognose des Rapserdflohs (Psylllodes chrysocephala L.). B e i t r . Ent. 3(5): 518-529. B e r l i n . 1953. 92. Cdum, E.P. Fundamentals of ecology. W.B, Saunders. P h i l . Pa. 1953. 93. Patterson, J.E. L i f e history of Recurvaria m i l l e r ! Busck, the lodgepole needle miner i n Yosemite National Park, C a l i f o r n i a . J . Agr. Res. 21: 127-142. 1921. 94. P e a r l , R. and J.R. Miner. Experimental studies on the duration of l i f e . XIV. The comparative mortality of c e r t a i n lower organisms. Quart. Rev. B i o l . 10: 60-79. 1935. 95. Penner, CM. A three-front model for synoptic analyses. Quart J. Roy. Met. Soc. 81 (347): 89-91. 1955. 96. P h i l i p , J.R. 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Bi-Monthly Progress Report 6(6): 3. 1950. 104. Smith, F.E. Experimental methods i n population dynamics: a c r i t i q u e . Ecology 33(4): 441-450. 1952. 105. Solomon, M.E. Mortality required to prevent population increase. Nature, V o l . 159, p.848. 1947. 106. The natural control of animal populations. J . Anim. Ecol. 18(1): 1-35. 1949. 107. Dynamics of insect populations. In, Ann. Rev. of Entomology. Vol . 1 1 , p.121-142. Palo A l t o , C a l i f . 1957. 108. Stark R.W. Lodgepole pine needle miner. Canad. Dept. A g r i c , Div. Ent. Bi-Monthly Progress Report 4(6): 3. 1948. 1°9» Lodgepole pine needle miner. Ibid. 4(5): 3. 1949. HO. Lodgepole needle miner. Ibid. 5(1): 3. 1949. H I . Lodgepole needle miner. Ibid. 5(3): 3. 1949. 112. Annual Technical Report. Canad. Dept. Agr., Div. Ent., Ottawa. Mimeographed. (1948) 1949. 113. Ibid. 1950. 114. Parasitism of the lodgepole needle miner. Canad. Dept. A g r i c Div. For. B i o l . Bi-Monthly Progress Report. 7(6): 3. 1951. 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Theory of natural and b io logica l contro l . J_a Ann. Rev. of Entomology. V o l . 1. p.379-402. Palo A l t o , C a l i f . 1956. 140. Trewartha, G.T. An introduction to weather and cl imate. Mcffiraw-H i l l , New York. 1943. 141. Turnock, W.J. The climates of North America. Proc. Ent . Soc. Manitoba. V o l . 11: 19-22. 1955. 142. Preliminary l i f e tables for the larch sawfly. Canad. Dept. Agr ic . For . B i o l . Div. Interim Report, Winnipeg, Manitoba. Mimeographed, (1955-7) 1956. 135. 143. U l l y e t t , G.C. Mortality factors i n populations of Piute11a  macullpennis Curtis (Tineidae:Lepidoptera) and t h e i r r e l a t i o n to the problem of control. South A f r i c a Dept. Agr. For. Ent. Mem. 2(6): 77-202. l44« Uvarov, B.P. Insects and climate. Trans. Ent. Soc. London 79: 1-247. 1931. 145. Varley, G.C. The natural control of population balance i n the knapweed g a l l - f l y (Urophora iaceana) J . Anim. Ecol. 16: 139-187. 1947. ^46. Population changes i n German forest pests. J . Anim. Ecol. 18: 117-122. 1949. 147. Voute, A.D. On the regulation of insect populations. Paper presented at the Xth. Int. Congr. Ent. Montreal. August 1956. 148. Wellington, W.G. Conditions governing the d i s t r i b u t i o n of insects i n the free atmosphere. I. Atmospheric pressure, temperature and humidity. Canad. Ent. 77(1): 7-15. 1945. 149. Ibid. IV. Di s t r i b u t i v e processes of economic significance Canad. Ent. 77(4): 69-74. 150. Effects of radiation on the temperature of insectan habitats. Sci.Agr. 30: 209-234. 1950. 151. Air-mass climatology of Ontario north of Lake Huron and Lake Superior before outbreaks of the Spruce budworm, Choristoneura fumiferana (Clem.) and the forest tent c a t e r p i l l a r , Malacasoma d i s s t r i a Hon. (Lepidoptera:Tortricidae;Lasio-campidae). Canad. J . Zool. 30: 1LV-127. 1952. 152. ____________ Atmospheric c i r c u l a t i o n processes and insect ecology. Canad. Ent. 86 (7): 312-333. 1954. 153. Weather and climate i n forest entomology. Met. Monogr. 2(8): 11-18. 1954. 154. jJ .J . Fettes, K.B. Turner and R.M. Belyea. Physical and b i o l o g i c a l indicators of the development of outbreaks of the spruce budworm, Choriatoneura fUTI)1 fArana (Clem.) (Lepidoptera: To r t r i c i d a e ) . Canad. J . Res. D,28: 308-331. 1950. 136. 6. APPENDICES Origin of weather data* Broadly speaking, weather reporting stations f a l l into two main categories: (a) those stations which report hourly, 3-hourly, or 6-hourly, for forecast purposes and are generally equipped with instruments for recording the various meteorological elements, and (b) those cl imatological stations where obser-vations are taken once or twice dai ly and reports submitted to the d i s t r i c t o f f ice at the end of each month. Instruments at a l l o f f i c i a l weather reporting stations In Canada under the supervision of the Meteorological Branch are supplied by that organization. Every ef fort i s made to ensure that the observers are supplied with good quality instruments and the accuracy of the instruments i s checked from time to time by meteorological inspectors, or in the Instrument Division of the Meteorological Branch. The three stations used in the extensive analyses above are c lass i f i ed as follows: Banff (a) Synoptic Founded in 1887 Calgary (a) Synoptic- Aviation forecast " " 1876 Lake Louise (b) Climatological s tat ion. n n 1915 The other stations used, Golden, Rocky Mountain House, Edson and Exshaw are a l l type (b) stat ions. 1. Personal Communication, C .C . Boughner, Acting Director, A i r Services Branch, Department of Transport. September 12, 1957. APPENDIX 1 DAILY MAXIMUM AND MINIMUM TEMPERATURES FOR WINTER-MONTHS BANFF, ALBERTA. 1920 to 1954-. B A N F F , ALBERTA - 1920 - 21 November December J « m r y February Mflrch Day Max. Min, Max. Min. Max. Min. Max. Min. Max. Min. 1 44 14 37 22 29 20 34 19 39 32 2 41 31 38 29 33 26 33 22 37 21 3 36 30 38 31 29 14 32 21 42 29 35 15 38 28 23 12 26 2 40 26 5 34 15 29 13 30 12 19 - 9 33 12 6 31 5 28 12 24 14 22 - 8 35 26 7 35 8 29 20 20 10 35 17 32 2 8 33 12 25 6 30 12 35 30 41 12 9 30 4 28 10 28 11 36 21 38 17 10 28 13 32 23 22 6 36 28 18 5 11 18 - 6 29 21 16 - 6 37 26 6 -12 12 25 - 6 27 9 23 11 36 21 13 -28 13 22 - 3 26 22 27 18 21 14 17 -19 14 33 10 24 11 34 21 19 14 14 - 9 15 38 25 23 0 33 13 16 2 29 10 16 42 29 21 4 14 -13 17 -19 39 22 17 39 27 24 7 21 -12 19 -12 43 28 18 43 31 28 20 32 19 13 - 4 42 24 19 40 32 27 12 27 12 25 -14 34 7 20 39 30 15 1 25 5 30 0 26 -14 21 33 20 4 -12 22 8 30 - 4 38 2 22 34 25 - 5 -23 26 13 34 10 35 15 23 39 26 14 -16 24 8 45 28 41 24 24 37 26 24 11 27 - 2 48 33 38 28 25 38 29 19 4 28 16 45 39 36 25 26 33 22 15 -12 33 5 42 25 26 7 27 35 26 35 13 34 24 46 26 39 0 28 35 28 36 30 29 7 41 36 45 32 29 39 30 34 24 18 -13 40 24 30 40 30 39 29 26 5 43 15 31 33 20 31 10 45 25 BANFF, ALBERTA - 1921 - 22 November December January February March Day Max. Min. Max. Min. Max. Min. Max. Min. Max. Min. 1 45 34 26 2 24 12 15 -21 38 9 2 51 36 26 19 25 14 18 3 41 23 3 56 36 34 12 16 - 9 20 8 37 26 4 49 28 34 23 11 -10 21 7 34 19 5 49 40 26 15 17 - 6 22 - 5 31 14 6 43 27 32 12 20 4 28 13 31 - 2 7 28 19 29 12 20 3 25 2 31 8 8 34 12 32 19 39 16 3 -15 40 22 9 39 24 35 28 39 21 - 1 -11 41 31 10 42 31 39 29 26 7 - 7 -24 37 25 1! 42 29 39 33 26 6 -15 -30 33 14 12 38 34 40 30 30 15 - 2 -31 29 23 13 36 29 41 27 32 12 18 -19 30 14 14 31 17 28 3 33 10 20 -16 35 0 15 27 15 13 - 2 19 - 1 24 - 3 38 9 16 16 6 14 - 7 18 0 34 17 36 13 17 10 - 6 14 - 5 2 -22 33 0 35 20 18 8 -14 9 -13 11 -28 30 0 38 9 19 4 -19 -11 -32 14 2 22 -21 37 28 20 3 -20 -17 -41 14 1 26 -11 36 5 21 - 5 -24 16 -27 15 3 22 - 2 32 24 22 7 -11 13 - 1 4 -16 4 - 8 30 8 23 25 -10 0 -23 15 -14 13 -27 24 10 24 22 1 8 -17 22 1 24 -24 30 8 25 33 9 8 -14 28 13 28 -16 24 0 26 34 23 5 -19 27 19 24 -25 12 1 27 34 28 22 - 3 27 8 32 -19 29 -21 28 31 21 24 3 9 -20 38 -12 35 -12 29 31 21 31 22 0 - 9 40 14 30 31 24 21 11 - 7 -35 41 10 31 14 -11 - 2 -36 42 13 BANFF, ALBERTA - 1922 - 1923 NnvpmhAT» Dftcfimhar January February Marsh. Day Max; ' " k i n . Max;- Min. Max. •Min. Max. ' • -Min. Max. - Min. 1 36 16 25 7 28 12 20 - 4 31 22 2 34 6 23 8 30 12 15 -21 23 14 3 35 5 11 - 3 34 24 26 2 27 -12 36 24 6 - 9 17 10 36 13 27 2 5 38 16 - 8 -29 26 5 38 25 31 6 i 6 36 6 - 2 -17 30 12 40 16 38 14 7 34 5 - 4 -25 33 23 37 21 38 25 8 38 20 - 4 -24 33 27 26 8 35 14 9 36 17 - 9 -16 33 25 24 11 31 9 10 31 5 -12 -19 30 22 13 -24 36 2? 11 30 3 -13 -36 29 20 6 - 9 37 22 12 38 2 - 6 -29 25 4 - 5 -22 29 - 1 13 37 19 -14 -33 24 16 -14 -43 25 - 5 14 38 0 -32 22 2 -10 -41 30 - 8 15 40 23 3 - 9 30 17 26 -15 42 8 16 39 34 1 -27 36 25 37 - 7 37 23 17 38 27 3 -16 35 11 38 24 30 - 8 18 32 9 13 - 2 11 - 8 45 30 43 8 19 29 12 31 11 23 - -10 42 13 42 31 20 33 9 37 26 27 19 42 12 35 21 21 33 23 40 31 11 2 V- 5 31 6 22 36 72 39 34 16 15 26 13 39 11 23 38 16 39 38 17 3 42 32 39 5 24 39 2D 38 32 20 - 3 40 29 43 26 25 41 26 38 31 18 - 3 36 16 43 18 26 42 33 35 27 21 4 42 17 50 34 27 38 28 39 28 19 - 9 50 25 54 24 28 32 16 37 27 3 - 2 54 32 54 28 29 17 - 1 32 20 - 2 -17 39 28 30 19 10 27 8 10 -21 60 26 31 26 12 U -21 62 21 BANFF, ALBERTA - 1923 - 1924 November December January February March Day Max. Min. Max. Min. Max. Min. Max. Min. Max. Min 1 39 31 28 0 - 4 -35 40 30 43 20 2 38 31 30 20 - 5 -29 33 18 35 3 3 44 31 38 20 -11 -32 32 9 32 18 4 49 30 38 30 9 -12 34 10 40 10 5 48 27 30 25 22 4 38 26 47 27 6 48 17 31 23 29 21 36 13 46 17 7 47 16 30 24 34 26 35 16 45 10 8 47 17 23 4 32 15 30 13 44 4 9 45 21 26 15 25 4 34 4 40 13 10 44 22 36 20 28 20 39 22 40 18 11 41 17 31 22 24 8 36 30 38 16 12 42 18 22 - 2 19 - 2 40 32 42 15 13 42 22 24 - 5 25 10 37 21 41 8 14 45 26 32 17 27 11 22 4 37 28 15 42 23 40 28 25 1 18 3 34 7 16 40 24 41 33 2 -20 30 - 8 39 - 1 17 42 29 40 33 6 -12 36 2 41 14 18 37 23 39 33 3 -25 33 8 39 13 19 39 33 25 14 8 -24 16 - 6 36 8 20 28 20 24 5 16 1 34 -10 32 16 21 25 8 24 13 24 12 35 18 27 7 22 34 22 29 17 22 5 40 9 31 - 5 23 37 25 29 11 28 - 4 41 10 35 6 24 38 29 27 9 9 -10 44 23 34 9 25 35 23 20 6 28 - 1 45 20 37 2 26 33 20 14 2 36 17 43 29 32 24 27 36 17 22 5 38 26 47 29 37 14 28 34 27 13 0 39 29 42 27 40 9 29 32 22 -15 -18 40 30 42 20 35 21 30 24 7 -15 -27 41 32 33 6 31 -15 -40 45 34 39 11 BANFF, ALBERTA - 3924 - 1925 Novftmber TteRgmher January F ^ h n w r y M a r c h Day Max. Min. Max. Min. Max. Min. Max. Min. Max. Min, 1 2§ 20 22 14 16 21 -12 45 2 B 2 3 So 39 H ii 8 9 « % 3B 30 4 23 18 34 22 25 16 36 23 38 30 5 22 1 25 13 29 17 36 27 38 12 6 22 5 9 1 25 14 36 17 35 5 7 37 9 18 - 1 21 11 33 12 39 - 3 8 17 - 1 14 - 7 19 - 2 30 0 39 - 1 9 11 -11 24 2 22 3 36 17 24 15 10 27 - 4 30 19 27 15 30 13 32 - 7 11 10 - J 43 28 22 4 31 19 32 19 12 15 -15 39 32 19 6 34 0 14 1 13 26 10 35 25 4 - 9 31 - 1 30 -22 14 29 17 32 -14 5 -18 16 8 33 14 15 34 18 -12 -31 18 -12 19 -15 38 24 16 42 26 -33 -47 26 10 37 - 4 40 24 17 31 21 -34 -54 2? 