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The aerobiology of the aecial state of the commandra blister rust, Cronartium comandrae Peck, in Alberta. Powell , John Martin 1969

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THE AEROBIOLOGY OF THE A E C I A L STATE OF THE COMANDRA B L I S T E R RUST, CRONARTIUM COMANDRAE PECK, I N A L B E R T A  by JOHN MARTIN POWELL B.Sc, M.Sc,  U n i v e r s i t y o f L o n d o n , 1956 McGill University, 1959  A T H E S I S SUBMITTED I N P A R T I A L F U L F I L M E N T OF THE REQUIREMENTS FOR THE DEGREE OF  DOCTOR OF PHILOSOPHY  i n t h e Department of BOTANY  We a c c e p t t h i s required  thesis  as conforming t o t h e  standard  THE U N I V E R S I T Y OF B R I T I S H COLUMBIA April,  0  I969  John Martin Powell  1969  In  presenting  for  this  an a d v a n c e d  that  degree  the L i b r a r y  study thesis  thesis  shall  I f u r t h e r agree for  Department  f u l f i l m e n t of  the U n i v e r s i t y  make  it  that  of  permission  or  representatives  w i t h o u t my w r i t t e n  of  this  thesis  for  permission  Z> 0>"T A NY  The U n i v e r s i t y o f B r i t i s h V a n c o u v e r 8 , Canada  Columbia  It  requirements I  reference  for extensive  p u r p o s e s may be g r a n t e d  by h i s  the  B r i t i s h Columbia,  freely available for  scholarly  or p u b l i c a t i o n of  Department  at  in p a r t i a l  agree  and  copying  of  this  by t h e Head o f my  is understood  f i n a n c i a l gain  shall  that not  be  copying allowed  i  '  ABSTRACT  Cronartium comandrae Peck i s a heteroecious native rust which i s damaging to several Pinus species i n North America.  I t grows peren-  n i a l l y i n the l i v i n g bark of hard pines producing pycniospores and aeciospores, and develops annually on species of Comandra and Geocaulon producing urediospores, teliospores and basidiospores. Studies were carried out to determine the e f f e c t of various meteorological and b i o l o g i c a l environmental factors on the aeciospore a e r o b i o l o g i c a l phase of the rust.  The main aspects considered included  the factors a f f e c t i n g aeciospore production and release; and the factors a f f e c t i n g aeciospore transport, dispersion, deposition, germination and viability. Aeciospore production occurred from mid May to late August, with the peak period between late May and mid June. produced  spores f o r 35-50 days, and i n d i v i d u a l cankers produced  for up to 95 days. trees.  An average aecium spores  Much v a r i a t i o n occurred from year to year and between  A e c i a l production was interfered with by the a c t i v i t y of rodents,  insects and other fungi on the canker and through r e s i n o s i s , which probably accounted for a 50-55$ reduction of the p o t e n t i a l aeciospore production i n any year.  Fresh rodent damage was recorded on U0-52$ of the  cankers at about 20 locations i n the years I966 to -I968, insect damage on 39-^6$, Tuberculina and Cladosporium i n f e c t i o n on YJ-33?o, on 67-71$.  and r e s i n o s i s  Rodent damage was mainly caused by s q u i r r e l s .  f l o r a l organisms were i s o l a t e d from the cankers and spores.  Some 6k microTuberculina  maxima was mainly responsible for k i l l i n g the infected canker bark and  ii  g r e a t l y reduced  the production o f aeciospores.  undescribed  Cladosporium  lesser role  i nreducing production.  insects, mites obliquus,  were a l s o prominent on t h e cankers  an u n i d e n t i f i e d  One h u n d r e d a n d s e v e n t e e n  cecidomyiidae  Various other  including Dioryctria,  Pissodes  species o f Epuraea  and Paracacoxenus g u t t a t u s were  s c h w a r z i and C y l i n d r o c o p t u r u s d e l e o n i . showed s p o r e s  t o be n o r m a l l y  dis-  0800 a n d 1900 h o u r s , w i t h some e v i d e n c e o f a d o u b l e p e a k  between  1000 a n d 1600 h o u r s , a n d l i t t l e d i s p e r s a l b e t w e e n 2000 a n d 0700  hours.  Turbulent  aeciospore  atmospheric  concentrations.  c o n d i t i o n s were a s s o c i a t e d w i t h a l l peak  Heavy r a i n s  initially  i n c r e a s e d spore  concen-  t r a t i o n s , b u t no d i s p e r s a l o c c u r r e d d u r i n g l o n g h u m i d c o o l p e r i o d s . deposition  c o n c e n t r a t i o n s were v e r y steep  t y p i c a l hollow  curve  deposition gradient.  