17 34 14 35 6 18 28 21 -10 -40 36 25 38 24 38 19 19 38 27 -10 -31 30 21 41 26 38 22 20 38 29 -21 -40 36 27 31 20 43 23 21 33 28 -18 -42 30 25 43 U 46 2? 22 33 24 4 -22 42 23 42 13 38 34 23 34 21 9 -12 36 16 41 32 38 18 24 36 20 6 - 4 16 - 4 36 4 43 33 25 30 24 16 - 1 16 -14 12 -16 37 27 26 28 5 10 -14 22 - 1 37 - 3 41 21 27 30 18 5 -28 32 19 39 26 50 18 28 35 20 14 4 31 10 39 8 51 25 29 39 25 17 10 34 3 43 17 30 36 21 3 - 5 - 6 -14 45 27 31 26 - 1 -10 -20 50 27 BANFF, ALBERTA - 1925 - 1926 November December January February March Day Max. Min. Max. Min. Max. Min. Max. Min. Max. Max. 1 40 17 47 35 26 15 27 3 52 24 2 37 13 39 24 24 17 27 1 57 24 3 22 16 30 21 23 37 12 55 21 4 32 1 37 26 18 8 a 28 42 29 5 u 29 42 30 25 15 38 28 38 26 6 a 19 43 34 34 15 37 27 42 10 7 38 15 39 29 36 26 36 27 44 16 3 34 18 38 31 33 17 a 30 46 11 9 39 30 32 23 26 19 43 27 50 14 10 38 22 36 25 21 10 43 28 50 14 11 42 27 43 30 26 9 34 28 44 29 12 38 26 41 25 26 11 29 24 44 20 13 38 17 32 12 33 16 27 10 50 24 U 35 15 30 14 24 17 32 - 1 53 16 15 36 24 33 24 32 20 35 10 59 23 16 39 26 32 28 29 23 23 14 42 25 17 a 32 35 21 31 23 34 1 45 24 18 45 31 34 22 14 2 36 19 43 30 19 46 37 31 10 21 1 39 29 48 25 20 31 20 19 7 22 7 a 20 50 33 21 25 6 17 - 1 16 - S 38 15 47 33 22 37 15 31 17 27 3 37 26 54 39 23 36 29 31 22 24 16 32 18 40 32 24 37 27 38 19 26 4 37 22 36 22 25 33 28 42 34 31 14 38 22 39 15 26 29 12 39 17 35 18 44 32 38 23 27 20 - 2 27 16 27 20 47 28 33 8 28 23 7 27 12 30 17 45 22 37 13 29 29 11 29 10 28 14 42 23 30 37 18 27 13 36 22 34 11 31 27 8 31 7 34 15 BANFF, ALBERTA - 1926 - 1927 November December January • February March Day Max. Min. Max. Min. Max. Min. Max. Min. Max. Min. 1 50 23 38 29 39 33 27 2 40 4 2 45 9 5 36 3? 36 9 a 21 3 51 21 14 - 3 33 26 32 19 39 11 4 51 28 21 - 5 27 15 29 5 35 - 1 5 49 31 27 18 16 4 30 3 33 10 6 43 36 22 10 20 -10 28 1 37 25 7 45 29 28 5 30 16 22 -13 43 19 8 42 21 26 8 26 10 28 - 9 31 22 9 44 32 33 19 16 - 4 26 -10 34 12 10 47 32 41 25 24 2 32 7 32 4 11 44 34 30 9 17 - 8 31 18 36 2 12 45 34 - 5 -11 5 - 1 27 10 43 31 13 40 25 -20 -37 2 -20 31 3 38 28 14 40 28 - 7 -33 17 -19 1 - 6 27 11 15 36 25 7 -17 28 14 - 5 -u 30 15 16 34 14 17 1 21 - 3 - 7 -19 34 8 17 32 22 17 4 - 6 - H 27 -30 34 9 18 22 10 25 - 2 - 5 -29 31 7 32 - 6 19 10 0 31 20 - 8 -37 35 25 40 - 5 20 22 - 9 33 23 -11 -38 34 26 46 24 21 23 8 25 19 1 -27 37 27 41 33 22 33 13 7 - 7 5 - 6 36 12 40 22 23 25 20 19 -17 18 2 36 13 40 21 24 16 10 20 8 23 13 33 8 38 21 25 8 3 19 4 25 3 31 4 38 7 26 19 - 1 22 11 30 18 37 21 37 15 27 29 8 25 17 35 20 31 15 44 3 28 29 13 37 17 33 23 37 - 2 43 11 29 24 4 36 29 32 20 47 15 30 38 1 44 28 27 10 48 22 31 42 30 26 6 32 28 BANFF, ALBERTA - 1927 - 1928 November December January February March Day Max. Min. Max. Min. Max. Min Max. Min. Max. Min. 1 31 12 34 2 - 1 -31 30 •i 39 4 2 42 2? 34 8 16 -29 37 *.• i P 5 3 a 36 34 19 23 10 36 24 45 12 4 34 27 40 31 37 21 45 23 43 17 5 30 20 25 2 39 28 44 27 36 15 6 27 24 - 7 -?2 38 30 43 31 39 7 7 24 11 - 8 -24 45 27 37 17 16 9 8 23 7 - 9 -24 49 35 35 13 24 3 9 28 8 - 6 -34 51 39 39 23 40 19 10 10 4 - 1 -27 42 38 47 28 40 27 11 9 - 9 10 - 4 39 33 36 26 38 25 12 7 - 5 7 - 3 35 30 30 8 35 13 13 7 1 - 1 ! -26 25 15 32 13 32 9 14 5 - 3 2 -22 22 14 28 7 39 8 15 15 - 8 12 - 5 21 - 3 35 4 42 7 16 31 - 2 18 0 24 4 40 21 46 28 17 38 4 19 - 1 26 5 37 15 50 16 IB 14 8 20 13 20 10 43 16 57 25 19 39 2 22 8 14 4 34 17 53 28 20 2 r 7 20 7 14 - 7 37 8 55 31 21 28 - 4 10 -10 19 1 14 6 54 32 22 31 17 12 - 9 20 2 15 -24 45 35 23 30 21 19 7 13 - 5 22 -20 42 28 24 32 24 22 7 19 - 4 30 -14 38 28 25 34 15 29 15 17 - 4 36 - 3 39 21 26 21 - 5 32 24 27 6 37 3 42 15 27 30 9 16 - 6 35 19 35 14 34 25 28 25 19 - 9 -17 35 27 30 12 39 22 29 27 - 2 - 9 -16 27 22 35 - 5 42 20 30 26 2 -20 -35 33 11 42 24 31 -22 -45 30 7 36 29 BANFF, ALBERTA - 1928 - 1929 November December January February March Day Max. Min. Max. Min. Max. Min. Max. Min. Max. Min. 1 39 5 25 15 23 10 0 -14 40 30 2 42 11 8 - 2 21 3 5 -11 41 29 3 43 12 0 -24 22 7 24 -20 43 33 •i 43 34 11 - 7 9 - 2 20 -18 47 36 5 39 28 20 5 17 - 3 11 -11 35 16 6 42 27 22 7 26 14 15 -26 29 10 7 a 20 34 16 27 15 20 -13 35 20 8 36 20 38 30 28 17 26 -13 38 4 9 39 30 37 32 32 12 28 6 48 17 10 38 29 35 27 36 25 28 2 46 33 11 48 44 31 18 43 18 31 9 41 15 12 35 30 25 8 36 21 34 9 45 8 13 39 21 24 10 34 24 33 15 48 14 14 28 1! 15 - 3 18 3 33 3 52 14 15 32 5 20 - 1 22 12 37 27 52 17 16 34 24 23 9 21 - 3 13 5 47 15 17 29 9 26 10 22 9 4 -15 42 29 18 30 8 17 4 6 - 8 12 -27 48 20 19 33 17 21 6 14 -14 25 4 49 24 20 40 31 27 13 10 - 1 27 13 43 32 21 50 36 24 15 6 - 2 26 1 47 24 22 45 33 27 18 - 6 -31 28 12 38 28 23 36 23 38 22 2 -32 34 13 26 14 24 33 12 29 14 1 -25 36 20 35 4 25 36 11 33 16 - 4 -18 28 18 37 15 26 37 23 29 15 -13 -21 27 1 42 30 27 30 16 23 13 -20 -31 32 7 45 30 28 35 6 15 -12 -20 -50A4 35 1 45 31 29 31 19 26 8 - 9 -35 n 23 30 28 13 28 7 6 -27 30 8 31 25 8 0 -31 29 5 -58 at Lake Louise BANFF, ALBERTA - 1929 - 1930 November December January February March Day Max. Min. Max. Min. Max. Min. Max. Min. Max. Min. 1 49 29 2l 11 2 5 8 38 27 18 -16 2 39 32 30 11 26 U 37 31 30 -14 3 39 15 33 15 25 7 31 9 38 2 A 43 32 35 17 - 5 -12" 38 23 41 15 5 32 8 40 23 -14 -24 38 32 37 23 6 37 21 31 7 -10 -32 30 23 36 - 1 7 40 20 13 6 4 -22 38 5 36 12 8 46 27 5 0 2 - 6 23 19 36 20 9 40 33 - 4 - 8 - 7 . -31 27 1 37 25 10 30 16 6 -12 - 2 -30 32 19 46 30 11 29 2 7 -14 - 6 -30 25 7 42 36 12 32 9 - 7 -16 - 5 -30 21 0 23 13 13 43 24 2 -20 4 -27 23 9 21 - 3 14 42 28 21 - 4 - 6 -12 29 -17 28 6 15 45 36 22 0 -19 -27 a 15 33 - 6 16 32 29 7 - 1 -21 -5lfcfr 43 33 41 - 4 17 30 23 4 - 5 2 -26 46 38 45 9 18 18 3 9 -16 8 - 9 37 34 51 27 19 19 -13 15 - 4 -10 -32 38 31 9 - 2 20 15 0 10 - 1 7 -34 a 27 31 -14 21 19 -11 22 - 5 10 -15 38 31 34 21 22 26 13 29 16 12 -12 36 8 39 27 23 27 6 33 24 17 -10 38 25 36 14 24 33 18 37 29 18 - 2 33 10 40 17 25 u 28 40 32 13 - 1 28 - 3 44 27 26 44 37 32 22 7 -17 25 15 43 29 27 43 37 27 19 14 -10 23 6 56 30 28 29 22 38 24 15 1 13 2 60 24 29 38 19 37 33 25 5 40 31 30 34 35 27 26 15 42 12 31 25 17 33 21 52 17 -60 at Lake Louise BANFF, ALBERTA - 1930 - 1931 November December January February March Day Max. Min. Max. Min. Max. Min. Max. Min. Max. Min. 1 51 31 33 24 28 20 42 12 44 36 2 54 33 37 29 30 19 41 15 47 30 3 50 36 37 27 33 25 41 14 47 34 4 43 32 37 28 33 18 36 7 25 18 5 50 26 33 18 35 23 37 16 35 6 6 46 27 30 24 32 25 32 25 35 21 7 50 29 33 23 22 8 27 — 1 38 9 8 52 26 39 27 19 - 1 31 - 5 39 15 9 47 37 33 13 27 11 36 14 16 7 10 50 37 35 25 31 18 36 23 6 - 3 11 38 33 31 20 22 4 35 13 11 - 7 12 26 22 28 17 22 4 38 10 38 5 13 14 6 31 25 25 12 42 10 28 11 14 16 - 9 30 14 36 22 42 13 44 4 15 22 5 37 21 31 23 39 7 45 26 16 27 9 33 25 26 8 36 25 46 29 17 24 9 28 16 30 14 34 24 43 27 18 25 1 28 13 27 9 35 27 41 29 19 27 12 28 1! 23 7 39 28 44 32 20 32 19 28 7 32 15 35 24 50 33 21 34 16 28 3 25 15 35 14 50 39 22 40 24 28 17 37 21 42 25 43 28 23 28 12 29 20 42 32 36 26 44 22 24 41 23 26 8 37 26 30 16 43 20 25 36 20 25 14 34 14 35 2 15 7 26 32 21 28 15 35 24 33 11 19 - 8 27 27 15 27 8 35 29 36 20 21 -11 28 28 15 27 15 47 33 39 15 42 - 4 29 33 20 26 10 51 u 41 19 30 35 28 26 11 44 26 43 22 31 29 15 44 16 50 33 BANFF, ALBERTA - 1931 - 1932 November December January February March Day Max. Min Max. Min. Max. Min. Max. Min. Max. Min. i 50 4? S IS - n 39 % '** 3 39 21 29 23 20 - 3 16 4 16 - 1 4 42 28 30 22 25 9 24 8 15 - 7 5 51 27 28 18 25 1 24 18 12 - 1 6 45 36 23 14 31 20 9 2 6 - 7 7 40 33 28 6 36 19 10 - 9 6 -15 8 32 18 18 8 38 27 29 - 4 17 -23 9 33 21 24 1 36 29 31 26 26 -14 10 33 16 21 11 35 21 28 15 34 -12 11 34 12 12 - 8 33 24 20 0 44 -11 12 37 27 16 - 4 - 5 -12 12 - 7 45 7 13 30 27 16 - 2 - 6 -26 16 -23 43 20 U 7 19 11 - 2 -22 24 - 1 48 28 15 s 5 - 8 24 11 11 -15 23 10 32 23 16 5 - 9 35 16 15 - 6 24 -14 38 14 17 13 - 9 33 25 16 - 8 35 8 43 29 18 20 - 5 40 29 23 10 33 9 42 26 19 23 11 38 31 29 12 32 15 44 24 20 14 6 34 22 28 5 29 13 40 25 21 14 -17 29 9 28 9 38 - 6 38 20 22 24 9 26 12 23 4 42 26 u 17 23 28 4 25 8 22 0 47 38 42 27 24 26 18 29 16 35 13 44 36 38 30 25 23 18 29 18 34 21 46 36 42 25 26 14 - 5 22 2 21 - 3 40 33 39 25 27 19 - 6 25 16 22 4 43 36 39 28 28 24 5 25 34 1 - 8 46 33 30 19 29 24 11 21 ' 11 - 4 -28 33 20 38 8 30 23 10 16 - 5 -11 -22 42 28 31 21 9 -13 -25 43 33 BANFF, ALBERTA - 1932 - 1933 November December *anuary February March Day Max. Min. Max. Min. Max. Min. Max. Min. Max. Min. 1 34 14 35 22 27 16 22 - 3 25 0 2 39 27 36 20 24 10 20 - 9 40 - 6 3 34 25 35 30 23 4 32 9 35 12 A 37 28 38 29 30 20 33 18 30 1 ?5 39 30 19 13 31 26 28 7 38 19 '6 38 29 0 -11 31 25 0 -20 42 27 7 34 11 - 4 -29 33 26 0 -12 29 21 8 34 23 - 7 -23 35 26 - 4 -24 11 3 19 29 18 2 -18 35 30 2 -30 22 -16 10 26 7 5 - 7 24 14 7 - 8 25 - 5 11 29 9 8 -23 30 12 11 - 4 37 8 12 32 12 11 2 37 26 7 -12 39 29 13 32 - 4 11 1 40 25 9 -16 37 20 14 9 -12 11 - 7 32 26 18 - 7 37 7 15 11 - 5 17 - 1 r5 - 4 24 7 39 4 16 32 0 18 - 1 8 -19 29 1 44 13 17 39 32 20 0 6 - 1 27 23 42 26 18 45 10 15 5 8 -17 27 3 39 10 19 42 27 24 11 11 - 5 29 19 39 16 20 30 16 31 26 19 7 35 25 42 32 21 30 18 29 20 23 6 35 28 36 25 22 33 25 32 25 22 22 33 22 32 19 23 40 25 33 21 26 14 31 19 31 19 24 40 31 26 14 27 9 29 15 34 10 25 29 7 26 11 22 3 37 25 37 6 26 a 32 29 18 22 - 3 33 25 40 4 27 44 37 26 16 20 13 26 15 42 19 28 46 36 26 18 20 - 7 13 5 43 29 2?yO 38 33 25 19 18 - 7 42 26 30 37 26 25 7 21 0 43 35 31 29 20 21 1 39 27 BANFF, ALBERTA r 1 9 3 3 - 1 9 3 4 November December January February March Day Max. Min. Max. Min. Max. Min. Max. Min. Max. Min. 1 34 17 40 30 - 5 -15 42 34 40 30 2 32 25 44 29 35 -12 40 29 40 33 3 30 16 31 26 37 31 30 12 33 29 4 34 9 24 14 37 29 31 5 29 23 5 36 15 31 19 32 28 38 3 26 8 6 34 26 31 24 23 4 37 13 23 12 7 37 19 28 16 25 1 44 23 34 4 8 35 13 26 16 28 11 40 34 40 18 9 43 23 25 12 39 17 37 25 43 9 10 47 38 6 - 6 32 26 37 18 51 24 11 49 40 0 - 6 26 20 42 30 51 30 12 48 32 - 4 -11 28 2 43 25 51 35 13 57 a - 2 -13 31 23 42 16 43 19 48 33 - 3 - H 24 20 a 16 45 34 15 45 27 - 4 -13 22 - 2 39 12 43 35 16 44 30 21 -23 35 17 43 11 27 14 17 36 28 31 12 34 21 35 20 40 16 18 44 31 6 -10 21 1 38 11 47 35 19 45 35 23 -12 32 10 34 7 46 37 20 40 33 39 - 7 33 24 22 14 41 28 21 44 33 28 5 33 25 17 - 3 33 21 22 47 34 13 - 7 31 24 19 - 8 32 8 23 49 41 - 3 -15 24 2 8 - 4 34 4 24 47 37 - 5 -31 22 -16 6 - 7 33 9 25 a 35 - 7 -44 23 14 15 -25 33 11 26 47 36 8 -35 42 21 24 - 8 48 7 27 26 18 6 -16 46 39 33 17 27 18 28 26 9 28 - 9 43 13 39 27 17 8 29 34 20 38 22 39 24 48 3 30 38 30 - 3 -15 41 20 53 31 31 - 6 -19 42 29 46 29 BANFF, ALBERTA - 1934 - 1935 November December January February March Day Max. Min. Max. Min. Max. Min. Max. Min. Max. Min. 1 47 38 32 25 37 19 46 32 39 27 2 37 31 28 16 13 - 5 47 34 38 28 3 38 30 29 16 34 - 6 46 29 37 13 4 41 30 29 6 29 26 u 25 7 - 2 5 44 30 30 17 27 -10 33 -13 33 - 8 •6 43 35 32 12 30 21 30 18 28 -34 7 44 29 35 34 20 8 31 - 2 31 - 9 8 39 33 36 15 15 - 4 36 - 2 29 - 3 9 39 20 36 17 20 1 40 5 32 9 10 43 21 32 17 27 15 38 13 30 - 6 11 46 23 35 26 18 4 32 16 37 23 12 47 30 32 26 - 4 -14 33 16 u 30 13 53 33 37 28 - 7 -31 32 9 45 35 34 50 29 39 30 5 -26 36 18 40 30 15 52 36 33 25 9 -12 36 19 34 13 16 43 29 30 20 0 -14 40 34 35 27 17 a 26 33 26 -21 -29 43 29 38 21 18 39 24 32 24 -23 -42 37 19 36 17 19 33 20 35 26 -17 -43 28 12 36 21 20 29 15 32 24 - 8 -42 42 21 33 2 21 29 9 29 24 - 5 -17 40 26 28 - 5 22 38 23 24 4 16 -23 a 28 32 0 23 42 30 4 - 6 33 -12 23 34 34 19 24 37 27 -10 -30 40 28 28 0 34 28 25 37 29 -16 -42 42 3 32 - 2 30 17 26 31 20 0 -22 42 32 35 - 1 29 16 27 30 16 - 7 -30 44 36 35 0 30 4 28 25 8 9 -18 46 33 39 22 28 9 29 30 19 11 - 8 43 29 24 2 30 34 21 14 - 5 43 24 24 -10 31 32 - 6 43 29 17 - 4 BANFF, ALBERTA - 1935 - 1936 A .ft * A A ft, vs tat S3t November December January February March Day Max. Min. Max. Min. Max. Min. Max. Min. Max. Min. 1 11 -19 31 32 21 8 -20 45 30 2 M -20 30 n 35 18 6 -25 48 37 3 - 8 31 12 30 12 6 -24 43 38 •4 32 17 30 13 26 0 4 -38 44 28 5 36 23 39 . 11 9 -18 - 4 -28 46 23 6 42 26 36 34 38 -12 -16 -28 43 24 7 44 32 39 29 32 13 -12 -45 40 26 8 37 34 38 34 32 21 1 -21 41 28 9 16 - 3 38 13 29 15 13 -17 34 23 10 24 0 39 32 24 5 15 - 2 32 6 11 29 17 37 28 22 10 0 -17 34 2 12 24 7 36 25 25 0 -10 -35 u . 23 13 24 4 32 34 26 - 2 - 4 -38 37 26 14 30 1 26 13 16 - 1 - 5 -24 34 13 15 31 34 23 15 7 - 9 - 2 -38 38 13 16 38 26 24 15 4 -11 - 5 -36 39 21 17 34 24 25 15 5 -13 2 -36 39 19 18 26 0 19 2 1 -12 12 -28 38 20 19 34 11 30 3 21 - 4 20 -15 44 26 20 41 30 30 10 29 13 22 -10 40 23 21 38 26 34 15 38 21 21 - 6 36 23 22 27 17 32 22 41 30 16 0 31 30 23 35 15 24 - 6 39 28 4 -34 34 11 24 34 20 24 - 1 32 21 3 -19 32 - 6 25 38 27 31 16 22 - 9 4 -29 32 - 3 26 38 27 36 22 22 - 4 24 -11 33 7 27 43 32 34 22 20 4 39 19 29 9 28 44 33 36 27 19 -16 40 0 38 - 6 29 39 35 28 10 20 -31 44 28 15 - 5 30 33 19 25 34 22 7 17 -34 31 33 31 20 - 2 34 - 9 4 * Banff not recorded, readings from Anthracite. BANFF, ALBERTA - 1936 - 1937 November December £2~'"'cf February March Day Max. Min. Max. Min. Max." Min. Max. Min. Max. Min. 1 19 - 9 36 27 3 -29 - 6 -12 40 8 2 30 -11 28 -19 -17 - 2 45 17 3 31 22 16 5 26 1 8 -26 44 30 4 34 22 - 5 -10 16 - 2 3 -12 54 31 5 13 - 1 7 -25 - 4 -12 15 -14 49 34 6 18 -17 11 -25 - 5 -40 18 - 6 43 28 7 30 - 5 28 1 1 -27 2 - 8 46 12 8 41 19 26 8 13 -I4 7 -33 48 12 9 40 20 26 3 7 - 2 17 - 7 49 18 10 40 27 31 H 15 - 6 27 9 50 31 11 a 29 30 12 16 - 4 29 19 37 26 12 52 33 30 24 20 5 25 18 34 19 13 56 44 35 24 - 1 -10 28 - 2 26 12 14 49 29 29 25 3 -28 30 11 30 - 3 15 50 27 10 0 2 -16 30 17 37 21 16 46 26 33 - 8 8 -25 34 22 43 10 17 48 31 32 25 12 - 2 28 19 43 9 18 56 38 45 26 6 -11 24 8 40 12 19 59 44 34 20 -12 -33 19 4 35 20 20 38 22 35 13 - 1 -33 18 -11 24 11 21 45 15 42 16 3 -14 23 5 29 - 7 22 40 28 40 30 14 - 4 37 3 33 7 23 43 18 26 17 12 4 34 3 39 8 24 39 14 16 4 10 -11 24 15 36 21 25 39 17 20 0 16 - 3 24 10 38 20 26 41 12 13 2 2 - 7 30 -11 46 2 27 29 12 7 -15 - 5 -18 40 -14 48 5 28 37 11 - 4 -21 -11 -26 37 2 37 13 29 38 23 10 -13 - 3 -36 43 10 30 35 U 10 - 2 10 -17 43 31 31 - 2 -14 2 -14 44 23 BANFF, ALBERTA - 1937 - 1938 November December January February March Day Max. Min. Max. Min. Max. Min. Max. Min. Max. Min. 1 43 40 15 26 5 34 22 6 - 7 60 12 2 8 30 5 34 22 7^  -14 33 26 3 45 28 23 13 33 7 30 - 9 25 19 4 41 21 27 2 27 5 34 16 26 9 5 44 26 36 22 22 7 7 - 9 29 18 6 36 29 33 22 24 0 7 - 9 41 4 7 36 24 5 - 8 28 2 9 -12 u 3 8 40 29 -10 -23 31 13 24 -23 42 3 9 36 30 9 -29 35 15 0 7 -25 a 15 10 29 24 29 -11 32 23 8 - 9 a 26 11 31 26 36 20 16 - 4 6 -19 40 32 12 23 13 20 7 28 11 - 5 -11 39 25 13 15 2 34 5 34 20 - 3 -16 50 27 14 21 0 37 25 34 27 - 3 -17 44 35 15 18 6 36 19 34 28 7 -28 40 28 16 20 5 36 30 23 3 25 -24 40 27 17 17 11 36 30 23 10 28 - 5 36 20 18 13 1 35 19 23 0 28 12 34 13 19 26 -18 33 10 27 6 34 - 5 33 25 20 33 12 39 20 30 - 9 34 5 32 3 21 35 21 28 16 35 23 40 21 31 5 22 35 15 18 8 33 21 39 5 33 8 23 40 28 0 -20 27 8 40 5 38 10 24. 36 25 - 3 -15 23 6 46 9 44 19 25 3? 30 8 -20 33 16 50 19 46 18 26 31 8 24 -20 29 20 53 24 44 29 27 28 - 2 6 -15 43 18 56 12 43 36 28 34 15 37 -11 7 0 54 11 37 27 29 30 11 36 31 - 6 -26 28 10 30 25 6 35 25 0 -33 37 20 31 33 24 7 -20 34 0 BANFF, ALBERTA - 1938 - 1939 November December January February March Day Max. Min. Max. Min. Max. Min. Max. Min. Max. Min. 1 36 24 41 32 36 32 19 -15 32 9 2 41 20 34 25 40 32 21 - 6 32 15 3 41 30 34 25 37 28 20 1 13 - 1 4 33 19 36 22 34 19 20 -12 27 -22 5 32 15 39 30 20 5 30 8 32 - 8 6 31 8 36 30 26 0 - 6 -14 36 13 7 40 26 40 32 37 21 -13 -24 36 17 8 ' 33 26 47 32 35 24 -14 -29 36 - 1 9 33 10 28 22 32 15 4 -36 36 - 8 10 27 12 22 8 34 10 11 -30 34 2 11 23 - 5 18 -11 37 26 22 0 33 19 12 27 15 22 0 33 13 32 15 22 8 13 30 20 26 14 36 26 31 22 18 1 14 39 26 26 16 28 8 36 25 36 -18 15 39 31 17 3 28 18 33 10 33 0 16 38 26 25 1 28 9 37 7 36 - 3 17 35 22 22 4 30 20 42 30 47 4 18 40 30 21 5 33 22 34 21 50 35 19 35 26 18 - 4 37 28 25 - 4 51 40 20 28 8 18 - 5 30 15 25 - 7 56 46 21 21 1 21 - 1 19 -10 32 -11 60 46 22 14 - 8 24 0 27 14 39 4 62 35 23 20 - 6 41 19 31 18 38 21 61 47 24 27 14 41 32 30 21 40 13 56 46 25 26 - 2 12 - 1 27 18 32 24 42 20 26 28 11 - 2 -25 27 - 1 27 - 2 33 11 27 35 9 - 3 -18 30 17 30 1 48 6 28 35 17 -11 -22 31 12 28 2 54 18 29 34 12 29 -24 27 8 52 18 30 41 30 33 - 1 12 4 47 27 31 35 26 18 -10 50 39 BANFF, ALBERTA - 1939 - 19^ 0 November December January February March Day Max. Min. Max. Min. Max. Min. Max. Min. Max. Min. 1 51 23 43 26 31 18 23 - 7 2 49 21 44 31 35 24 20 7 3 45 31 47 39 26 9 25 - 3 4 41 31 47 38 18 11 38 15 5 42 31 47 27 31 14 38 31 6 41 31 42 33 22 3 26 22 7 36 26 46 35 16 - 7 33 18 8 ' 36 27 45 37 9 -10 36 8 9 34 26 34 32 21 -10 43 31 10 39 28 38 27 25 15 35 28 11 44 32 36 27 18 6 33 10 12 52 37 34 24 19 r-12 31 0 No 13 48 26 36 15 24 10 35 6 14 48 37 38 21 30 - 3 41 12 15 50 41 40 29 33 - 1 33 - 4 record 16 50 40 34 25 14 - 3 35 4 17 49 35 28 20 12 - 5 38 23 18 45 30 27 5 5 -22 28 21 f o r 19 45 27 34 20 0 -24 30 10 20 42 27 35 26 5 -24 31 9 21 46 27 31 26 22 -10 21 13 March 22 43 33 20 5 16 - 3 7 1 23 40 24 8 - 5 2 -18 11 -18 24 39 20 7 -14 - 2 -29 29 -14 25 35 18 2 - 6 15 -30 11 0 26 37 17 9 -14 26 3 6 - 7 27 34 15 12 -16 38 19 40 - 6 28 42 22 26 9 46 32 40 10 29 45 36 36 5 49 28 44 20 30 43 34 34 23 31 11 31 27 1 27 - 1 BANFF, ALBERTA, « 1940 - 1941 November December January February March Max. Min. Max. Min. Max. Min. Max. Min. Min. Min. \ 41 21 21 6 20 0 44 29 16 10 I 34 20 40 2 22 8 42 35 37 4 3 34 19 45 33 14 - 2 39 23 36 14 4 33 14 u 26 40 -11 42 21 44 16 5 28 5 39 31 12 - 5 40 15 45 9 6 21 10 36 29 22 2 39 4 44 22 7 9 3 36 30 24 - 5 37 2 49 34 8 3 -11 36 30 41 9 39 9 41 25 9 4 - 6 31 17 44 22 36 8 38 6 10 6 -18 19 6 51 38 39 21 41 4 11 4 -28 15 2 45 32 36 14 40 2 12 14 -17 15 -16 40 26 32 18 u - 2 13 21 0 13 - 9 38 27 34 8 48 - 3 14 31 6 19 - 3 23 15 32 - 2 48 15 43 21 14 - 4 22 14 38 3 39 8 16 36 15 21 -10 15 - 1 38 3 46 1 17 36 15 33 -13 31 2 39 3 53 24 18 30 11 36 9 36 25 30 14 44 27 19 28 6 40 22 38 17 21 10 45 29 20 27 11 42 30 30 12 18 1 a 22 21 21 7 41 26 21 - 6 6 - 3 44 14 22 23 - 6 48 25 20 0 6 - 9 45 18 23 26 10 34 16 - 1 -11 11 -31 44 27 24 31 21 32 22 5 -21 19 -12 52 20 25 33 21 36 20 36 -12 27 0 52 25 26 32 18 36 26 36 2 37 2 47 32 27 32 18 30 16 43 27 43 30 54 19 28 35 18 31 16 44 34 45 32 54 20 29 32 23 29 11 35 16 53 23 30 34 18 25 10 46 27 54 27 31 24 14 41 25 44 31 BANFF, ALBERTA - 1941 - 1942 November December January February March Day Max. Min. Max. Min. Max. Min. Max. Min. Min. Min. i g S a 28 35 xl -2| % 20 8 fo 2$ 3 36 25 40 27 6 -22 36 10 35 23 4 44 28 27 22 - 6 -26 34 22 40 26 5 46 34 32 14 7 -21 26 16 34 24 6 43 25 36 27 - 8 -26 32 11 33 7 7 41 20 36 29 5 -24 33 10 42 21 8 49 27 34 28 4 -14 33 8 45 26 9 47 26 15 11 20 - 8 32 - 3 42 29 10 49 24 24 2 31 14 34 12 38 13 11 47 35 26 14 33 20 35 18 41 21 12 38 23 19 0 39 24 36 9 37 22 13 39 16 18 2 36 24 34 4 33 2 14 43 32 29 6 30 17 39 13 37 11 15 37 26 33 10 32 9 29 12 37 10 16 34 19 43 24 33 14 10 -16 37 14 17 25 10 30 24 31 8 23 -26 38 28 18 27 3 38 15 33 15 28 - 7 36 9 19 30 11 33 24 30 16 32 6 35 0 20 36 15 32 27 31 8 18 11 42 21 21 19 17 22 9 34 6 13 0 48 33 22 23 - 7 St 9 36 2 10 -12 33 22 23 24 6 32 23 34 12 16 -21 24 6 24 38 15 25 21 -40 21 24 -11 29 - 5 25 42 29 15 5 38 24 28 -14 31 - 2 26 44 38 - 2 -21 38 21 30 - 6 36 - 4 27 40 20 38 -20 38 2$ 35 -13 44 - 3 28 44 18 15 -16 33 26 \ 36 12 52 - 1 29 52 38 - 5 -18 26 0 51 8 30 44 40 - 1 -20 28 12 54 21 31 - 4 -38 27 - 4 57 25 BANFF, ALBERTA - 1942 - 1943 November December January February March Day Max, Min. Max. Min. Max. Min. Max. Min. Max. Min. 1 35 19 9 - 9 6 -13 24 10 25 - 8 2 33 7 14 4 6 -20 32 19 35 -10 3 27 20 19 - 4 10 -10 29 9 37 6 4 19 7 9 - 6 15 -10 30 17 6 - 5 5 21 - 1 3 -16 23 1 32 18 13 -28 6 29 2 10 -10 33 6 36 22 28 -14 7 29 - 2 7 -17 30 18 36 - 4 26 6 8 26 1 19 - 7 36 24 1 -29 27 4 9 28 - 2 23 6 41 22 19 -28 31 - 6 10 38 21 34 12 36 21 28 14 30 16 11 44 27 34 18 26 0 37 23 30 15 12 41 26 40 29 32 9 43 12 36 21 13 52 29 40 32 44 29 47 29 12 2 14 43 34 39 31 40 30 46 28 9 - 9 15 38 29 41 27 38 -12 47 24 13 -24 16 25 17 39 32 - 5 -30 43 22 19 -21 17 11 6 37 18 -14 -47 49 18 27 -21 18 19 - 4 30 3 - 9 -33 48 22 27 - 8 19 27 2 34 9 -21 -30 43 32 33 5 20 24 4 34 21 -26 -36 46 19 37 - 2 21 34 15 34 29 -23 -44 31 24 42 22 22 40 29 29 25 - 4 -32 41 4 44 30 23 30 27 12 1 - 4 -36 29 0 47 33 24 30 13 24 - 3 7 -35 34 - 1 45 39 25 23 6 24 - 3 4 -18 44 8 34 24 26 20 5 14 -10 20 - 7 45 10 30 20 27 12 - 5 19 -10 21 - 1 48 9 40 22 28 14 -10 32 2 28 8 39 22 35 26 29 23 - 6 30 - 3 18 - 8 26 22 30 28 6 24 - 2 21 -14 38 19 31 16 2 21 1 41 20 BANFF, ALBERTA - 1943 - 1944 November December January February March, Day Max. Min. Max. Min. Max. Min. Max. Min. Max. Min. 1 32 4 32 16 26 12 32 9 31 -14 2 38 18 38 27 20 12 36 18 30 - 2 3 38 27 40 31 19 0 34 14 20 1 4 39 23 27 16 18 0 20 6 18 0 5 39 26 26 - 6 20 - 3 34 8 25 - 2 6 34 9 34 8 18 - 8 40 26 36 3 7 36 11 31 22 25 8 33 24 44 3 8 39 25 26 2 27 16 30 - 1 44 28 9 40 11 31 6 25 - 2 16 1 52 35 10 43 28 29 10 19 - 4 30 -13 48 20 11 41 22 29 19 24 - 3 31 18 29 5 12 42 16 ^ 21 - 2 28 16 30 0 28 10 13 42 23 27 7 35 25 28 14 23 - 5 14 42 23 30 10 40 24 21 11 40 0 15 42 20 32 8 38 29 28 - 5 46 15 16 46 23 29 14 40 28 28 16 47 33 17 45 30 34 17 40 34 29 - 4 44 32 18 34 16 33 28 38 31 32 7 40 20 19 40 25 13 - 8 50 36 31 - 5 39 26 20 40 19 21 - 4 38 28 28 8 38 18 21 45 31 26 9 33 10 20 7 34 7 22 36 22 26 6 41 15 28 -12 38 26 23 30 13 30 16 37 23 33 6 34 15 24 40 23 36 23 34 22 31 15 16 - 4 25 38 9 34 25 24 9 34 0 25 - 5 26 38 U 27 9 23 - 3 33 5 20 - 5 27 32 19 27 9 23 - 4 31 - 3 28 -15 28 32 10 23 10 26 - 6 15 8 45 6 29 34 25 10 -3 23 2 20 - 7 53 23 30 38 25 20 - 1 21 - 6 51 25 31 25 6 30 1 47 25 BANFF, ALBERTA - 1944 - 1945 November December January February March Day Max. Min. Max. Min. Max. Min. Max. Min. Ma::. Min. 1 48 22 34 26 28 - 4 28 27 8 2 32 19 32 20 29 9 18 § 34 9 3 26 19 29 8 32 10 19 3 12 8 4 42 21 39 24 32 18 40 12 13 -24 5 48 32 44 25 36 26 28 14 25 -14 6 45 35 42 36 36 10 34 1 28 10 7 42 30 33 24 33 1 38 28 32 2 8 46 32 28 18 39 22 46 32 37 18 9 45 33 24 6 42 32 33 17 40 28 10 30 20 29 6 37 18 33 17 34 26 11 31 18 26 4 40 32 36 13 36 23 12 34 15 22 8 38 30 2o n 40 28 13 28 - 1 32 8 34 24 27 i i 34 18 14 34 1 26 13 33 19 16 - 7 38 22 15 32 14 26 4 27 8 0 - i i 38 20 16 40 12 24 10 31 16 15 -26 35 4 17 37 14 28 9 32 20 16 -25 37 5 18 32 10 23 10 18 13 27 -16 39 5 19 38 7 24 8 13 - 8 36 - 2 41 30 20 32 16 24 - 6 19 -12 32 14 44 34 21 38 24 20 6 28 2 31 2 40 27 22 42 31 0 -17 33 11 32 16 46 28 23 4° 28 8 - 9 34 14 35 5 44 28 24 32 25 - 4 -17 2? 