close t o source,  largely by diffusion. calm a i r .  forest paction  Rapid  Aeciospores  from  w i t h d i s t a n c e from  h a d a n a v e r a g e v e l o c i t y o f 3.23  d e p l e t i o n o f spore  canopy, m a i n l y b y s e d i m e n t a t i o n ,  Spore  a n d showed a  Spore c o n c e n t r a t i o n s  a r t i f i c i a l r e l e a s e p o i n t s were s i m i l a r l y reduced  in  a  s p e c i e s c a u s e d damage t o t h e c a n k e r ,  D a i l y aeciospore p e r i o d i c i t y between  but played  and s p i d e r s were a s s o c i a t e d w i t h t h e r u s t canker.  true mycetobionts.  persed  P e n i c i l l i u m spp. and an  source, cm/sec  c o n c e n t r a t i o n o c c u r r e d under t h e although  aeciospores  h a d good i m -  efficiency. Aeciospores  germinated  on w a t e r a g a r o v e r  1-30°C, w i t h o p t i m u m f o r g e r m i n a t i o n Most a e c i o s p o r e s  germinated  germ tube e l o n g a t i o n a f t e r F r e e w a t e r was n e c e s s a r y germination.  Aeciospores  range  a n d g e r m t u b e g r o w t h c l o s e t o 15°C.  w i t h i n k-5 h o u r s , 8 hours and l i t t l e  f o r germination; germinated  t h e temperature  with a reduction i nrate of i n c r e a s e a f t e r 2k  a l l spores  hours.  swelled prior t o  e q u a l l y w e l l i n the dark  and l i g h t ,  i i i  a n d g e r m i n a t e d o v e r t h e pH r a n g e k.5-8.  G e r m i n a t i o n response on sugar  m e d i a was b e t t e r t h a n o n some o t h e r m e d i a , b u t a d d i t i o n o f a l t e r n a t e host m a t e r i a l t o media d i d not improve g e r m i n a t i o n . D a i l y a e c i o s p o r e c o l l e c t i o n s gave h i g h g e r m i n a t i o n p e r c e n t a g e s for  2-k w e e k s , b u t much l o w e r p e r c e n t a g e s d u r i n g l a t e r  sporulation  period.  A e c i o s p o r e g e r m i n a t i o n was l o w e r f r o m e x p o s e d t h a n p r o t e c t e d a e c i a , a n d wet  spores germinated p o o r l y .  from associated fungi. posed t o temperatures favourable.  V i a b i l i t y was r e d u c e d b y c o n t a m i n a t i o n  A e c i o s p o r e s l o s t v i a b i l i t y v e r y r a p i d l y when e x a b o v e 25°C; t e m p e r a t u r e s  High humidity affected v i a b i l i t y ,  were a l s o adverse.  Direct  retention.  but u l t r a dry conditions  sunlight reduced v i a b i l i t y  d a i l y c o n d i t i o n s f a v o u r i n g d i s p e r s a l were l e a s t and v i a b i l i t y  c l o s e t o 0°C w e r e m o s t  rapidly.  Generally,  favourable f o rgermination  iv  TABLE OF CONTENTS  Page ABSTRACT  i  TABLE OF CONTENTS  i v  L I S T OF TABLES  i x  L I S T OF F I G U R E S  xiii  ACKNOWLEDGMENTS  xxiv  INTRODUCTION  1  THE ORGANISM  5 5  HISTORY OF THE FUNGUS  10  D I S T R I B U T I O N AND HOSTS Distribution of the rust  on P i n u s  11  Distribution o f the rust  on Santalaceae  Ik  L I F E CYCLE  2k  SYMPTOMS  37  DAMAGE  kl  1+8  AREA OF STUDY LOCATION, PHYSIOGRAPHY  AND GEOLOGY  48  SOILS  50  CLIMATE  50  VEGETATION  6l  STUDY LOCATIONS  62  V  65  AECIOSPORE PRODUCTION PERIODS OF AECIOSPORE PRODUCTION AND ENVIRONMEfflTAL FACTORS  65  A F F E C T I N G SPORULATION Methods  65  Results  66  Discussion  75  B I O L O G I C A L FACTORS A F F E C T I N G AECIOSPORE PRODUCTION  78  Methods  78  Results  82  Microflora  82  Microfauna  91  Rodents  102  Response o f t h e t r e e  107 109  Discussion  128  AECIOSPORE D I S P E R S A L  128  METEOROLOGICAL FACTORS A F F E C T I N G D I S P E R S A L Methods and M a t e r i a l s  128  Experimental sites  128  Spore c o l l e c t o r s  131  Meteorological  136  instruments  139  Results Diurnal p e r i o d i c i t y of meteorological and  the microclimate  factors,  o f study l o c a t i o n 1  D i u r n a l spore p e r i o d i c i t y  139 l 4 l  E f f e c t o f r e l a t i v e h u m i d i t y and temperature  146  Effect  150  of rainfall  vi  15I4.  