3* 39 >S 53 27 25 28 10 11 -14 32 8 43 10 42 18 26 31 15 13 1 10 - 4 38 18 48 24 27 34 14 18 - 2 23 -10 31 22 46 17 28 28 6 30 10 23 3 35 4 44 30 29 28 7 21 4 20 -15 44 22 30 36 24 13 4 20 -14 44 37 31 0 - 9 11 -15 30 23 BANFF, ALBERTA - 194-5 - 1946 November December January February March Day Max. Min. Max. Min. Max. Min. Max. Min. Max. Min. 1 32 19 27 6 33 25 26 3 38 28 25 18 30 10 35 23 2 41 30 51 31 39 31 36 13 - 4 40 19 1 36 25 40 26 36 26 21 - 7 38 34 5 15 8 36 28 32 19 20 -15 34 23 $ 6 - 7 29 22 31 10 26 12 31 20 7 3 - 7 25 15 28 22 28 - 5 34 31 k 7 -26 4 - 3 30 12 29 34 38 24 9 15 - 7 15 -12 30 17 30 20 40 7 10 22 - 1 20 6 30 22 25 1 49 ?6 11 28 - 2 13 4 21 1 15 -34 41 32 12 19 7 2 -17 28 9 26 2 38 28 13 31 4 15 -14 34 21 34 19 37 38 14 24 8 14 0 32 13 44 26 36 8 15 33 - 2 16 4 40 20 36 27 34 18 16 31 - 2 15 2 32 8 34 2 36 6 17 33 14 11 - 4 37 26 37 9 46 3 18 30 22 11 -15 39 28 37 25 48 26 19 30 12 17 5 31 21 34 - 1 41 29 20 26 2 19 13 21 -31 40 9 50 12 21 23 - 3 0 - 6 30 31 40 15 57 15 22 31 34 2 - 9 33 20 39 26 50 20 23 35 16 31 - 2 28 17 4» 25 41 28 24 36 21 31 21 25 16 45 30 39 24 25 37 22 31 21 9 - 3 38 28 40 30 26 32 24 31 10 16 -10 36 11 40 30 27 31 16 36 20 27 10 38 26 30 24 28 25 3 38 27 24 8 38 28 33 17 29 35 20 38 26 16 1 42 25 30 30 12 34 22 20 -13 44 28 31 34 22 24 31 46 14 BANFF, ALBERTA - 1946 - 1947 November December January February March Day Max. Min. Max. Min. Max. Min. Max. Min. Max. Min. 1 2 % 22 1 f? -£ - 3 -19 -13 11 - 2 -34 -20 1 8 - 1 3 42 24 39 25 - 3 12 -27 6 - 9 4 47 21 44 27 24 11 36 8 13 -31 5 46 26 40 30 29 - 4 42 26 22 -26 6 45 30 38 27 32 3 24 8 26 -20 7 35 24 38 29 31 13 23 -20 32 -16 8 33 11 35 21 28 12 20 -24 33 8 9 34 9 24 3 40 19 21 -23 40 6 10 35 23 44 22 36 22 36 -12 31 1 11 33 9 23 5 37 28 35 20 32 —4 12 36 10 3 - 8 18 10 39 29 40 - 3 13 39 15 4 - 8 1 - 5 46 30 49 25 14 37 18 15 -12 0 -21 45 30 56 33 15 25 18 21 - 3 15 -11 43 23 59 24 16 22 5 11 -18 33 8 38 4 57 22 17 8 3 14 -18 40 28 33 21 56 22 18 - 2 - 7 17 7 40 27 34 13 54 20 19 - 6 -14 35 12 u 12 25 4 56 18 20 - 8 -24 35 20 25 8 34 - 4 55 24 21 6 -27 34 22 39 20 u 20 55 37 22 17 -12 17 8 54 31 44 27 43 26 23 27 8 34 3 a 33 42 31 35 10 24 30 4 42 25 43 27 34 25 37 5 25 34 18 43 34 43 24 17 9 26 24 8 38 14 25 16 26 -11 27 33 19 - 6 -15 20 - 5 25 12 28 29 10 10 -31 17 - 7 24 -18 29 31 10 10 - 2 -11 -19 54 - 2 30 24 8 11 -14 -20 -25 53 19 31 12 - 4 -11 - a 46 33 BANFF, ALBERTA - 1947 - 1948 November December January February March Day Max. Min. Max. Min. Max. Min. Max. Min. Max. Min. 1 2 $ M i ? i f 18 -17 19 3 40 22 25 17 33 19 24 1 25 -21 4 34 20 19 6 32 15 8 1 35 - 1 5 25 - 1 29 7 28 12 2 -22 45 6 6 28 6 28 13 42 21 14 -24 46 12 7 38 23 26 16 28 8 20 2 25 10 8 35 26 22 4 32 10 8 - 2 15 - 7 9 34 24 24 7 25 4 0 -14 18 -23 10 33 20 24 6 29 13 14 -29 28 -18 11 28 10 30 18 36 5 12 34 13 30 13 26 - 7 42 15 13 32 20 31 25 30 8 43 21 14 25 18 29 24 35 18 36 25 15 21 11 22 3 19 5 37 16 16 21 - 6 31 4 29 - 3 30 9 17 27 1 35 25 29 14 4 -12 30 - 8 18 27 19 38 31 30 - 2 0 -12 30 20 19 21 9 35 20 39 17 16 -27 32 7 20 18 12 37 16 24 -12 32 - 5 21 18 - 9 47 32 34 8 42 28 22 28 2 40 25 29 19 35 27 23 34 14 38 25 25 0 29 - 1 24 37 24 50 25 25 17 31 3 34 5 25 41 24 54 23 16 - 4 37 26 36 - 8 26 42 27 42 25 18 -21 37 22 34 7 27 35 12 44 34 27 10 34 16 32 - 7 28 30 11 40 34 28 13 25 - 5 48 28 29 36 8 38 17 37 10 27 -19 45 35 30 32 12 19 - 1 29 18 38 12 31 18 1 29 1 40 30 BANFF, ALBERTA - 1948 - 1949 November December January February March Day Max. Min. Max. Min. Max. Min. Max. Min. Max. Min. 1 40 19 32 23 %X 16 _ 9 50 15 2 38 0 33 22 10 - 6 15 - l o 47 13 3 40 29 18 11 10 -31 14 - 6 46 19 4 37 23 20 - 9 19 - 3 4 - 6 48 27 5 36 27 20 - 6 40 4 14 -24 50 32 6 33 14 16 - 3 43 14 19 - 4 44 25 7 30 9 17 1 19 14 2 - 6 40 24 8 35 16 3 -15 - 2 -17 20 -16 25 10 9 39 15 10 -27 2 -35 22 3 33 -15 10 u 20 19 1 10 -31 23 12 39 - 6 11 42 25 24 1 21 - 2 1 -11 43 1 12 35 30 8 0 21 3 14 -31 19 15 13 37 29 - 6 -17 24 8 16 -11 15 - 4 14 43 23 5 -28 28 12 18 2 27 -19 15 36 31 12 -24 27 8 20 - 7 19 0 16 34 20 11 -17 21 - 5 32 11 32 3 17 35 23 11 -12 25 14 4 - 1 40 7 18 30 19 18 - 3 11 - 7 -11 -18 45 29 19 28 22 27 14 - 3 -25 6 -39 46 29 20 31 21 26 18 1 -29 16 -31 41 25 21 32 10 22 6 9 -12 34 2 40 30 22 35 26 8 -12 - 6 -22 40 22 40 24 23 35 26 12 - 9 - 1 -44 43 8 45 20 24 36 26 19 -19 8 -37 40 17 45 18 25 28 14 14 1 10 -12 42 26 41 27 26 23 2 18 0 17 5 46 12 37 23 27 21 2 18 7 14 3 46 14 37 26 28 32 12 18 4 19 -21 50 18 40 17 29 27 18 22 8 22 9 39 20 30 29 17 25 8 24 4 a 25 31 24 18 9 2 49 20 BANFF, ALBERTA - 1949 - 1950 November December January February March Day Max. Min. Max. Min. Max. Min. Max. Min. Max. Min, 1 58 24 42 28 -21 3 -18 2 2 56 27 35 26 -19 "11 10 - 7 39 16 3 57 29 28 9 - 8 -48 13 - 9 43 31 4 58 32 19 8 - 1 -21 23 - 1 38 33 5 60 32 32 10 3 -11 39 19 30 16 6 54 31 33 17 5 - 6 37 16 27 22 7 53 28 22 1 4 - 8 30 2 27 12 8 45 24 20 - 2 2 -26 30 5 17 10 9 41 28 26 14 - 4 -23 25 34 10 - 1 10 41 23 8 - 2 -10 -25 32 12 11 - 8 11 36 29 14 -10 -11 -17 32 9 24 -29 12 37 31 18 7 -18 -21 32 0 23 -20 13 u 25 18 - 1 -20 -28 39 22 29 4 14 42 26 23 10 -13 -30 40 21 34 11 15 43 34 26 16 -13 -36 49 29 35 17 16 39 23 14 5 -18 -50 45 29 21 9 17 54 34 - 1 - 4 - 8 -38 35 4 10 - 1 18 48 37 - 7 -12 - 1 -20 44 25 40 3 19 42 28 1 -17 7 - U 37 26 38 26 20 39 11 14 -11 8 2 33 7 37 19 21 42 22 18 3 30 3 29 18 38 24 22 42 29 23 10 - 3 -12 30 18 38 21 23 42 32 21 - 1 -20 -22 31 15 37 19 24 43 33 17 - 2 -19 - u 37 17 38 14 25 43 33 21 - 6 -12 -60 39 17 37 23 26 a 32 20 8 - 6 -31 39 29 32 21 27 46 33 - 3 -13 4 -43 33 18 38 15 28 40 31 - 2 -13 1 -44 29 - 1 36 22 29 35 30 39 -14 1 -25 34 16 30 35 18 - 8 -17 - 1 -36 34 20 31 - 7 -18 2 -13 38 7 BANFF, ALBERTA - 1950 - 1951 November December January February March Day Max. Min. Max. Min. Max. Min. Max. Min. Max. Min. 1 32 10 5 - 6 29 5 12 - 5 22 -11 2 32 6 16 -15 16 - 3 27 2 19 - 2 3 51 26 11 4 6 0 32 14 22 -12 4 54 41 6 -20 7 -29 27 3 3 - 6 5 37 31 11 -13 16 -29 9 - 7 -10 -23 6 37 28 22 4 17 1 10 -73 -12 -40 7 32 11 27 6 19 6 22 -16 - 4 -34 8 22 8 29 19 26 12 37 1 - 5 - a 9 25 - 3 29 21 7 40 29 - 3 -25 10 32 15 38 25 20 5 20 0 6 -20 11 31 18 37 28 27 12 11 -21 24 -29 12 34 8 35 30 28 5 21 -26 35 11 13 31 10 34 24 27 16 23 - 8 40 24 14 26 11 26 - 3 25 17 30 6 41 23 15 15 1 29 14 27 6 33 10 40 30 16 18 12 35 23 20 9 32 7 30 12 17 4 - 2 33 21 28 15 32 11 30 -15 18 13 -12 24 12 15 -18 35 6 36 - 5 19 21 0 30 7 8 -24 32 11 46 22 20 28 10 40 23 24 - 7 32 12 50 19 21 33 10 44 35 23 11 25 12 43 32 22 1 -rlO 41 33 28 10 32 -12 37 21 23 22 -28 43 32 29 13 34 - 5 35 25 24 33 14 36 33 17 14 32 14 42 30 25 32 18 35 24 0 - 6 32 18 45 31 26 42 28 23 6 -13 -17 24 11 a 30 27 45 31 26 14 - 4 - a 24 - 1 35 19 28 34 29 32 22 3 -30 26 - 9 42 14 29 23 4 32 12 7 -21 49 25 30 15 5 33 27 4 -38 46 32 31 25 9 5 -32 46 30 BANFF, ALBERTA - 1951 - 1952 November December January February March Day Max. Min. Max. Min. Max. Min. Max. Min. Max. Min. 1 23 -13 36 30 0 -16 37 25 13 6 2 40 16 30 6 - 7 35 22 18 -10 3 30 16 31 16 10 - 4 36 20 22 -12 4 41 25 29 21 17 4 33 25 36 2 5 41 22 24 6 20 3 34 22 35 0 6 39 16 22 6 26 13 36 26 34 7 7 42 22 16 - 6 21 5 "28 23 36 2 8 40 28 19 3 21 - 1 34 15 46 22 9 38 28 20 7 24 9 37 26 33 19 10 32 17 31 16 25 10 39 25 19 12 11 40 24 u 22 6 -12 40 19 26 1 12 35 25 35 23 15 -19 35 16 28 -13 13 37 20 18 4 26 6 29 15 34 2 14 26 19 - 4 -16 - 1 -12 28 1 35 6 15 28 14 8 -21 - 9 -20 30 11 40 2 16 20 - 3 14 - 3 20 -29 24 5 38 6 17 25 9 6 - 4 25 16 13 8 39 8 18 33 12 - 6 -12 27 19 13 - 6 36 20 19 35 17 -10 -16 25 - 3 22 -15 32 - 7 20 28 20 3 -26 3 -12 20 -11 32 6 21 19 7 9 - 2 -15 -23 22 -11 33 3 22 30 14 12 - 2 -12 -32 22 - 8 40 1 23 33 19 13 - 6 - 7 -34 24 -13 44 21 24 30 11 10 - 6 15 -15 30 0 34 27 25 38 16 8 - 6 30 4 28 11 46 20 26 36 28 0 -20 35 0 37 22 48 25 27 38 28 16 -12 38 31 30 1 50 31 28 34 29 24 6 46 32 26 - 2 47 30 29 32 25 10 - 5 39 28 28 4 38 24 30 35 15 -14 -21 36 32 39 26 31 - 2 -39 36 31 35 17 BANFF, ALBERTA - 1952 - 1953 November December January February March Day Max. Min. Max. Min. Max. Min. Max. Min. Max. Min. 1 38 25 25 12 28 18 35 30 22 0 2 39 15 28 10 23 10 38 28 28 4 3 49 32 32 12 36 21 35 29 35 9 4 47 41 32 23 36 28 35 28 38 28 5 32 17 33 27 14 2 32 38 30 13 6 38 12 32 20 6 -16 32 9 40 8 7 41 24 27 13 17 -17 38 26 44 32 8 38 18 30 12 32 -15 28 2 46 26 9 38 22 28 35 36 - 4 26 - 1 48 26 10 40 29 28 20 29 7 29 18 36 22 11 44 33 24 7 26 2 29 22 32 6 12 38 28 30 11 1 -11 34 12 37 17 13 38 19 39 24 -14 -18 32 26 35 9 34 35 26 38 31 - 5 -25 29 9 33 4 15 28 19 25 21 28 -20 32 23 38 11 16 34 9 26 7 31 22 29 5 41 21 17 36 20 25 12 22 5 31 21 33 25 18 30 17 20 11 25 4 27 - 3 33 6 19 40 23 17 - 3 33 14 24 - 5 36 16 20 34 21 15 - 1 35 28 28 - 4 32 12 21 27 5 24 4 35 7 35 34 35 34 22 30 10 22 8 33 28 32 23 35 6 23 25 9 15 0 30 19 32 0 42 7 24 19 9 20 2 37 27 35 7 54 25 25 U - 3 16 3 31 21 42 34 44 32 26 19 - 3 15 2 12 4 40 24 42 22 27 22 - 3 20 - 1 21 - 1 30 16 46 20 28 18 7 26 12 34 34 22 14 48 26 29 16 4 30 21 37 28 u 17 30 17 5 35 22 35 15 45 26 31 33 28 39 4 34 24 BANFF, ALBERTA - 1953 - 1954 November December January February March Day Max. Min. Max. Min. Max. Min. Max. Min. Max. Min. \ 11 n a 22 28 K 32 48 33 30 11 -zi 3 39 19 29 18 30 23 49 36 36 3 4 44 20 27 16 30 23 50 36 42 - 2 5 50 29 29 19 33 20 48 30 u 9 6 36 32 29 23 32 19 45 27 29 24 7 40 25 24 2 17 9 48 29 35 8 8 49 34 25 12 17 12 50 33 43 27 9 46 36 35 21 17 - 4 35 24 51 32 10 47 29 29 13 17 - 1 18 12 34 25 11 50 31 29 20 21 8 6 - 3 33 8 12 43 30 32 23 17 - 5 6 - 5 37 2 13 43 26 28 18 18 9 38 —1 39 5 14 48 26 34 24 5 -15 25 - 6 34 22 15 43 33 21 3 -22 -26 31 9 30 - 3 16 36 30 23 0 1 -39 38 27 41 10 17 29 23 28 12 4 -13 45 32 35 19 IB 25 6 40 15 6 -19 32 22 41 14 19 26 10 32 28 - 9 -18 30 20 41 22 20 31 13 25 22 -11 -33 36 26 35 19 21 32 16 24 8 -18 -26 37 24 41 12 22 38 24 22 3 -16 -23 40 31 35 9 23 34 26 28 11 -15 -25 43 32 40 10 24 37 26 33 23 -12 -21 35 28 31 19 25 33 26 35 29 - 1 -20 34 27 35 3 26 35 24 32 28 1 -14 38 • 14 15 10 27 30 13 35 9 17 -24 31 16 9 0 28 35 27 31 27 27 10 31 25 20 -22 29 35 27 31 19 18 - 3 27 9 30 42 28 35 20 38 9 31 - 2 31 32 26 40 32 25 11 APPENDIX 2 DAILY MAXIMUM AND MINIMUM TEMPERATURES FOR WINTER MONTHS, LAKE LOUISE, ALBERTA. 1932 to 1954 LAKE LOUISE, ALBERTA - 1932 - 1933 November December January February March Day Max. Min. Max. Min. Max. Min. Max. Min. Max. Min. 1 30 4 31 22 20 4 14 -20 23 0 2 35 25 3* 26 20 - 1 17 -13 37 - 6 3 30 16 20 19 9 28 -10 33 19 4 31 20 30 26 24 13 27 - 1 30 20 5 35 23 17 - 1 27 16 24 12 34 20 6 33 21 - 1 - 3 26 21 7 -31 36 21 7 29 - 6 - 3 -44 29 20 4 -12 31 22 8 31 14 - 6 -36 29 12 - 6 -25 31 - 1 9 30 11 15 -34 30 20 - 3 -47 23 -26 10 24 -12 9 -22 25 11 2 -18 22 -12 11 25 -11 2 -35 24 9 7 -17 31 - 8 12 26 17 3 - 8 29 22 4 -27 37 24 13 5 - 2 3 -24 33 11 4 -31 34 24 14 12 -17 9 -27 30 25 10 -24 33 -15 15 16 2 7 -28 4 2 IS - 3 35 -17 16 28 2 11 -28 4 -27 18 -14 37 - 3 17 34 22 10 -22 1 -23 23 15 34 - 1 18 40 32 11 -22 3 -29 23 -10 36 7 19 38 30 17 4 8 -16 25 8 37 - 3 20 28 10 24 12 14 -13 33 18 39 26 21 26 9 25 11 16 3 31 16 32 18 22 29 19 24 2 18 - 7 30 16 30 6 23 31 12 26 16 24 2 28 14 32 9 24 32 26 20 3 20 - 5 27 7 29 2 25 25 -12 24 7 18 - 4 29 18 34 — 2 26 36 17 24 8 18 - 8 29 26 38 -10 27 40 25 21 10 18 8 22 1 39 2 28 40 31 24 13 18 - 7 18 4 39 26 29 35 30 21 8 16 -18 39 16 30 28 26 19 - 8 16 -18 39 28 31 24 5 18 - 5 35 25 LAKE LOUISE, ALBERTA - 1933 - 1934 November December January February March Day Max. Min. Max. Min. Max. Min. Max. Min. Max. Min. 1 fo 32 32 20 22 -10 32 22 27 18 i 16 32 18 22 2 40 32 37 21 3 32 - 3 32 10 32 22 32 -10 39 32 4 42 - 5 22 6 31 20 30 -18 26 - 8 5 41 - 1 30 10 30 6 31 -18 23 - 8 6 30 32 20 0 29 -10 32 - 2 27 6 7 30 2 22 6 19 -18 38 5 28 12 8 60 27 20 6 21 -32 36 27 33 5 9 58 24 24 5 20 -12 36 18 24 -10 10 38 27 19 -13 31 20 36 50 6 11 41 32 10 0 32 8 38 10 48 8 32 42 34 0 - 8 21 -34 38 - - 2 - 46 23 13 34 47 48 24 22 7 0 - 8 26 26 32 12 37 28 -_2~ -_2- 11 al ft 42 20 2 -1? 22 -22 37 - 5 43 30 16 37 38 - 3 -24 24 22 35 - 8 53 32 17 40 20 6 0 29 10 38 1 37 34 18 30 34 20 2 21 -16 37 - 9 u 24 19 37 25 34 -10 24 -11 36 - 8 49 29 20 36 28 32 - 5 34 18 27 -18 38 24 21 38 28 28 - 6 32 18 24 -18 32 22 22 37 26 31 -10 28 16 25 -19 35 5 23 38 22 39 -13 27 32 15 -21 40 10 24 40 28 30 - 5 34 -24 9 -21 36 8 25 50 23 - 5 -49 24 -21 8 -40 35 6 26 27 13 - 5 -46 26 20 11 -34 40 4 27 30 5 1 -36 31 24 24 5 32 22 28 30 - 1 19 - 6 a 10 23 8 46 2 29 19 10 32 -14 41 7 40 4 30 32 24 - 8 - 2 30 3 44 21 31 34 - 6 32 22 42 22 LAKE LOUISE, ALBERTA - 1934 - 1935 November December January February March Day Max. Min. Max. Min. Max. Min. Max. Min. Max. Min. 1 41 32 21 8 30 15 40 9 36 18 2 40 19 21 0 30 -31 40 31 37 20 3 38 28 23 0 22 -17 36 14 33 16 4 35 22 21 1 21 19 a 22 21 - 1 5 40 26 27 6 20 12 40 - 4 20 -15 6 40 21 22 0 20 12 33 - 4 27 -23 7 35 28 28 0 23 -32 22 -19 31 -17 8 38 14 28 0 23 23 32 -15 35 -10 9 35 27 0 22 - 6 32 -31 27 - 7 10 34 25 6 20 3 32 -32 22 -16 11 32 29 14 19 5 30 - 5 34 19 12 39 10 54 20 32 -19 28 - 4 35 29 13 47 12 34 28 12 -20 29 - 5 a 36 14 39 27 34 12 -12 -42 28 - 5 39 31 15 42 30 34 8 1 -30 31 5 38 23 16 38 31 32 6 2 -15 34 32 30 4 17 40 22 34 34 -31 -34 34 24 34 15 18 30 18 34 34 -15 -40 30 4 33 3 19 30 33 31 18 -17 -54 27 0 34 4 20 20 12 32 18 -31 -55 35 12 25 1 21 19 10 27 12 - 5 -18 34 12 26 -12 22 31 13 25 8 15 -20 35 34 30 -10 23 30 34 22 0 19 15 39 34 32 3 24 38 19 25 1 24 22 27 -18 34 21 25 38 20 25 0 35 32 28 -32 25 0 26 21 10 34 -19 35 15 30 -18 26 5 27 20 8 12 -21 36 32 30 -32 26 -13 28 22 - 5 12 -24 38 18 34 10 30 2 29 20 31 8 - 2 35 6 28 - 9 30 28 8 15 - 6 36 8 24 -26 31 16 - 3 41 10 25 -26 LAKE LOUISE, ALBERTA - 1935 - 1936 November December January February March Day Max. Min. Max. Min. Max. Min. Max. Min. Max. Min. 1 i i -30 32 " 4 H -37 34 2 -30 33 - 6 16 -38 44 32 3 29 -15 32 0 17 -35 44 32 4 26 10 28 - 9 18 -36 38 25 5 25 15 21 -10 12 -25 42 14 6 33 19 26 13 0 -26 35 22 7 39 26 30 - 6 - 4 -55 36 23 8 35 17 35 25 -14 -40 36 24 9 15 -13 34 9 -12 -36 29 2 10 17 -19 30 20 - 7 -22 30 - 1 11 22 12 26 9 -12 -22 21 -12 02 27 20 29 16 - 3 -14 32 21 13 24 -13 28 - 4 - 8 -42 31 14 14 22 - 3 22 - 5 32 15 - 8 -52 34 14 15 26 0 18 - 7 29 - 3 - 7 -29 35 16 16 30 10 20 - 5 26 - 3 - 4 -52 34 21 17 30 11 19 - 7 22 - 7 - 2 -50 30 17 18 21 -16 20 -11 13 -32 10 -45 32 27 19 31 20 17 -12 19 -20 18 -32 40 22 20 34 18 21 - 7 23 - 3 14 -10 35 25 21 32 12 26 - 2 27 14 18 -11 48 22 22 21 7 27 -10 35 25 20 - 7 31 - 2 23 30 3 29 10 35 26 10 0 20 9 24 31 10 35 3 10 -25 27 -10 25 30 15 24 -23 40 -41 25 14 26 32 23 20 -23 18 - 2 27 0 27 36 27 13 -10 32 10 17 1 28 34 29 14 -32 35 27 16 -26 29 35 30 16 -31 37 25 9 -10 30 34 0 18 - 5 14 -19 31 18 -31 11 -33 LAKE LOUISE, ALBERTA - 1936 - 1937 November December January February March Day Max. Min. Max. Min. Max. Min. Max. Min. Max. Min. 1 4° -21 ,27 10 - 2 -37 4 -10 37 2 i 39 -17 25 10 1 -27 0 -11 39 9 3 29 14 22 - 8 10 -23 34 -30 38 22 4 30 34 15 -10 9 -22 34 -10 43 15 5 20 8 14 -10 32 -20 14 - 8 47 18 6 18 -22 14 -28 6 -47 17 -13 35 26 7 21 -17 18 -10 2 -39' - 8 -17 40 - 3 8 26 8 21 0 1 -26 0 -47 40 - 4 9 32 13 22 - 7 1 -20 -12 -20 43 4 10 31 13 22 - 3 4 -23 19 3 42 24 11 33 22 23 - 3 7 -21 25 16 46 6 12 39 26 26 8 16 -13 20 12 38 18 13 44 30 30 9 7 -21 23 -20 40 - 6 42 19 26 21 34 -37 23 - 1 26 - 6 5 42 20 26 -21 7 -34 26 - 3 36 16 16 40 13 21 -23 0 -27 28 14 u 0 17 40 24 25 4 0 -39 24 12 40 - 4 18 43 26 37 20 7 -15 17 1 43 4 19 54 33 38 «. 5 5 -27 17 0 a 10 20 42 16 37 - 8 - 3 -45 34 -26 33 0 21 44 7 38 0 - 5 -47 18 -34 41 -16 22 46 6 37 24 - 3 -29 30 -11 40 0 23 34 7 28 10 5 -20 30 - 1 a 1 24 34 7 22 -10 7 -17 29 15 .40 20 25 32 6 22 -17 8 -12 34 - 7 44 12 26 26 3 22 -38 5 -13 31 -17 45 4C 27 30 0 17 -30 - 2 -34 33 -34 45 0 28 31 6 6 -28 - 2 -43 33 -26 40 10 29 34 8 0 -35 6 -16 a 14 30 30 6 7 -16 4 -34 43 23 31 - 9 -21 4 -10 44 24 LAKE LOUISE, .ALBERTA - 1937 - 1938 November December January February March Day Max. Min. Max. Min. Max. Min. Max. Min. Max. Min. i i i 12 4 & zil 18 B if :9 0 25 3 46 26 15 4 16 -14 26 - 3 35 22 4 38 11 21 0 22 -15 28 15 44 11 5 39 11 25 16 17 - 9 28 13 35 25 6 35 21 30 10 13 -13 26 - 4 36 4 7 37 19 20 -20 14 -13 15 - 8 35 - 7 8 41 25 13 -33 22 - 9 22 -21 38 0 9 44 25 0 -38 22 2 19 -29 36 7 10 39 24 15 -26 26 17 19 - 7 36 24 11 34 24 30 11 18 -21 15 -20 36 23 12 31 14 20 - 8 18 - 5 16 - 9 46 25 13 20 3 25 - 9 25 11 - 2 -13 45 24 14 7 2 27 9 32 21 - 1 -19 42 25 15 22 8 32 9 31 21 9 -34 35 20 16 17 3 30 20 23 -15 22 -28 35 25 17 16 3 38 16 21 —7 23 -25 32 17 18 18 - 2 36 13 19 -13 24 -10 30 18 19 11 -24 32 - 4 19 2 32 -14 29 16 20 25 8 32 - 2 18 -19 22 - 6 31 - 4 21 16 10 24 12 27 12 34 0 30 -11 22 34 6 13 -10 27 6 36 - 8 29 1 23 35 22 0 -28 23 - 4 37 - 8 36 10 24 33 22 0 -10 19 - 3 39 - 6 38 16 25 32 21 0 -22 25 7 44 0 46 23 26 25 17 6 -18 19 3 47 0 u 16 27 28 0 13 - 5 31 - 7 49 0 36 29 28 28 3 32 0 27 0 53 0 34 26 29 28 4 38 32 3 -40 26 14 30 17 33 32 1 -42 32 14 31 30 22 6 -27 31 -10 LAKE LOUISE, ALBERTA - 1938 - 1939 November December January February March Day Max. Min. Max. Min. Max. Min. Max. Min. Max. Min. 1 38 17 34 22 33 26 14 -30 26 - 3 2 34 14 29 15 33 23 17 -15 29 8 3 35 27 28 14 30 24 17 -10 20 - 6 4 31 18 26 7 27 10 10 -17 24 -30 5 27 5 35 24 15 - 1 15 - 3 25 -27 6 27 - 3 32 22 18 -17 15 -12 31 - 5 7 33 17 33 26 28 12 5 -24 30 12 8 27 12 33 23 27 9 - 4 -38 31 -17 9 26 2 33 17 26 9 -10 -47 30 -20 10 23 8 21 - 4 27 - 1 - 1 -45 31 -14 11 21 -10 14 -17 23 19 10 -13 31 14 12 24 14 13 -23 23 19 26 9 28 9 13 25 13 18 - 6 31 18 25 10 25 3 U 30 20 19 - 4 27 - 8 29 16 30 -22 15 32 25 13 -17 25 11 28 10 30 5 16 30 25 16 -16 24 - 4 27 - 2 33 - 2 17 29 22 14 -15 25 8 33 23 40 - 7 18 34 23 13 -20 26 12 18 - 6 50 23 19 31 21 12 -21 30 16 26 -24 54 27 20 25 4 9 -25 25 13 25 -26 53 30 21 19 0 11 -18 16 -21 26 -24 52 35 22 17 -20 22 -14 18 - 1 32 -19 57 20 23 16 -21 33 16 25 5 35 6 60 24 24 21 -19 33 17 25 15 34 - 2 50 35 25 32 - 5 29 - 3 37 5 29 16 40 2t 26 21 - 7 29 -35 29 7 26 -13 39 5 27 50 - 5 19 -15 26 5 26 - 8 43 - 4 28 26 0 11 -22 26 11 24 -13 49 8 29 27 10 4 -27 26 7 44 8 30 32 19 26 4 21 - 1 40 31 31 27 19 17 -17 44 35 LAKE LOUISE, ALBERTA - 1939 - 1940 November December January February March Day Max, Min. Max. Min. Max. Min. Max. Min. 1 36 24 29 14 24 -20 9 32 30 28 16 23 - 7 -20 4 37 28 26 12 31 14 5 36 22 30 14 34 20 6 39 28 14 -16 30 25 7 37 20 12 -17 37 22 8 NO 39 24 10 -27 34 - 8 9 36 11 12 -28 38 27 10 32 26 18 - 5 31 24 11 34 24 16 -10 28 3 12 RECORD 30 18 14 -24 25 -16 13 28 2 19 - 5 30 -10 14 30 24 14 5 34 5 15 32 20 27 - 8 28 -18 16 FOR 33 22 30 10 28 - 8 17 26 16 15 -16 32 20 18 21 -10 10 -31 30 2 19 28 18 10 -35 30 5 20 NOVEMBER 30 22 8 -36 32 0 21 30 26 13 -29 22 10 22 24 10 16 -20 32 16 23 22 -29 12 -37 30 18 24 20 -10 5 -41 26 8 25 15 -28 10 -42 22 5 26 8 -26 16 -22 16 - 8 27 15 -10 29 8 37 - 7 28 32 10 37 15 38 16 29 25 22 41 18 38 19 30 27 15 30 - 8 31 18 -18 18 -14 Max. Min. NO RECORD FOR MARCH LAKE LOUISE, ALBERTA - 19Z>0 - 1941 November December January February March Day Max. Min. Max. Min. Max. Min. Max. Min. Max. Min. 1 36 23 2$ 8 - 8 32 23 35 31 2 34 15 31 8 - 4 38 27 42 7 3 33 19 36 25 34 -17 41 7 41 27 4 27 19 33 6 13 -18 39 4 45 24 5 26 - 2 32 25 32 - 9 41 - 6 42 4 6 27 31 32 19 15 - 6 49 - 7 42 16 7 22 7 29 23 21 -26 47 -10 47 25 8 18 - 8 33 26 30 - 7 41 - 6 41 24 9 13 - 4 29 10 32 15 39 - 7 35 - 2 10 9 -22 22 - 9 37 24 35 34 44 -10 11 8 -29 18 -10 36 22 37 32 50 -10 12 9 -25 15 -28 41 11 33 6 40 -16 13 15 -15 22 -10 36 20 37 7 48 -16 34 23 - 3 18 -20 31 5 33 -18 43 4 15 28 12 15 -25 28 8 