E f f e c t o f dew Effect  155  o f wind  Availability  o f spores  Seasonal spore p e r i o d i c i t y AECIOSPORE D I S P E R S A L FROM A NATURAL POINT SOURCE  155 156 159  Methods and M a t e r i a l  159  Results  162  OTHER DATA ON DISTANCE OF AECIOSPORE D I S P E R S A L  171  AECIOSPORE D I S P E R S A L EXPERIMENTS FROM POINT SOURCES  173  Materials  and Methods  173  Experimental  arrangement  175  Experimental  procedure  179  Analysis of data  18U  Results Concentration  patterns  Change o f c o n c e n t r a t i o n w i t h d i s t a n c e RATE OF F A L L OF AECIOSPORES I N CALM A I R Materials  l8l  and Methods  Results AECIOSPORE D I S P E R S A L D I S C U S S I O N  l8k l8h 197 197 200 205  M e t e o r o l o g i c a l f a c t o r s and spore p e r i o d i c i t y  205  Spore d i s p e r s a l  213  AECIOSPORE GERMINATION  228  FACTORS A F F E C T I N G AECIOSPORE GERMINATION Methods and M a t e r i a l s  230 230  vii  Spore m a t e r i a l and methods o f h a n d l i n g  230  Methods f o r e x p l o r a t o r y experiments  231  Methods f o r subsequent experiments Results  . 23^ 2k0  E f f e c t o f temperature  on g e r m i n a t i o n  2k0  E f f e c t o f h u m i d i t y on g e r m i n a t i o n  2k7  S w e l l i n g o f spores on ... l i q u i d media  2^9  E f f e c t o f h y d r a t i o n o f spores on g e r m i n a t i o n  251  Effect of light  252  on g e r m i n a t i o n  E f f e c t o f hydrogen i o n c o n c e n t r a t i o n on g e r m i n a t i o n  255  E f f e c t o f s u b s t r a t e on g e r m i n a t i o n  257  a.  V a r i o u s agar media  257  b.  A d d i t i o n o f s u c r o s e t o media  259  c.  Presence o f h o s t l e a v e s on, o r l e a f e x t r a c t s i n t h e media  2.6o 262  Discussion DAILY AECIOSPORE GERMINATION  270  Methods and M a t e r i a l s  270  R e s u l t s and D i s c u s s i o n  272  283  AECIOSPORE VIABILITY Methods and M a t e r i a l  283  Results  287  E f f e c t o f temperature  •  on spore v i a b i l i t y  287  E f f e c t o f h u m i d i t y on spore v i a b i l i t y  292  Effect of light  29U  on spore v i a b i l i t y  viii  Discussion  297  SUMMARY AM) CONCLUSION  302  LITERATURE CITED  321  APPENDIX I  353  APPENDIX I I  356  ix  LIST OF TABLES  TABLE  Page M o n t h l y and a n n u a l c l i m a t i c summaries f o r K a n a n a s k i s , l a t . 51°02'W, l o n g . 115°03'W e l e v . k,560 f t MSL, f o r t h e p e r i o d o f r e c o r d (1939-1968).  53  II  M o n t h l y and a n n u a l c l i m a t i c summaries f o r s e l e c t e d s t a t i o n s i n t h e Marmot Creek R e s e a r c h B a s i n , l a t . 50°57'N, l o n g . 115° 1 0 % e l e v . 5,300-^8,000 f t MSL (1962-1967).  54  III  M o n t h l y t e m p e r a t u r e and p r e c i p i t a t i o n averages a t Kana n a s k i s Boundary Ranger S t a t i o n , l a t . 50°55'N, l o n g . 115°08'W, e l e v . 4,800 f t MSL (1962-1968)., and P i g e o n M o u n t a i n Lookout, l a t . 51°03'N, l o n g . 1 1 5 ° 0 4 % e l e v . 6,000 f t MSL (1960-1968), compared w i t h t h o s e a t K a n a n a s k i s f o r a s i m i l a r p e r i o d (1963J-I968).  55  IV  Date o f b e g i n n i n g and end o f C r o n a r t i u m comandrae a e c i o s p o r e p r o d u c t i o n , and main s p o r u l a t i o n p e r i o d a t a number o f l o c a t i o n s i n t h e y e a r s 1964 - 1968' inclusive.  67  V  Date o f b e g i n n i n g and end o f spore p r o d u c t i o n , and t o t a l number o f days o f p r o d u c t i o n from i n d i v i d u a l cankers a t l o c a t i o n 1 f r o m 1965 t o 1968 i n c l u s i v e . M a i n spore p r o d u c t i o n p e r i o d i s i n c l u d e d f o r 3 t o 6 cankers f o r t h e y e a r s 1965 t o I967.  68  VI  Average number o f days and range o f days o f a e c i o spore p r o d u c t i o n f o r i n d i v i d u a l p u s t u l e s on s e v e r a l cankers i n 1966 and I967 a t l o c a t i o n 1.  74  VII  Summary o f sequence o f s p o r u l a t i o n f o r 30 and 20 p u s t u l e s on two cankers a t l o c a t i o n 1 i n 1967.  74  VIII  The r e c o r d e d i n c i d e n c e o f T u b e r c u l i n a maxima on C r o n a r t i u m comandrae cankers on l o d g e p o l e p i n e and the t o t a l number o f a c t i v e and i n a c t i v e cankers obs e r v e d a t v a r i o u s l o c a t i o n s i n southwest A l b e r t a d u r i n g t h e y e a r s 1964 t o 1968.  83  IX  The i n c i d e n c e o f T u b e r c u l i n a maxima on o b s e r v e d a c t i v e and i n a c t i v e C r o n a r t i u m comandrae cankers a t 7 l o c a t i o n s d u r i n g t h e y e a r s 1964 t o 1968.  87  I  ;  :  X  X  The number o f C r o n a r t i u m comandrae a e c i a l cankers ( l ) p r o d u c i n g a e c i o s p o r e s , (2) s p o r u l a t i n g b u t i n f e c t e d w i t h T u b e r c u l i n a , (3) w i t h i n a c t i v e a e c i a l zones i n f e c t e d w i t h T u b e r c u l i n a , (k) w i t h i n a c t i v e o r dead c a n k e r s , a t two l o c a t i o n s d u r i n g t h e y e a r s 1966 t o 1968.  