35 -14 40 0 16 31 - 1 20 -22 34 15 41 -14 45 - 1 17 31 10 22 -18 22 - 1 43 -12 49 22 18 29 6 27 - 5 31 20 38 1 45 27 19 32 - 3 30 5 28 11 26 15 44 17 20 28 - 7 32 22 36 4 31 5 40 9 21 29 - 5 32 19 30 -19 17 - 2 42 - 2 22 22 -19 36 21 20 7 15 - 2 43 4 23 21 - 3 31 - 2 21 0 16 -38 47 16 24 29 12 28 9 16 -18 21 -20 49 31 25 30 17 32 10 26 - 8 32 - 2 50 16 26 30 31 34 8 33 4 31 32 52 28 27 27 32 36 10 33 12 38 22 53 31 28 29 38 32 5 34 25 44 31 52 9 29 30 20 26 5 36 - 1 55 13 30 29 12 25 6 37 2 50 30 31 19 1 37 13 47 33 LAKE LOUISE, ALBERTA - 19a - 194-2 November December January February March Day Max. Min. Max. Min. Max. Min. Max. Min. Max. Min. 1 33 18 35 22 0 -35 28 9 36 24 2 29 5 39 29 4 -12 34 4 34 21 3 29 19 36 20 7 -31 35 1 31 5 4 39 21 37 11 • -38 32 18 36 14 5 a 29 34 - 1 0 - 4 « 30 17 32 21 6 40 18 29 21 7 -37 54 3 29 5 7 43 8 30 14 2 -40 33 3 35 7 8 44 17 30 14 10 -27 32 2 40 15 9 47 21 22 12 10 -25 31 -16 36 26 10 43 13 25 - 5 19 3 32 3 11 37 21 21 - 2 22 3 33 12 12 39 17 14 -16 30 14 34 - 5 13 29 8 12 -19 36 14 32 -11 14 36 29 21 -14 29 - 2 38 - 2 15 31 16 26 0 24 - 7 29 15 16 32 1 32 10 27 - 7 20 -30 17 30 -10 26 5 26 - 4 19 -36 18 23 -15 28 2 28 8 20 -24 19 30 - 4 32 22 29 - 3 27 -15 20 25 - 1 29 0 22 - 8 35 t 21 21 18 30 21 33 -11 20 -20 22 19 -17 22 1 32 -11 12 -25 23 18 -17 25 12 35 - 9 17 -24 24 28 - 2 17 - 3 37 4 20 - 6 25 33 24 24 -10 35 11 23 -17 26 a 28 11 -29 37 13 28 -19 27 36 19 11 -32 34 22 31 2 28 39 7 13 -30 35 15 30 - 1 29 41 27 11 -30 27 -10 30 a 26 5 -27 29 4 31 31 -46 27 -12 LAKE LOUISE, ALBERTA - 1942 - 1943 November December January February March Day Max. Min. Max. Min. Max. Min. Max. Min. Max. Min. 1 30 -1 21 -20 117 -22 15 - 9 33 -18 23 19 - 5 15 -30 26 12 36 -20 3 34 18 17 -14 17 -29 25 3 32 -14 4 24 4 18 -19 10 -31 26 13 26 -15 5 18 0 13 -20 22 -12 27 9 26 -28 6 27 9 18 -19 21 1 33 18 28 -30 7 31 -11 15 -22 23 4 24 - 8 26 - 6 8 28 -10 14 -13 29 17 15 -36 29 4 9 23 -15 18 - 4 26 4 14 -30 28 -21 10 29 16 26 4 21 12 21 -10 27 2 11 35 14 28 14 24 -15 30 15 26 6 12 39 9 34 23 21 - 4 35 7 35 11 13 40 3 36 26 33 17 39 8 14 46 29 34 21 34 27 40 14 15 33 28 35 20 31 - 2 45 12 16 30 17 36 27 22 -32 43 9 17 19 - 1 35 22 10 -*2 46 8 18 27 -14 27 - 3 2 -48 44 4 19 24 -10 33 12 - 4 -24 38 22 20 26 -15 29 10 -14 -36 40 0 21 25 - 9 30 22 -19 -49 35 21 22 31 10 28 19 - 4 -38 36 -15 23 35 31 22 -16 0 -46 31 - 5 24 33 10 17 -19 4 -45 36 -20 25 23 9 23 11 9 -38 48 - 3 26 19 1 12 -28 13 - 7 46 - 5 27 27 -16 13 -21 20 -12 50 - 6 28 17 -25 25 5 26 0 59 13 29 18 -16 16 - 7 28 -20 30 24 9 27 0 25 -29 31 21 3 16 -15 LAKE LOUISE, ALBERTA - 1943 - 1944 November December January February March Day Max, Min. Max. Min. Max. Min. Max. Min. Max. Min. i % -io 30 29 12 S 1 30 34 " I 31 29 ~ 2 i 3 35 24 37 25 18 - 5 35 - 4 30 - 2 4 34 19 28 4 13 - 3 27 4 21 - 1 5 34 24 27 -14 16 -18 28 8 25 -10 6 35 - 1 22 - 6 18 -26 30 20 31 -11 7 34 1 25 12 19 9 28 21 39 -16 8 34 14 24 -10 28 5 34 -15 38 23 9 42 5 28 - 6 25 -17 22 -15 48 25 10 32 19 21 - 9 20 -21 30 -25 36 16 11 43 16 20 - 2 13 -23 30 3 29 - 7 12 42 3 27 -11 23 2 25 -17 20 8 13 45 11 25 -10 29 17 24 2 24 -27 14 40 17 20 -11 32 13 27 4 38 -11 15 42 6 25 - 8 32 18 22 -19 37 2 16 43 4 23 - 8 33 20 25 6 35 27 17 34 16 28 - 4 35 28 26 -17 37 26 18 36 3 29 - 8 32 24 30 - 6 39 4 19 38 16 23 -20 45 38 27 -22 38 19 20 35 17 23 -18 43 20 23 - 7 35 11 21 39 25 26 - 8 35 - 9 20 5 31 -10 22 48 20 23 -13 38 2 28 -29 31 21 23 49 7 22 10 31 6 23 -10 29 15 24 45 22 27 17 30 14 35 11 22 - 9 25 36 - 2 29 18 25 - 7 24 -15 22 -10 26 33 - 7 31 2 28 -22 30 - 9 18 -12 27 36 - 1 24 -11 26 -22 32 -14 20 -30 28 23 - 3 22 -10 23 -25 25 - 4 40 18 29 29 19 14 -15 28 -24 29 -12 48 32 30 33 23 13 -16 34 -22 42 23 31 19 -10 35 -18 43 20 LAKE LOUISE, ALBERTA - 1944 - 1945 November December January February March Day Max. Min. Max. Min. Max. Min. Max. Min. Max. Min. i » S l\ E 26. 23 zl a i S 11 -h 3 29 20 28 - 7 25 2 21 5 26 6 4 38 21 28 19 27 9 36 8 18 -39 5 39 29 36 21 30 18 30 16 17 -35 6 38 28 37 27 29 17 32 - 3 27 - 6 7 39 27 29 6 27 7 30 18 29 14 8 40 24 30 8 32 8 39 26 31 3 9 36 28 23 -12 36 25 32 12 39 10 10 36 21 26 -12 30 4 29 8 t l 27 17 28 -10 34 22 29 - 2 12 35 7 30 -10 32 17 28 14 13 33 -12 27 - 8 29 5 25 7 14 33 -10 25 - 3 35 13 19 -18 15 37 3 29 - 8 25 - 7 13 -10 16 37 - 3 19 - 8 26 7 17 -43 17 32 2 18 - 6 25 13 24 -35 18 27 2 15 - 7 24 10 30 -32 19 36 0 21 - 2 29 -17 27 -20 20 28 1 19 -10 18 -30 28 0 21 30 13 18 -15 28 -11 24 - 9 22 34 23 14 -28 28 -10 26 - 8 23 35 26 8 -30 32 - 9 34 -10 24 33 13 7 -30 33 -16 37 -15 25 26 - 3 2 -33 30 - 8 46 8 26 28 13 12 -22 29 -20 43 3 27 29 13 10 -18 25 -25 34 18 28 24 -10 24 8 21 -20 28 -10 29 28 -10 24 0 25 -26 30 30 11 18 - 5 22 -24 31 15 - 7 27 -25 LAKE LOUISE, ALBERTA - 1945 - 194-6 November December January February March Day Max. Min. Max. Min. Max. Min. Max. Min. Max. Min. 1 32 29 -10 2§ 12 20 - 1 33 21 i 27 32 -10 29 7 21 - 2 41 20 3 42 23 31 -11 28 16 26 - 7 35 1 4 41 29 31 22 30 12 23 -24 33 - 1 5 31 5 33 12 29 9 22 -30 31 16 6 22 - 6 26 14 30 - 9 21 7 33 14 7 14 - 8 23 10 24 11 22 -20 31 8 a 13 -35 19 -22 27 6 26 6 33 18 9 14 -16 12 -24 24 9 24 10 36 - 8 10 25 - 8 21 - 3 27 10 20 - 5 47 5 n 24 8 23 - 8 24 -14 19 -21 43 23 12 28 7 15 -27 20 - 2 21 - 7 13 27 1 10 -25 26 16 27 16 14 26 6 9 -15 31 2 34 20 15 37 3 12 -12 32 6 34 21 16 29 - 3 19 -14 26 - 8 37 -15 17 27 16 12 -20 29 19 31 - 5 18 28 11 11 -31 31 16 33 10 19 29 8 10 -18 18 18 39 -17 20 27 1 16 7 21 -29 33 - 6 21 28 -20 13 - 3 22 -12 40 6 22 24 - 4 15 - 8 30 16 35 18 23 34 2 27 11 25 10 36 20 24 30 5 31 10 22 14 38 25 25 32 8 30 14 20 - 7 35 22 26 28 12 31 11 14 -30 34 - 4 27 28 10 32 13 19 1 33 17 28 23 -10 29 11 21 7 31 23 29 36 4 32 16 21 5 30 26 10 34 17 18 -26 31 28 10 19 12 LAKE LOUISE, ALBERTA _ 1946 - 1947 November December January February March Day Max. Min. Max. Min. Max. Min. Max. Min. Max. Min. 1 29 16 22 -15 - 2 -30 7 -28 25 3 2 28 -12 30 14 13 -30 13 - 6 19 2 3 36 20 32 6 12 -30 11 -12 14 - 6 4 49 9 36 24 14 -12 22 2 20 -38 5 47 7 34 28 18 6 32 18 26 -35 6 40 22 31 21 25 - 8 36 9 28 -31 7 36 13 33 23 22 - 7 22 -30 28 -21 8 30 - 3 29 16 26 - 6 33 -31 33 8 9 28 - 6 20 - 6 28 8 42 -30 32 15 10 30 15 34 14 34 20 35 -28 34 14 11 37 - 5 34 20 35 16 27 17 36 5 12 37 - 3 22 -11 16 6 37 18 33 - 5 13 38 - 3 17 -16 6 - 5 45 21 45 17 14 34 0 15 -13 4 -35 36 22 55 25 15 27 6 19 4 5 -30 39 19 56 15 16 20 4 14 -24 20 2 40 -12 60 16 17 17 5 15 -31 35 13 42 -11 54 9 18 9 - 5 18 - 1 31 25 37 5 51 12 19 0 -14 22 5 27 14 25 8 52 13 20 - 2 -39 29 12 20 13 30 -10 48 17 21 9 -41 27 13 29 14 33 15 44 32 22 14 -29 28 15 34 20 40 21 36 25 23 17 7 25 4 36 25 43 27 37 1 24 27 5 32 13 35 28 35 23 39 4 25 28 9 33 25 28 21 32 11 39 16 26 26 15 29 18 28 14 28 -20 37 5 27 27 11 18 -10 18 -12 27 6 42 5 28 27 7 4 -40 16 4 26 -29 51 18 29 26 3 9 -19 9 -17 52 15 30 30 5 5 -27 -14 -23 50 16 31 8 -19 - 5 -51 46 31 LAKE LOUISE, ALBERTA - 194-7 - 194-8 November December January February March Day Max. Min. Max. Min. Max. Min. Max. Min. Max. Min. 1 36 20 30 k ^ 11 28 - 2 31 4 2 37 23 29 9 27 9 31 ~2o 23 0 3 32 18 24 9 22 5 22 -11 31 -31 4 31 6 29 6 23 18 15 4 36 -19 5 31 -14 20 -11 24 - 4 15 -37 38 -13 6 26 0 26 - 5 30 7 20 -34 40 - 6 7 33 17 18 - 9 21 - 9 14 -10 33 13 8 36 19 20 5 25 - 2 12 9 21 - 6 9 32 - 2 20 - 2 27 -13 5 -12 19 -34 10 33 8 21 - 3 24 - 4 30 -36 30 -35 11 35 13 22 3 21 11 25 -24 24 -16 12 29 16 21 4 17 -22 18 -10 13 32 17 29 17 18 - 3 24 8 14 25 10 29 10 28 4 26 5 15 22 9 20 - 8 29 - 2 29 22 16 26 2 19 -15 25 - 3 24 17 17 24 -11 35 13 32 3 22 12 18 30 14 31 12 23 -17 20 - 6 19 19 -12 27 8 31 13 19 -36 20 20 11 27 - 4 28 5 23 -27 21 23 -20 28 7 34 23 28 0 22 19 -10 25 6 37 20 27 13 23 25 13 33 21 34 21 28 - 3 24 27 21 34 22 29 7 32 12 25 30 9 36 - 2 21 -16 34 14 26 36 28 38 23 18 -30 32 17 27 32 - 5 35 28 23 - 9 32 13 28 33 - 2 30 23 21 0 26 -18 29 32 - 8 24 11 31 -10 24 -28 30 ?5 - 3 19 -11 24 3 31 20 3 26 - 9 LAKE LOUISE, ALBERTA - 1948 - 1949 November December January February March Day Max. Min. Max. Min. Max. Min. Max. Min. Max. Min. 1 25 12 22 8 15 -15 59 - 9 2 26 10 16 -10 14 -32 46 - 5 3 21 - 4 8 -38 12 -24 u 7 4 14 - 6 13 -19 9 -17 42 22 5 15 - 5 12 -14 14 -38 48 24 6 16 -12 32 9 12 -22 41 14 7 18 2 37 11 6 - 4 36 17 8 VOLUME 15 6 9 -25 16 -18 27 11 9 6 -27 7 -43 20 - 7 35 -24 10 14 - 2 8 -38 23 11 40 -18 11 MISSING 18 - 2 14 -16 19 -10 43 -14 12 12 - 7 17 -18 11 -42 33 13 13 5 -16 20 -14 10 -31 25 - 4 14 FOR 13 -34 24 0 12 -14 26 -33 15 11 -31 25 - 2 16 -18 21 -12 16 12 -29 2£ -19 23 5 38 2 17 NOVEMBER 6 -32 22 9 27 7 37 11 18 12 -22 15 -15 -10 -15 41 25 19 12 0 7 -38 8 -43 40 24 20 22 11 - 6 -48 13 -42 AA 25 21 17 - 8 6 -31 31 9 A2 27 22 10 -19 1 -34 38 21 38 24 23 13 -28 - 6 -49 37 24 47 18 24 20 -26 5 -48 38 3 41 6 25 13 -23 4 -34 45 9 39 15 26 14 -27 13 - 2 48 - 6 36 16 27 14 - 8 16 -10 47 - 8 35 19 28 18 -19 18 -32 46 - 6 36 2 29 15 - 5 18 2 35 5 30 18 - 3 25 -18 42 20 31 19 3 21 -13 AA 6 LAKE LOUISE, ALBERTA - 194-9 - 1950 November December January February March Day Max. Min. Max. Min. Max. Min. Max. Min. Max. Min. 1 55 25 33 12 -10 -36 2 52 19 34 28 -21 -54 33 - 3 3 53 18 24 4 -11 -53 34 19 4 54 20 18 - 6 - 1 -42 35 17 5 57 21 26 0 - 3 -34 29 - 2 6 53 20 27 0 4 -33 31 10 7 51 19 22 3 8 - 5 32 9 8 40 22 18 -20 17 -18 NO 23 7 9 37 24 23 0 - 5 -26 13 - 5 10 38 23 18 -18 - 6 -22 17 -21 11 31 21 14 -22 RECORD 26 -40 12 35 20 16 -13 19 -32 13 37 16 15 -20 -18 -36 25 - 4 14 38 11 17 - 2 -18 -35 FOR 29 3 15 37 21 21 12 -14 -52 31 9 16 38 9 18 0 -18 -54 34 16 17 43 27 16 - 4 - 6 -39 5 FEBRUARY 19 2 18 40 29 8 -17 - 5 -44 39 - 1 19 39 21 0 -38 4 -26 38 19 20 37 6 6 -31 14 - 1 39 7 21 39 7 11 -14 30 7 33 10 22 40 17 20 4 8 -11 38 8 23 27 18 2 -10 -22 38 14 24 5 13 "4 -21 -55 40 - 5 25 36 28 17 -20 -13 -63 40 17 26 34 24 14 - 6 0 -44 38 18 27 42 28 -11 4 -51 38 7 28 36 26 -11 -13 3 -51 35 16 29 30 22 32 -15 - 1 -42 30 4 30 29 13 31 0 - 1 -46 33 11 31 8 -11 - 4 -38 38 - 8 LAKE LOUISE, ALBERTA - 1950 - 1951 November December January February March Day Max. Min. Max. Min. Max. Min. Max. Min. Max. Min. 1 33 - 2 12 - 5 23 5 5 -22 26 -24 2 37 -12 13 - i i 15 - 8 11 - 2 25 -15 3 35 8 10 7 13 0 26 8 21 -19 4 40 30 8 -28 6 -30 26 12 5 - 4 5 36 26 4 -31 15 -24 22 5 - 5 -20 6 30 16 20 -10 17 -22 14 -28 - 6 -48 7 45 12 26 - 8 18 -13 18 -23 - 2 -45 8 22 - 6 27 3 19 - 4 28 12 - 5 -39 9 22 -17 23 - 4 20 - 9 37 25 - 6 -23 10 29 5 34 21 21 -13 32 14 4 20 11 31 13 31 14 25 - 9 22 -29 25 -33 12 33 - 7 33 23 28 1 24 -28 36 0 13 23 - 6 33 14 22 5 27 -27 35 13 34 20 7 23 0 30 10 32 -20 38 22 15 18 2 24 4 21 2 28 -11 