XI  The r e c o r d e d i n c i d e n c e o f C l a d o s p o r i u m t a x . sp. 1 on C r o n a r t i u m comandrae cankers observed a t v a r i o u s l o c a t i o n s i n southwest A l b e r t a d u r i n g t h e y e a r s I965 t o 1968.  XII  The i n c i d e n c e o f i n s e c t damage on C r o n a r t i u m comandrae cankers on l o d g e p o l e p i n e and t h e numb e r o f cankers observed a t v a r i o u s l o c a t i o n s i n southwest A l b e r t a d u r i n g t h e y e a r s 1964 t o 1968.  XIII  The i n c i d e n c e o f new r o d e n t damage on C r o n a r t i u m comandrae c a n k e r s , and t h e number o f cankers observed a t v a r i o u s l o c a t i o n s i n southwest A l b e r t a d u r i n g t h e y e a r s 1966 t o I968.  XIV  The p e r c e n t a g e i n c i d e n c e o f f r e s h r o d e n t chewing of C r o n a r t i u m comandrae cankers a t s e l e c t e d l o c a t i o n s d u r i n g t h e y e a r s 1966 t o 1968.  106  XV  The i n c i d e n c e o f f r e s h r e s i n o s i s on C r o n a r t i u m comandrae cankers on l o d g e p o l e p i n e and t h e number o f cankers observed a t a number o f l o c a t i o n s i n southwest A l b e r t a d u r i n g t h e y e a r s 1966 to 1968.  109  XVI  The h o u r l y mean temperature, r e l a t i v e h u m i d i t y and wind speed a t two canker s t a t i o n s , compared w i t h r e c o r d s from t h e Bay s t a t i o n f o r temperature and h u m i d i t y , and from K a n a n a s k i s f o r wind speed, f o r the p e r i o d May 17 t o J u l y 11, 1966.  iko  XVII  Number o f a e c i o s p o r e s d e p o s i t e d on n a t u r a l l e a f surfaces c o l l e c t e d along the four c a r d i n a l r a d i i at v a r i o u s d i s t a n c e s from a s p o r u l a t i n g C r o n a r t i u m comandrae canker on J u l y 7, 1968, a t l o c a t i o n 3-  170  XVIII  Average number o f a e c i o s p o r e s d e p o s i t e d p e r square c e n t i m e t e r on spore c o l l e c t o r coated s l i d e s a t d i s t a n c e s o f 5, 10 and 15 f e e t a l o n g e i g h t r a d i i around a s p o r u l a t i n g C r o n a r t i u m comandrae canker a t l o c a t i o n 3 on 9 days between June 2k and J u l y 15, 1968.  170  3  92  10k  xi  XIX  P a r t i c u l a r s o f experiments on d i s p e r s i o n o f Crona r t i u m comandrae a e c i o s p o r e s from a p o i n t s o u r c e .  l80  XX  T o t a l number o f spores t r a p p e d a t one f o o t above ground a t each d i s t a n c e on a l l r a d i i .  185  XXI  T o t a l number o f spores t r a p p e d a t 5 and 10 f e e t above ground a t 20, 50, 100 and 150 f e e t o n f i v e r a d i i .  185  XXII  R e s u l t s o f E x p e r i m e n t X, showing number o f spores t r a p p e d on an a r e a o f 13-5 s q cm a t t h e one f o o t l e v e l a t each s a m p l i n g p o i n t .  190  XXIII  Comparison o f t h e observed and e x p e c t e d c o n c e n t r a t i o n o f spores a t f o u r d i s t a n c e s from t h e 5 f o o t release point during eight tests.  195  XXIV  Percentages o f i n d i v i d u a l t e s t s o f aeciospores of C r o n a r t i u m comandrae d e p o s i t e d 3 meters from p o i n t o f l i b e r a t i o n on g l a s s s l i d e s exposed succ e s s i v e l y f o r 15 second p e r i o d s i n a c l o s e d c y l i n der.  201  XXV  Average r a t e o f f a l l i n s t i l l a i r o f t e n a e c i o spore t e s t s .  202  XXVI  Number o f a e c i o s p o r e clumps d e p o s i t e d 3 meters from p o i n t o f l i b e r a t i o n on g l a s s s l i d e s exposed s u c c e s s i v e l y f o r 15 second p e r i o d s i n a c l o s e d c y l i n d e r .  203  XXVII  Average r a t e o f f a l l i n s t i l l a i r o f d r y and wet a e c i o s p o r e s d u r i n g two t e s t s .  20k  XXYIII  Average l e n g t h and w i d t h (|a) o f a e c i o s p o r e s d e p o s i t e d on s l i d e s d u r i n g t h e d r y and wetspore t e s t s o f r a t e o f f a l l .  205  XXIX  Average number o f germ tubes p e r spore, and t h e range o f t h e average f o r f o u r t e s t s e r i e s a t v a r i o u s temperatures.  2^3  XXX  Average l e n g t h and w i d t h ( i n m i c r o n s ) o f 25 a e c i o spores from 5 f r e s h o r s t o r e d samples. Measurements were made on d r y spores and when spores were p l a c e d on a Czapek-Dox agar medium.  250  XXXI  Average g e r m i n a t i o n p e r c e n t a g e s f o r spores s t o r e d f o r v a r i o u s p e r i o d s o f time a t -k°C, and t h e n germinated, d r y o r a f t e r 2k hours hyd r a t i o n i n a s a t u r a t e d h u m i d i t y , on Czapek-Dox agar a t 15°'C.  252  Xll  E f f e c t o f l i g h t and dark c o n d i t i o n s on p e r c e n t g e r m i n a t i o n and germ tube growth o f t h r e e s e r i e s o f a e c i o s p o r e samples, germinated on Czapek-Dox agar a t 15-C o r under f l u c t u a t i n g o u t s i d e temp e r a t u r e s f o r 2k h o u r s . E f f e c t o f d a r k and t h r e e c o l o r e d l i g h t wave bands on p e r c e n t g e r m i n a t i o n and germ tube growth o f t h r e e s e r i e s o f a e c i o s p o r e samples, germinated a t 15°C on Czapek-Dox agar f o r 2k hours. Average p e r c e n t a g e and range o f g e r m i n a t i o n o f two s e r i e s o f a e c i o s p o r e s a f t e r 2k hours on d i f f e r e n t media a t 15°C. Average number o f germ tubes and l e n g t h s o f germ tubes p e r spore f o r t h r e e samples on seven d i f f e r e n t media a f t e r 2k hours a t 15°C. Average g e r m i n a t i o n p e r c e n t a g e s o f f i v e a e c i o spore samples s t o r e d a t s i x temperatures and germinated on water agar a t 15°C a f t e r v a r i o u s i n t e r v a l s o f storage. Average g e r m i n a t i o n p e r c e n t a g e s o f a e c i o s p o r e samples s t o r e d f o r f o u r t o s i x days a t v a r i o u s temperatures and germinated on water agar a t 15°C Average p e r c e n t a g e g e r m i n a t i o n o f a number o f spore samples g i v e n two methods o f r a p i d c o o l i n g and s t o r e d a t -20°C f o r p e r i o d s between 7 and 18 days, and t h e n germinated on water agar a t 15°C f o r 2k hours. A p o r t i o n o f the l i q u i d n i t r o g e n c o o l e d spores was r a p i d l y thawed f o r one minute a t 38°C p r i o r t o s e e d i n g on the agar. Average number o f germ tubes p e r spore and average l e n g t h o f l o n g e s t germ tube i n t h r e e dry-humid treatments a f t e r v a r i o u s i n t e r v a l s o f storage.  xiii  LIST OF FIGURES  Figure  Page  1  D i s t r i b u t i o n o f C r o n a r t i u m comandrae on P i n u s s p e c i e s i n w e s t e r n Canada.  15  2  D i s t r i b u t i o n o f C r o n a r t i u m comandrae on P i n u s s p e c i e s i n e a s t e r n Canada.  16  3  A e r i a l stems o f Geocaulon l i v i d u m , w i t h a x i l l a r y greenish flowers.  17  h  A e r i a l stems o f Comandra u m b e l l a t a ssp. p a l l i d a , w i t h t e r m i n a l w h i t e f l o w e r s about t o b r e a k open.  17  5.  S e r i e s o f a e r i a l stems o f Comandra u m b e l l a t a s s p . p a l l i d a , w h i c h branch j u s t below the s o i l , and are a t t a c h e d t o an underground rhizome. Some a e r i a l stems dead from p r e vious year.  17  6  Large h a u s t o r i u m o f Comandra u m b e l l a t a s s p . p a l l i d a a t t a c h e d t o Populus t r e m u l o i d e s Michx. r o o t , w i t h s m a l l e r h a u s t o r i u m on s m a l l e r r o o t .  17  7  D i s t r i b u t i o n o f C r o n a r t i u m comandrae on the S a n t a l a c e a e , Comandra u m b e l l a t a and Geocaulon ' l i v i d u m , i n w e s t e r n Canada.  22  8  D i s t r i b u t i o n o f C r o n a r t i u m comandrae on t h e S a n t a l a c e a e , Comandra u m b e l l a t a and Geocaulon l i v i d u m , i n e a s t e r n Canada.  23  9  T r a n s v e r s e s e c t i o n s t a k e n from the c e n t r a l p o r t i o n o f f o u r C r o n a r t i u m comandrae cankers on l o d g e p o l e p i n e showing e c c e n t r i c growth caused by the r u s t . Note t h e r e s i n impregn a t i o n o f the o u t e r sapwood.  26  10  T r a n s v e r s e s e c t i o n s t a k e n from the c e n t r a l p o r t i o n o f f i v e C r o n a r t i u m comandrae cankers on l o d g e p o l e p i n e showing e c c e n t r i c growth caused by the r u s t .  