36 32 16 22 12 29 7 25 8 34 -12 28 18 17 33 1 27 12 23 9 28 - 3 28 -18 18 12 -14 25 - 3 18 -17 35 - 8 34 -15 19 22 -15 20 - 8 10 -30 29 1 45 7 20 25 8 30 5 19 - 5 28 3 48 27 21 29 23 35 25 20 - 5 29 2 39 25 22 26 -10 36 27 20 4 30 -25 38 11 23 12 -34 35 26 25 2 31 -24 33 18 24 29 8 34 26 19 4 29 - 4 38 24 25 27 9 33 21 10 -15 31 8 42 22 26 35 23 24 -16 11 -19 31 - 6 37 27 27 38 26 22 - 6 - 3 -49 27 - 1 34 31 28 32 14 28 14 4 -30 25 -15 38 0 29 18 - 5 24 3 4 -43 46 17 30 13 - 2 30 21 7 -45 42 27 31 23 6 6 -43 47 25 LAKE LOUISE, ALBERTA - 1951 - 1952 November December January February March Day Max. Min. Max. Min. Max. Min. Max. Min. Max. Min. 1 22 -22 32 26 —33 -m "°32 19 22 8 2 29 12 29 12 2 -30 32 13 18 - 8 3 26 2 28 8 9 -27 25 9 26 - 5 4 36 16 25 5 15 -15 31 18 32 5 5 36 6 24 - 5 16 -13 29 9 32 8 6 37 3 20 - 4 20 3 34 20 35 -12 7 a 7 14 -22 18 -14 30 18 34 -16 a 34 22 17 -17 10 -16 28 3 38 10 9 33 16 16 -12 21 - 3 34 14 35 8 10 30 4 19 6 19 11 32 8 33 14 11 34 18 30 18 13 -14 40 - 3 33 - 8 12 32 20 34 25 11 -20 31 4 37 5 13 34 18 25 1 16 -17 31 6 30 4 14 27 14 15 -15 - 5 -12 28 -20 36 - 9 15 30 10 7 -24 -10 -20 29 2 38 -16 16 23 -23 9 -22 11 -24 26 -12 36 - 8 17 24 -13 12 -14 18 5 21 5 34 0 18 28 -10 3 - 8 20 13 20 - 9 33 10 19 35 2 - 4 -18 21 3 22 -25 30 0 20 35 16 - 2 -40 13 -13 23 -29 32 - 6 21 24 - 9 7 -15 -14 -22 22 -28 30 -21 22 28 5 9 - 7 -16 -38 22 -24 36 -11 23 28 3 13 -24 2 -35 26 -32 39 24 24 29 - 7 6 -24 11 - 6 29 -20 40 20 25 26 2 7 -32 23 2 32 - 2 42 25 26 29 16 3 -30 29 17 32 6 43 16 27 34 18 12 -22 32 22 27 -15 48 26 28 32 22 11 -13 37 24 26 -14 a 23 29 28 16 10 2 37 21 31 -14 36 10 30 27 3 5 -22 33 22 33 20 31 -15 -48 34 20 30 16 LAKE LOUISE, ALBERTA - 1952 - 1953 November Dec ember January February March Day Max. Min. Max. Min. Max. Min. Max. Min. Max. Min. i li 22 24 0 - 5 28 19 -1 26 33 % -8 3 38 26 5 32 12 30 20 28 - 5 4 42 29 13 30 15 29 17 34 22 5 40 34 13 22 -10 27 7 34 24 6 39 23 18 13 -30 32 - 7 36 11 7 39 25 5 -15 -17 30 14 42 27 8 39 NO 26 8 29 -15 30 -15 48 15 9 40 23 6 34 4 26 -20 52 14 10 34 25 17 33 8 26 9 42 22 11 43 RECORD 27 0 25 6 42 12 46 12 12 40 -25 -4 33 4 31 8 39 13 13 35 39 15 -14 -16 32 18 35 - 5 14 33 FOR 33 8 - 7 -29 25 - 6 30 -13 15 41 24 9 17 -34 30 18 32 - 9 16 37 22 5 31 14 25 - 4 39 5 17 35 MIN. 26 11 25 10 29 8 37 22 13 28 22 5 22 - 7 30 -20 29 - 9 19 33 24 -22 28 1 27 -25 35 6 20 31 14 -17 32 20 32 -21 35 -10 21 25 21 -18 29 11 37 - 4 39 - 2 22 28 25 3 29 22 32 14 35 0 23 33 19 -12 26 9 34 -15 39 - 8 24 29 18 -14 36 23 32 - 8 53 11 25 22 9 -15 28 11 36 - 4 40 27 26 17 14 -20 22 14 36 28 42 10 27 23 15 -18 19 - 4 33 5 44 6 28 24 20 - 2 26 13 23 3 45 25 29 15 27 5 36 22 37 6 30 21 29 14 32 26 40 18 31 31 12 34 15 32 15 LAKE LOUISE, ALBERTA - 1953 - 1954. November December January February March Day Max. Min. Max. Min. Max. Min. Max. Min. Max. Min. i 8 23 22 a 25 22 4 1^  ?l 8 3 40 7 24 8 26 16 40 20 32 -13 4- 42 3 23 4 24 19 42 25 36 -22 5 40 18 23 13 31 6 47 24 34 -15 6 35 25 27 10 31 19 46 13 32 21 7 38 23 21 2 27 17 46 11 35 5 8 42 25 20 5 30 4 45 14 37 18 9 42 29 27 2 18 -25 32 12 46 23 10 65 19 25 5 18 -26 20 6 40 22 11 46 21 22 2 26 - 8 10 - 7 34 -19 12 37 20 28 19 15 -23 9 - 4 35 -14 13 41 20 25 7 16 7 33 2 35 -16 14 36 14 29 18 - 8 -16 33 - 1 28 12 15 35 28 22 -17 -21 -35 27 15 29 -19 16 33 26 22 -18 - 3 -52 35 24 36 - 4 17 29 14 24 - 8 - 2 - 8 39 23 42 19 IB 29 - 3 29 1 ?8 -31 32 16 40 14 19 22 2 30 19 -15 -32 25 9 39 19 20 30 2 24 16 -12 -47 34 20 37 2 21 26 - 1 22 3 -14 -23 36 15 39 1 22 31 11 18 -14 - 6 -25 34 22 32 - 7 23 30 20 22 - 8 -14 -22 36 24 40 - 4 24 31 20 28 17 -12 -19 33 25 33 7 25 30 4 32 24 - 7 -20 30 19 34 - 8 26 38 10 27 14 15 -12 27 8 25 9 27 27 3 28 - 8 14 -21 26 15 20 - 5 28 30 19 28 23 19 7 29 2 17 -41 29 32 18 26 8 17 -21 26 - 2 30 34 12 26 14 26 -11 27 -18 31 30 12 36 20 28 11 APPENDIX 3 DALLY MAXIMUM AND MINIMUM TEMPERATURES FOR WINTER MONTHS, CALGARY, ALBERTA. 1954 to 1956 CALGARY, ALBERTA - 1954 - 1955 November December January February March Day Max, Min. Max. Min. Max. Min. Max. Min. Max. Min. 1 2 61.4 57.6 23.5 32.2 32.1 42.1 W 15.0 19.0 6.2 3.7 m m 3 53.8 25.9 46.5 32.7 33.1 -5i4 37.4 13.6 -7.6 -15.8 4 59.9 35.3 40.0 29.8 35.1 21.9 38.0 15.0 10.7 -23.9 5 67.1 47.3 45.8 16.0 36.3 10.9 29.8 18.6 25.2 -5.5 6 51.7 36.1 49.5 21.5 33.8 21.5 36.8 6.7 42.9 11.8 7 43.0 19.6 38.0 15.6 45.4 24.5 42.0 24.3 52.1 27.8 8 56.5 25.0 32.0 11.9 27.5 16.7 21.5 15.7 37.4 30.0 9 60.5 25.5 43.2 19.7 19.0 -1.8 14.0 2.5 42.3 23.4 10 53.9 24.7 34-2 17.5 27.6 0.6 26.2 -10.7 41.5 32.0 11 43.2 21.5 43.2 13.7 10.7 -8.9 35.1 -4-4 40.2 21.6 12 47.4 26.2 47.4 15.8 43.1 3.4 38.4 3.7 36.6 17.2 13 34.6 21.6 40.1 24.3 39.2 24.3 46.5 24.3 16.2 -1.6 14 54.6 24-3 39.6 16.5 28.2 2.2 46.1 21.6 15.9 4.6 15 56.2 32.7 39.4 19.0 29-0 6.7 41.1 13.7 29.2 -5.5 16 48.6 29.0 48.2 27.1 23.0 10.7 46.1 19.5 28.4 14.4 17 47.8 31.0 56.7 35.8 15.2 9.7 21.1 13.6 39.5 7.6 18 56.4 26.2 58.3 30.0 15.9 -3.9 21.8 -3.7 27.2 19.8 19 52.5 32.8 62.5 25.4 20.3 4.7 29.6 11.9 12.9 5.8 20 53.4 33.3 57.2 28.3 24.1 0.4 34.0 5.8 16.5 -3.7 21 67.6 37.6 60.8 38.1 30.6 1.8 34.0 17.1 25.2 -0.8 22 58.2 32.0 45.3 31.8 42.0 2.9 5.0 -2.9 3.1 -3.7 23 42.2 20.8 27.7 9.8 37.0 21.2 -2.6 -19.8 -5.8 -12.9 24 57.2 ?3.7 28.5 15.2 34.6 5.8 1.9 - 9.4 1.8 -22.9 25 58.2 35.1 17.6 11.5 32.8 16.2 -2.0 -10.5 17.2 -11.5 26 46.6 30.0 41.2 9.3 41.2 13.8 -9.8 -16.9 32.5 -6.0 27 33.3 20.6 34.1 10.0 38.5 22.5 10.0 -27.2 52.0 18.2 28 29.3 19.9 a . 2 24.3 45.0 16.8 24.0 -19.8 49.1 25.5 29 27.7 20.3 -7.2 -11.0 44.2 21.7 56.5 29.9 30 25.5 17.6 16.2 -14.6 47.0 19.5 37.0 28.0 31 29.5 3.6 40.2 15.4 56.1 31.0 CALGARY, ALBERTA - 1955 - 1956 November December January February March Day Max. Min. Max. Min. Max. Min. Max. Min. Max. Min. 1 19.1 13.6 23.1 9.9 34.9 9.7 29.6 -1.4 39.7 4.7 2 18.A 6.4 10.1 - 7.1 28.0 4.5 2JI8 29.6 17.0 3 26.0 11.9 23.1 35.0 4.4 38.9 33.0 20.8 4 20.1 10.7 12.3 -10.3 17.8 7.4 44.2 14.5 14.3 - 1.3 5 34.0 4.2 25.4 - 9.8 3.2 - 4.8 35.2 21.9 2.6 - 1.1 6 38.2 12.2 31.8 8.4 19.6 - 9.8 37.8 23.2 9.4 -12.0 7 57.9 18.6 30.9 8.4 7.3 - 5.4 30.4 14.8 34.7 - 5.8 8 57.0 37.5 35.3 6.6 22.1 - 7.0 43.0 11.3 35.3 5.8 9 50.7 33.1 35.3 10.0 22.2 - 1.8 39.9 28.4 5.2 - 0.8 10 39.6 13.5 30.4 7.6 13.6 - 0.1 36.1 27.2 - 5.0 -15.4 11 - 5.4 -13.5 34.0 14.9 4.2 - 3.1 30.8 11.0 29.3 -18.4 12 - 7.1 -20.7 27.1 5.6 13.2 - 2.9 19.2 9.5 30.1 15.1 13 - 6.2 -25.7 . 14.8 2.1 0.1 - 4.7 - 8.4 -14.2 28.6 13.0 14 - 4.8 -12.7 14.0 -10.7 - 3.2 -12.7 -19.0 -30.0 35.9 10.9 15 6.2 - 9.1 16.6 - 6.4 - 5.0 -16.3 -13.4 -31.0 44.1 18.6 16 9.7 -15.9 15.6 0.7 - 7.8 -26.3 22.9 -27.8 47.7 25.4 17 4.0 -14.9 2.2 -13.1 6.9 -15.0 22.8 10.5 47.6 27.1 18 22.6 -15.3 -10.6 -33.2 40.1 - 3.6 20.5 - 7.8 57.3 31.2 19 36.8 - 1.6 - 9.8 -16.7 26.3 9.0 23.3 2.9 57.3 31.3 20 37.4 0.1 21.9 -12.4 31.2 3.4 0.2 -10.8 33.9 24.5 21 20.3 1.3 24.0 - 8.0 20.6 9.7 - 0.8 - 5.7 31.2 22.4 22 3.2 - 9.7 32.2 - 2.8 23.4 4.6 0.4 -12.2 49.1 25.2 23 - 5.1 -15.6 -10.6 -22.8 33.8 - 0.5 5.0 - 7.0 54.7 25.5 24 - 1.1 -14.8 -12.6 -28.8 28.2 13o2 2.0 -14.1 45.0 27.5 2§ 0.4 - 7.7 33.2 -18.8 24.7 13.4 1.1 - 5.1 44.1 21.7 26 3.0 -16.0 33.6 - 2.5 8.8 - 1.3 16.0 -20.1 32.8 27.1 27 12.6 -10.8 - 0.7 - 7.5 - 0.6 - 4.4 29.1 - 5.5 34.3 23.2 28 7.1 -12.3 20.0 -10.3 - 2.8 - 6.8 35.0 - 4.5 33.8 10.3 29 16.1 - 0.4 35.4 - 3.1 - 6.6 -16.4 40.1 19.9 43.0 22.3 30 35.9 3.5 29.6 6.4 5.4 -26.3 44.1 22.7 31 38.4 0.4 17.2 -15.7 25.8 21.4 195 APPENDIX 4 Mean Monthly Temperatures, Banff, Alberta Year November December January February March 1920-21 27.1 19.1 17.8 22.1 23.1 1921-22 22.2 11.8 8.6 4.1 22.8 1922-23 25.4 7.8 15.4 17.8 26.1 1923-24 31.0 17.8 12.4 25.1 25.0 1924-25 22.1 8.8 15.5 22.6 28.1 1925-26 27.4 27.2 19.8 28.0 33.2 1926-27 26.2 21.2 9.9 16.2 26.1 1927-28 16.0 2.4 19.5 22.6 29.2 1928-29 28.6 17.0 4.3 12.0 30*7 1929-30 26.6 14.3 - 3.9 24.5 25-4 1930-31 28.2 24.0 25.1 25.9 26.7 1931-32 20.2 18.8 10.6 18.4 23.0 1932-33 26.5 13.8 16.4 12.8 25.0 1933-34 34-0 9.2 22.8 22.6 29.3 1934-35 32.4 16.4 9.2 13.5* 20.4** 1935-36 23.4 23.4 - 8 .0* * 1936-37 28.8 14.1 - 4.1 10.7 28.1 1937-38 23.0 15.4 16.8 10.1 28.0 1938-39 23.8 17.0 22.4 12.3 27.8 1939-40 35.8 25.0 10,5 19.0 32.2 1940-41 17.1 22.2 20.0 20.6 30.6 1941-42 30.0 16.8 14.8 16.0 27.2 1942-43 19.7 15.6 2,9 24.6 18.1 1943-44 29.0 19.4 20.0 17.4 23.1 1944-45 27.2 14.9 19.0 17.8 27.0 1945-46 18.1 15.2 20.7 21.5 30.4 1946-47 18.1 16.4 14.9 17.8 21.6 1947-48 22.9 22.9 21.0 7.1 20.6 1948-49 26.2 7.6 4 .0 9.4 27.2 1949-50 36.8 9.0 -16.0 22.0 21.6 1950-51 19.7 21.4 6.0 14.0 16.8 1951-52 25.4 6.4 9.0 19.4 22.6 1952-53 24.3 19.4 15.9 23.1 27.4 1953-54 31.6 23.4 3.5 28.8 21.1 Anthracite, Banff not recorded. Lake Louise, Banff not recorded. 196 APPENDIX 5 Mean Monthly Temperatures, Lake Louise, Alberta Year November December January February March 1932-33 21.1 5.7 10.0 6.2 20.2 1933-34 28.2 5.1 15.3 12.9 25.4 1934-35 25.6 14.7 6.0 16.6 16.2 1935-36 17.1 15.6 11.3 - 8.0 20.4 1936-37 22.1 8.0 -11.1 5.1 24.7 1937-38 20.0 9.8 7.8 7.8 25.0 1938-39 17.8 9.9 16.2 4.6 22.0 1939-40 20.4 4.0 17.1 30.0 1940-41 13.9 15.7 15.1 17.6 27.9 1941-42 22.7 11.5 7.1 11.1 1942-43 15.3 12.1 - 1.2 17.6 1943-44 23.8 10.6 13.0 16.6 18.0 1944-45 22.2 8.2 12.6 13.3 1945-46 15.2 9.9 14.0 15.6 1946-47 16.5 12.4 9.0 15.5 23.0 1947-48 17.8 15.7 12.8 8.9 1948-49 1.8 2.4 5.0 23.0 1949-50 30.5 5.6 -19.4 17.5 1950-51 15.7 14.8 2.0 9.6 13.6 1951-52 18.3 1.4 3.7 12.6 19.3 1952-53 12.5 13.4 16.8 15.4 1953-54 25.2 16.0 - 0.4 23.6 15.2 197 APPENDIX 6 Comparative Records, Calgary, Alberta Monthly and Annual Averages and Extremes for Total Period Observations Have Been Taken (1885 - 1955) Month Mean Max. Mean Min. Monthly Mean Absolute Year Min. Lowest Monthly Mean - Year Jan. 25.3 4.4 14.8 - 48 1893 -13.6 1950 Feb. 27.0 6.4 16.7 - 49 1893 -12.0 1936 Mar. 36.9 15.4 26.1 - 34 1896 8.8 1899 Apr. 52.3 27.4 39.8 - 22 1954 25.1 1954 May 62.1 36.8 49.4 2 1954 U.2 1907 June 68.2 43.7 55.9 26 1904 49.4 1902 J u l y 75.7 48.2 61.9 32 4 Years 56.3 1912 Aug. 73.1 45.6 59.3 28 1886 54.3 1911 Sept. 63.7 38.0 50.8 8 1926 42.8 1926 Oct. 54.. 