26  xiv  Transverse s e c t i o n taken through the a e c i a l zone o f a C r o n a r t i u m comandrae canker showing t h e a e c i a s i t u a t e d i n t h e b a r k t i s s u e s . Note t h e r e s i n i m p r e g n a t i o n o f t h e b a r k between a e c i a , and the s w e l l i n g o f t h e o u t e r r i n g - o f sapwood caused by t h e fungus. P y c n i a l drops on the p y c n i a l zone o f a C r o n a r t i u m comandrae canker on the stem and branches o f a l o d g e p o l e p i n e . Note t h e s w e l l i n g o f the stem a s s o c i a t e d w i t h the canker. Group o f p y r i f o r m p y c n i o s p o r e s o f C r o n a r t i u m comandrae. X 2500. P y r i f o r m a e c i o s p o r e s o f C r o n a r t i u m comandrae showing s i z e and shape v a r i a t i o n o f t h e spore t a i l . X 700. P y r i f o r m a e c i o s p o r e s o f C r o n a r t i u m comandrae showing the spore w a l l o r n a m e n t a t i o n . X 700. Canker.of C r o n a r t i u m comandrae on a young b r a n c h o f P i n u s c o n t o r t a , w i t h the p e r i d i a o f the a e c i a s t i l l u n r u p t u r e d and f o r m i n g t y p i c a l b l i s t e r s p u s h i n g t h r o u g h the b a r k . Canker o f C r o n a r t i u m comandrae w i t h abundant r u p t u r e d a e c i a c o v e r i n g the a e c i a l zone. B r i g h t orange-colored aeciospores of Cronartium comandrae c o v e r i n g the exposed a e c i a o f t h e canker. Young canker o f C r o n a r t i u m comandrae on P i n u s c o n t o r t a w i t h rough b a r k c o v e r i n g t h e o l d a e c i a l zone around the b r a n c h stub t h a t a c t e d as e n t r y p o i n t t o the stem. Abundant a e c i a were produced around t h e o l d a e c i a l zone. S l i g h t h y p e r t r o p h y o f t h e young P i n u s c o n t o r t a stem caused by C r o n a r t i u m comandrae, w i t h c h a r a c t e r i s t i c rough, c r a c k e d b a r k , i n t h e o l d e r canker a r e a caused by a e c i a r u p t u r i n g . T y p i c a l b a s a l stem canker o f C r o n a r t i u m comandrae showing o l d e r rough b a r k zone and c u r r e n t y e a r a e c i a l zone w i t h many d i s p e r s e d a e c i o s p o r e s caught i n the c r e v i c e s o f the b a r k .  XV  22  A e c i o s p o r e s o f C r o n a r t i u m comandrae, s t a i n e d ' w i t h HC1 - Giemsa, each w i t h two n u c l e i . X 700.  33  23  A e c i o s p o r e o f C r o n a r t i u m comandrae, s t a i n e d w i t h HC1 - Giemsa, w i t h two n u c l e i . X 1200,  33  2k  I n i t i a t i o n of aeciospore germination, with four germ tubes d e v e l o p i n g t h r o u g h the spore w a l l , a f t e r 1 hour. X 1200.  33  25  C r o n a r t i u m comandrae a e c i o s p o r e w i t h s e v e r a l germ t u b e s , but o n l y one germ tube w e l l developed a f t e r 6 h o u r s . Note the shortm u l t i p l e branches on the n o n - s e p t a t e germ tube. X 380.  33  26  Group o f germinated a e c i o s p o r e s w i t h t y p i c a l b r a n c h i n g o f germ tubes a f t e r 2k h o u r s . X 125.  33  27  A e c i o s p o r e o f C r o n a r t i u m comandrae w i t h one developed germ tube, a f t e r 3 h o u r s . Note t h e n o n - s e p t a t e c o n d i t i o n o f the germ tube and t h a t the two n u c l e i have m i g r a t e d towards t h e t i p o f the germ t u b e . X 830.  3^  28  Germ tubes o f two C r o n a r t i u m comandrae a e c i o - spores a f t e r 3 hours, showing development o f a g l o b o s e and club-shaped a p p r e s s o r i u m on t h e developed germ tube, i n t o w h i c h the two n u c l e i have m i g r a t e d . X 700.  3^+  29  Germ tube development o f a C r o n a r t i u m comandrae a e c i o s p o r e a f t e r 3 hours, showing a g l o b o s e app r e s s o r i u m w i t h two n u c l e i and an i n f e c t i o n peg. x 1080.  3^+  30  Germ tube development o f two C r o n a r t i u m comandrae a e c i o s p o r e s a f t e r 3 hours, showing two t y p e s o f a p p r e s s o r i a each w i t h two n u c l e i and an i n f e c t i o n peg. X 6l0.  3^  31  U r e d i a o f C r o n a r t i u m comandrae on c e n t r a l p o r t i o n o f a Comandra u m b e l l a t a s s p . p a l l i d a l e a f . Dark s t r u c t u r e s on bottom are young t e l i a .  35  32  T e l i a o f C r o n a r t i u m comandrae on b o t h s u r f a c e s o f a p o r t i o n - o f a Comandra u m b e l l a t a ssp. pallida leaf.  35  xvi  Branch i n f e c t i o n o f C r o n a r t i u m comandrae on P i n u s c o n t o r t a , w i t h mycelium c a u s i n g s w e l l i n g i n the main stem around the b r a n c h s t u b . Tree w i t h dead s p i k e - t o p above a canker o f C r o n a r t i u m comandrae g i r d l i n g t h e t r e e ; lower branches a r e p r o g r e s s i v e l y k i l l e d by downward growth o f r u s t . Heavy r e s i n o s i s f o r m i n g a d r i e d c r u s t over a p o r t i o n o f a C r o n a r t i u m comandrae stem canker. Note rough b a r k o f o l d b r a n c h canker (upper l e f t ) w h i c h s e r v e d as e n t r y f o r the r u s t t o t h e stem. A n n u a l r o d e n t damage on a l a r g e C r o n a r t i u m comandrae stem canker on P i n u s c o n t o r t a . Note the s t r i p o f d r i e d dead b a r k not removed on each a n n u a l v i s i t , and the abundant e x u d a t i o n of r e s i n . Rodentcdamage on a C r o n a r t i u m comandrae stem canker around a b r a n c h w h i c h a c t e d as an e n t r y p o i n t f o r t h e r u s t i n t o t h e stem. Sporul a t i o n o f t h e a e c i a l zone can be seen o u t s i d e the chewed a r e a . Map o f the lower K a n a n a s k i s R i v e r V a l l e y , A l b e r t a , showing the l o c a t i o n s o f the main study areas. Study l o c a t i o n 1, on t h e K a n a n a s k i s F o r e s t Experiment S t a t i o n , near t h e n o r t h e a s t shore o f B a r r i e r Lake, showing the l o c a t i o n o f i n f e c t e d l o d g e p o l e p i n e , Comandra p l a n t p l o t s , and i n struments used d u r i n g the s t u d y . Summary o f t h e d a i l y maximum and minimum t e m p e r a t u r e s and r e l a t i v e h u m i d i t y t a k e n from hygrothermograph r e c o r d s , t h e d a i l y r a i n f a l l , and the phenology o f the a e c i a l , p y c n i a l , u r e d i a l and t e l i a l s t a t e s o f C r o n a r t i u m comandrae, May t o August i n I965 and 1966. Summary o f t h e d a i l y maximum and minimum temp e r a t u r e s and r e l a t i v e h u m i d i t y t a k e n from hygrothermograph r e c o r d s , t h e d a i l y r a i n f a l l , and the phenology o f the a e c i a l , p y c n i a l , u r e d i a l and t e l i a l s t a t e s o f C r o n a r t i u m comandrae, May to August i n I967 and 1968.  XV11  A p l a s t i c s c r e e n i n g c y l i n d r i c a l - s l e e v e cage used f o r c o l l e c t i n g i n s e c t s from C r o n a r t i u m comandrae cankers on stems o f P i n u s c o n t o r t a . Rough, c r a c k e d b a r k o f the a e c i a l zone o f a C r o n a r t i u m comandrae canker i n f e c t e d w i t h the p u r p l e mold, T u b e r c u l i n a maxima, w h i c h i s conspicuous as a d a r k e r a r e a where the s u r f a c e b a r k has been removed, o r c r a c k e d . B a s a l canker o f C r o n a r t i u m comandrae w i t h t y p i c a l rough b a r k i n a e c i a l zone and showing e v i d e n c e o f i n s e c t damage. Note e x i t h o l e s and L e p i d o p t e r a f r a s s a t top o f canker, and f u r t h e r f r a s s i n lower rough zone. P u p a l chambers o f P i s s o d e s s c h w a r z i s c o r e d i n t o the sapwood throughout the C r o n a r t i u m comandrae canker a r e a on s m a l l P i n u s c o n t o r t a stem. View o f s i t e no. 2 at s t u d y l o c a t i o n 1, showing a 24-hour i m p a c t i o n spore c o l l e c t o r by C r o n a r t i u m comandrae canker no. 2721 on a s m a l l P i n u s c o n t o r t a , and i n s t r u m e n t s f o r r e c o r d i n g the weather. I n s t r u m e n t s a r e , from l e f t t o r i g h t , b l a c k porous d i s c atmometer, i n s t r u m e n t s h e l t e r c o n t a i n i n g hygrothermograph, Wallin-Polhemus dew d u r a t i o n r e c o r d e r on ground, mast w i t h anemometer cups f o r wind speed r e c o r d e r and spore c o l l e c t o r . ( R a i n gauge i s out o f t h e picture) S t a n d a r d Stevenson s c r e e n i n an opening, cont a i n i n g hygrothermograph and thermometers used as r e f e r e n c e weather s t a t i o n a t s t u d y a r e a 1, with a bi-metal actinograph f o r recording i n coming r a d i a t i o n seen a t t h e back. A r e c o r d i n g r a i n gauge and a wind d i r e c t i o n r e c o r d e r (out o f the p i c t u r e ) were m a i n t a i n e d a t t h i s open s i t e . A H i r s t spore t r a p w i t h s a m p l i n g o r i f i c e one f o o t above ground, c l o s e t o a Comandra umbellata plot.  xviii 49  Seven-day p o l l e n sampler o f t h e Sarvas t y p e , used t o c o l l e c t a e c i o s p o r e s a t s e t d i s t a n c e s from s p o r u l a t i n g C r o n a r t i u m comandrae c a n k e r s . Spores pass t h r o u g h the sampler o r i f i c e and are d e p o s i t e d on a v a s e l i n e - c o a t e d ' s a m p l i n g band p l a c e d around a c l o c k - d r i v e n drum housed w i t h i n the i n t a k e c y l i n d e r .  