0 29.9 41.9 - 8 1887,1939 32.8 1919 Nov. 38.7 17.9 28.3 - 31 1893 2.4 1896 Dec. 28.6 9.2 18.9 - 45 1924 3.1 1933 No. of years obs. in 5 3 53 53 69 69 198 APPENDIX 7 BANFF, ALBERTA Annual and 5 - Year Running Mean Temperatures YEAR ANNUAL 5 - YEAR YEAR ANNUAL 5 - YEAR 1896 34.3 1926 38.1 36.8 1897 34.7 1927 33.2 36.5 1898 36.2 1928 37.7 36.5 1899 34.0 1929 35.2 36.5 1900 37.3 35.3 1930 36.3 36.1 1901 36.5 35.7 1931 38.1 36.1 1902 34.8 35.8 1932 35.6 36.1 1903 35.1 35.5 1933 35.4 36.1 1904 36.4 36.0 1934 39.4 37.0 1905 37.3 36.0 1935 35.5 36.8 1906 37.5 36.2 1936 36.5 1907 34.8 36.2 1937 36.8 1908 37.6 36.7 1938 36.7 37.2 1909 33.4 36.1 1939 37.9 36.7 1910 37.0 36.1 1940 37.4 37.3 1911 33.5 35.3 1941 38.0 37.5 1912 35.9 35.5 1942 35.7 37.1 1913 34.5 34.8 1943 35.0 36.8 19U 37.0 35.6 1944 37.0 36.6 1915 37.9 35.8 1945 34.8 36.1 1916 32.3 35.5 1946 36.2 35.7 1917 34.9 35.3 1947 36.6 35.9 1918 37.0 35.8 1948 34.5 35.8 1919 34.9 35.4 1949 35.2 35.4 1920 35.7 35.0 1950 32.6 35.0 1921 36.0 35.6 1951 31.8 34.1 1922 34.6 35.6 1952 36.4 34.1 1923 37.7 35.8 1953 39.7 35.1 1924 35.5 35.9 1954 37.4 35.6 1925 38.1 36.4 1955 34.9 36.0 APPENDIX 8 1 9 9 Mean Monthly Temperatures for Selected Stations - WINTER MONTHS -Year Month GOLDEN ROCKY MOUNTAIN HOUSE EDS ON EXSHAW 1939 Jan. 23.4 17.8 19.3 26.3 Feb. 11.4 1.0 11.2 13.6 Mar. 28.7 23.7 25.0 28.6 1939-40 Nov. 32.8 38.6 34.7 40.7 Dec. 26.1 27.4 25.0 30.7 Jan. 13.2 - 3.1 6.0 15.2 Feb. 26.0 14.2 19.0 Mar. 40.1 26.8 32.0 1940-41 Nov. 18.6 12.5? 15.2 20.0 Dec. 26.8 16.0P 16.6 27.0 Jan. 22.8 8.6P 7.8 21.8 Feb. 23.9 14. OP 15.5 22.8 Mar. 37.9 23.8P 29.5 32.0 1941-42 Nov. 29.1 31.6 26.6 34.3 Dec. 22.2 15.7P 10.6 21.0 Jan. 11.0 20.7P 23.0 20.9 Feb. 19.7 13.2f 17.4 19.6 Mar. 33.2 29.9^ 27.8 30.8 1942-43 Nov. 25.9 15.5P 15.6 21.3 Dec. 17.8 5.4 21.0 Jan. 0.2 - 3.OP* - 1 . 9 Feb. 23.7 16.7 22.5 26.1 Mar. 24.2 12.6 18.2 18.4 1943-44 Nov. 28.4 32.4 30.6 34.4 Dec. 17.8 20.0 23.6 25.2 Jan. 15.6 13.8 18.4 25.9 Feb. 19.8 13.0 16.8 19.0 Mar. 27.7 20.9 23.8 22.6 1944-45 Nov. 32.0 24.8 22.6 27.8 Dec. 13.5 17.1 15.5 19.2 Jan. 16.5 11.5 14.4 23.0 Feb. 20.3 9.8 15.5 20.4 Mar. 31.8 26.5 28.4 30.0 1945-46 Nov. 28.6 9.7 10.8 17.8k Dec. 19.8 8.6 10.3 21.0k Jan. 21.4 12.1 16.2 24.6k Feb. 22.7 11.7 16.8 24.6k Mar. 35.5 28.2 31.2 32.2k p = Penhold k = Kananaskis 200 Mean Monthly Temperatures for Selected Stations (cont.) TEAR MONTH GOLDEN PENHOLD EDS ON KANANASKIS 1946-47 1947-48 1948-49 1949-50 1950-51 1951-52 1952-53 Nov. 23.4 15.9 17.0 20.4 Dec. 17.6 9.2 9.2 20.0 Jan. 15.1 10.4 14.0 19.4 Feb. 22.0 5.6 8.9 15.6 Mar. 35.4 18.2 23.1 23.5 Nov. 26.8 21.9 24.2 24.4 Dec. 24.0 17.6 19.3 25.2 Jan. 16.3 19.1 19.6 25.4 Feb. 15.2 3.3 7.5 10.6 Mar. 29.9 13.2 18.7 19.6 Nov. Dec. 7.3 3.5 3.0 14.0 Jan. 0.6 2.4 7.6 11.7 Feb. 12.4 - 2.4 2.6 9.0 Mar. 32.1 24.2 26.6 27.5 Nov, 34.0 36.4 35.2 Dec. 16.4 2.4 1.0 8.0 Jan, -12.4 -19.2 -16.4 -19.2 Feb. 19.8 10,4 11.1 Mar. 31.1 17.6 19.4 Nov. 26.4 15.2 9.4 20.8 Dec. 23.6 12.2 14.6 26.7 Jan. 10.2 0.0 2.4 10.4 Feb. 16.0 7.6 10.2 15.1 Mar. 25.5 8.0 13.2 16.2 Nov. 27.0 23.0 23.1 28.7 Deo. 9.1 4.8 6.8 12.4 Jan. 10.5 - 2.8 - 1.2 7.9 Feb. 24.5 12.1 20.2 24.0 Mar. 31.7 14.2 22.8 23.2 Nov. 26.0 28.8 29.8 29.8 Dec. 23.3 18.3 17.8 25.6 Jan. 23.8 4.5 1.6 19.5 Feb. 27.3 19.0 25.6 25.5 Mar. 34.6 22.9 25.4 28.8 APPENDIX 9 A i r Mass Sim-wry - B«u Vallav Drainage 201 Year Month Total No. a l l types No. cP (cA) No. days area in f rontal passages. cold f ronts . cP (cA)alr. 1920-21 1921-22 1922-23 1923-24 1924-25 1925-26 1926-27 Nov. 6 2 6 Dec. 5 3 14 Jan. 9 3 17 Feb. 9 2 11 Mar. 8 3 11 Nov. 6 2 14 Deo. 10 3 13 Jan* 12 5 14 Feb. 4 2 22 Mar. 8 2 5 Nov. 9 2 3 Dec. 7 2 10 Jan. 10 5 16 Feb. 10 3 12 Mar. 16 7 12 Nov. 6 0 0 Dec. 8 2 8 Jan. 8 3 15 Feb. 7 2 8 Mar. 10 5 8 Nov. 9 2 10 Deo. 7 4 20 Jan. 11 4 16 Feb. 10 5 15 Mar. 8 1 16 Nov. 9 2 8 Deo. 11 4 9 Jan. 7 1 0 Feb. 6 2 6 Mar. 5 1 9 Nov. 5 1 7 Dec. 11 5 11 Jan. 11 3 ia Feb. 6 4 9 Mar. 5 0 0 202 A i r Mass Summary - Bow Valley Drainage (cont.) Year Month Total No. a l l types No. cP (cA) No. days area in f rontal passages. cold f ronts . oP (cA) a i r . . 1927- 28 Nov. 7 4 15 Dec. 4 3 23 Jan. 10 4 7 Feb. 7 1 5 Mar. 4 1 7 1928- 29 Nov. 9 1 0 Dec. 6 3 3 Jan. 6 3 21 Feb. 5 2 14 Mar. 12 3 9 1929- 30 Nov. 12 1 4 Dec. 7 2 17 Jan. 1 0 30 Feb. 7 2 5 Mar. 12 4 14 1930- 31 Nov. 7 1 8 Dec. 9 0 0 Jan. 9 2 0 Feb. 10 1 1 Mar. 8 2 11 1931- 32 Nov. 8 4 11 Dec. 9 3 8 Jan. 10 4 18 Feb. 8 3 18 Mar. 8 4 17 1932- 33 Nov. 8 2 5 Dec. 6 3 8 Jan. 11 5 19 Feb. 7 3 14 Mar. 4 1 7 1933- 34- Nov. 8 3 7 Dec. 4 2 28 Jan. 9 2 5 Feb. 6 2 13 Mar. 12 6 16 A i r Mass Summary - Bow Valley Drainage 203 Year Month Total No. a l l types No. cP (cA) No. days area in frontal passages, cold f ronts . eP (oAjair. 1934-35 Nov. 12 3 3 Dec. 3 2 9 Jan. 9 5 17 Feb. 7 2 2 Mar. 8 4 18 1935-36 Nov. 5 1 12 Dec. 8 3 7 Jan. 5 2 22 Feb. 1 0 26 Mar. 6 3 11 1936-37 Nov. 9 0 7 Dec. 8 4 18 Jan. 6 3 22 Feb. 5 3 13 Mar. 8 3 6 1937-38 Nov. 8 2 5 Dec. 8 2 12 Jan. 32 6 6 Feb. 4 2 18 Mar. 6 2 0 1938-39 Nov. 7 3 6 Dec. 5 1 5 Jan. 32 4 8 Feb. 5 2 17 Mar. 10 3 13 1939-40 to 1944-45 missing. 1945-46 Nov. 13 4 32 Dec. 13 5 7 Jan. 12 5 4 Feb. 6 2 3 Mar. no cA or cP a i r . 1946-47 Nov. 10 3 6 Deo. 8 5 13 Jan. 8 3 11 Feb. 8 3 17 Mar. 6 1 13 Air Mass Summary - Bov Val ley Drainage (eont.) 204 Year Month Total No. a l l types No. cP (cA) No. days area f ronta l passages cold f ronts . in eP (cA) a i r . 1947-4.8 Missing 1948-49 Nov. 13 7 10 Dec. 12 7 16 Jan. 16 9 15 Feb. 13 6 16 Mar. 7 2 11 1949-50 Nov. 4 0 0 Dec. 9 5 14 Jan. 3 2 25 Feb. 3 4 8 Mar. 4 1 4 1950-51 to 1951-52 missing 1952-53 Nov. 6 3 4 Dec. 7 4 5 Jan. 21 10 15 Feb. 13 2 4 Mar. 26 6 10 1953-54 Nov. 23 4 4 Dec. 21 5 9 Jan. 20 8 22 Feb. 17 6 6 Mar. 18 7 19 1954-55 Nov. 20 3 3 Dec. 16 3 4 Jan. 13 4 12 Feb. 18 6 10 Mar. 13 3 16 1. Sequential sampling of the lodgepole needle miner. For . Chron. 28(2)157-60. 1952. 2. Analysis of a population sampling method for the lodgepole needle miner in Canadian Rocky Mountain Parks. Canad. Ent . 84.(lO)*316-321 1952. 3. Distribution and l i f e history of the lodgepole needle miner, Recurvaria sp. (Lepidoptera*Gelechiidae) in Canadian Rocky Mountain Parks. Canad. Ent. 86(l):l-13. 1954-4. l i f e tables for the lodgepole needle miner, Recurvaria starki, Free. (Lepidoptera:Gelechiidae) Proc. Tenth Int. Congr. Ent . August, 1956. In Press. 5. (with Henson, W.R. and W.G. Wellington). Effects of the weather of the coldest month on winter mortality of the lodgepole needle miner, Recurvaria, sp. In Banff National Park. Canad. Ent. 86(l):13-19. 1954. 6. (with K. Graham). Insect population sampling (General consider-at ions) . Forest defoliators by R.W. Stark. Proc. Ent. Soc .B.C. 1954. 7 . (with J . A . Cook). The effects of defol iat ion by the lodgepole needle miner (Recurvaria starkjL F r e e . ) . In Press. Forest Science. In addition to the above publications appearing in s c i e n t i f i c journals,17 sc ien t i f i c notes have appeared in the Bi-Monthly Progress Report, Division of Forest Biology, Department of Agriculture. A lso , yearly Annual Reports (nine) have been prepared which are available for s c i e n t i f i c use with the authors* permission. R 19 116°30' R 18 R 17 116V R 16 A R 15 Km I. Bah .V, \ , 10802 ^ J, U»MFMJtf0UND Mt.yCOirLkE ^ \ ^olhire-Ccfl" MiTN'. ;cvpi\ M.TN ?i'"XH hi O PK. 1056 '^-••9234 ' MT. McARTHUFf 8569 47 VBioWeWiv^ ffifOLLTI N DER ^\ fl \ MTN. *7V, YilLlLipUT W^ W^i —T""'>-116°oo' R 14 R 13 115°45' R 12 115°30' R l l R 10 115V R 9 ATA^ACl PK 5 S - l * U 'ULP.IT MARPOLE ® 2 \ \ \$f >*, MICHAEL Tp 28 Tp 27 . MT. C ^ R V O N ' l ^ f e - ™ VANGUARD ftK 8086 )/ !i-CATHEDRAL/MTN. l 6 4 W | ( f W4! irt! J 0706' ARDSON A5PTARMIG( 9BB5 - -^  mm FOSSIL PIKA" .EG • . 8604 "l% • 796<> MT. ODARAY aa7(M o'Bara I i\MT. HI MT. OWEN • 10128 Tp 37 Tp 36 7, i '/ , -iT^-H MT.VSY AVENS)\\| UWET MTN: i.-WfiV';  1 r4>— ! - H — :|RR6S MTN. MT. COCKSCOMB ;^|MT. yfiYLME 1375-K-_. JWJKanu Ore* Tp 34 Tp 33 Tp 31 Tp 30 p062 i, MT. DHEWSTER . ,' ;.Y - -•'•• - \ v ' j 87-20,'• \\ / N O R T H W E S T ^ T V p ^ f f c ^ y ^ P O R T I O N On same scale "' ••'^••Vbiv+siON M;TN THE DECLINATION OF THF. COMPASS NEEDLE 1954 Tho declination of the compass needlo al any place along a red lino is the declination given on thai red line. At other places the declination is between those given on the neighbouring ted linos; thus at the place marked A, the declination is between 23°00'E and 23°30'E. The declination ol the compass needle is decreasing A% mlnules annually. ..5§r ^OLDENS'GLE PK! JULIEN ^  ;,!(WT. ^AMBRAI \ ST VA^ENCIEN- r " ^ T-''rQI60 ^ f;A riGUt: M'f\N' I M N/.;eB '^ v tf-2 ,'r. i j l^';X J / l "V''"MJ.cos" i t ) o — ;M6!.' , .^ -.STRAHA Frr.nJirtel<!~ v,.....NNy • ^ s - - ' ' P R I O R P MT. CONWAY 1 .'(• 101.7$ ••. v^ .lvtTvD'E MARGE Sk^ f; ••• 9890 BREAKER MTN,Ki (•••/( , :'•. <a»^ "uT:-. ""•''/ Mummery Olacinrsj '\ v ib320'',; i • ' ' -• R 24 117°iS' R 23 R22 117V R 21 116 V R 19 ft TNCTAN PK.? / Pass/ 9817 i OCTOPUS MTN. ,N 9600 ' ^•.NASSWAL'D'PK 11 i l l S-G«lden \VaHejL A ValleVpf the Rocks I 9163 •¥ES.TOR PK Mt THE MARSHAL NU'B PK. 90H6 • / ^  \/ \ -v— S -I—/ 1 ' i s " i f / V! / _,' "=^ARK ENTRANCE -/'" //--' V_. Z,» v .1 I !./~^ 1^ V / 1-%^ ' / *«nB V „ ,'V ( ^ \ V/ —v ' */.>wv' •-, K - 1 - A -* I A - \ — - v > ,\ vy* \ 1 ' '^VVal'e-y'-cr \ V T \ ^ ^ ^ ' L ^ ^ V J A K L E ^ ; " #NT A&INIpNE PA'f?r/fe;^-^nPIJ p, E D G W O O D v ,7 '• W 0 N D , E 5- P^ VI iP'ftOVlNCIALl - / l H E T O W i R . S ' ' JTt;i'K Coal M 4297 Y / 4 - G r W T O MTN.t '--—J NV 1 ' , IN IjB^NJ^ ; / >• , vMARVEL / ^ T i . All.CANmR'A;^  IJ^ RED MA '.MT. LOllGHEED 1 \ 10190 ^ ' V - - : ' / - ' -PIGEO>r-"MTN TTV3T SPARROWHAWK >_/ \ \ BULLER1 7 ' MT. GALATEA-115V R13 R 12 MT. KING ALBERT , • i 1 CO 9 8 0 0 115 30' MT. QUEEN ELIZABETH, 9349 •MT. BOGARjl0 10316 \ MT. KIDT7 S J J ' 9 6 0 5 \ , VP r l N F L E X I B L E -MT. LAWSON MT. MURRAY 9920 • , VT\t-RENlCH P J DOJjiatj5Sf^J^LT. HC^ ErlTSON f\^T. KENT . SMITH-DORIRIE^ I , irj3nn MT WILLIAMS Tp 28 Tp 27 51°15' Tp 26 Tp 25 Tp 24 51°oo' Tp 23 Tp 22 Tp 21 R 10 115V R 9 REFERENCE — < L ? Railway. Standard Gauge, Single Track Road Fire Road,not open to public Trail. Telephone Line along Road or Trail I I I I I n i i i Boundary, Provincial — • — • — • Boundary, National Park Sur.nv.d Unsurvwd Boundary, Township Glacier ggggSggj' Sand or Gravel tu**-^^j~-±~^i Contours, interval 250 teet i0°° Post Office ® Telegraph Office © Wardens Cabin with Telephone * Warden's Cabin without Telephone £ Camera Station 426© Park Boundary Monument tD Spot Elevation, in leet above moan sea level ... 928-1 Compiled, drawn and printed at the office ol the Surveyor General, July, 1932. Reprinted with corrections, 1946, Reprinted with corrections at the Surveys and Mapping Branch, Ottawa, 1955. Miles S Kilometres 5 A L B E R T A Scale 1:190,080 1 Inch to 3 Miles 5 20 Kilometres Copies may be obtained from the Map Distribution Office. Department ot Mines and Technical Surveys, Ottawa, at 25 cents each. 50' 115° K E Y M A P 

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