130  50  A 24-hour i m p a c t i o n spore c o l l e c t o r , w i t h the s e a l e d l i d removed t o show t h e a c r y l i c p l a s t i c d i s c h o l d i n g 24 microscope s l i d e s upon w h i c h spores are d e p o s i t e d , and the f a n f o r drawing a i r t h r o u g h t h e box at a c o n t r o l l e d r a t e .  133  51  A 24-hour i m p a c t i o n spore c o l l e c t o r i n operat i o n , w i t h i t s sampling o r i f i c e p l a c e d c l o s e t o the s p o r u l a t i n g s u r f a c e o f a C r o n a r t i u m comandrae canker.  133  52  C e n t r a l i n s t r u m e n t power and r e c o r d i n g box, w i t h T h o r n t b m t e f o u r - u n i t wind speed r e g i s t e r r e c o r d e r ( l e f t ) , wind d i r e c t i o n r e c o r d e r , and d i g i t a l p r i n t o u t r e c o r d e r w i t h p o l a r o i d camera f o r wind speed r e c o r d e r system ( r i g h t ) . Behind are the t r a n s f o r m e r s and r e c t i f i e r s f o r r e d u c i n g t h e 110 v o l t power s u p p l y and c o n t r o l l i n g an o u t p u t o f 12 v o l t s f o r o p e r a t i o n o f wind i n struments, dew i n s t r u m e n t s and 24-hour i m p a c t i o n spore c o l l e c t o r s . A standby 12 v o l t b a t t e r y i s also present.  133  53  S e t o f anemometer cups o f the T h o r n t h w a i t e wind speed r e g i s t e r r e c o r d e r system o p e r a t e d at canker h e i g h t near a s p o r u l a t i n g C r o n a r t i u m comandrae canker.  133  54  Mean d i u r n a l a e c i o s p o r e p e r i o d i c i t y curves f o r t h r e e cankers over v a r y i n g p e r i o d s o f t r a p p i n g i n the y e a r s 1964 t o I967, e x p r e s s e d as a p e r c e n t a g e o f the peak g e o m e t r i c mean hourly concentration.  143  55  Frequency o f h o u r l y maximum a e c i o s p o r e concent r a t i o n s from t h r e e C r o n a r t i u m comandrae cank e r s on 91 d r y and 40 r a i n y days i n 1966 and 95 d r y and 29 r a i n y days i n 1967, d u r i n g the main spore p r o d u c t i o n p e r i o d s .  l45  56  Average h o u r l y a e c i o s p o r e c o n c e n t r a t i o n s c o l l e c t e d from t h r e e C r o n a r t i u m comandrae cankers i n June 1966.  147  xix  Average h o u r l y a e c i o s p o r e c o n c e n t r a t i o n s c o l l e c t e d from t h r e e C r o n a r t i u m comandrae cankers from June 6 t o 25, I967. Average h o u r l y a e c i o s p o r e c o n c e n t r a t i o n c o l l e c t e d from C r o n a r t i u m comandrae canker no. 2713 d u r i n g p e r i o d s w i t h o u t r a i n (average 19 t o 2k days) i n June 1966, compared w i t h t h e a v e r age h o u r l y a i r temperature, r e l a t i v e h u m i d i t y and wind speed. H o u r l y number o f a e c i o s p o r e s c o l l e c t e d from C r o n a r t i u m comandrae canker no. 25l6 d u r i n g t h e p e r i o d June 7 t o J u l y 3, I965, r e l a t e d t o h o u r l y a i r temperature, r e l a t i v e h u m i d i t y , r a i n f a l l , incoming r a d i a t i o n and wind speed. H o u r l y number o f a e c i o s p o r e s c o l l e c t e d from C r o n a r t i u m comandrae canker no. 2721 d u r i n g t h e p e r i o d May 29 t o June 9 1966, r e l a t e d t o a i r temperature, r e l a t i v e h u m i d i t y , r a i n f a l l , incoming r a d i a t i o n and wind speed. 5  D a i l y number o f a e c i o s p o r e s c o l l e c t e d from two C r o n a r t i u m comandrae cankers d u r i n g t h e 1966 spore p r o d u c t i o n p e r i o d , p l o t t e d on semi-log s c a l e . D a i l y number o f a e c i o s p o r e s c o l l e c t e d from C r o n a r t i u m comandrae canker no. 2713 d u r i n g I965, I966 and 1967, p l o t t e d on s e m i - l o g scale. P a t t e r n o f a e c i o s p o r e d e p o s i t i o n around C r o n a r t i u m comandrae canker no. 2713 d u r i n g a ICH5- hour p e r i o d on June 23, 1967The average percentage change o f a e c i o s p o r e d e p o s i t i o n c o n c e n t r a t i o n w i t h d i s t a n c e on e i g h t r a d i i around C r o n a r t i u m comandrae canker no. 2713 on June 23, 1967, and t h e percentage change on t h e s o u t h e a s t r a d i i , p l o t t e d on s e m i - l o g s c a l e . P a t t e r n o f a e c i o s p o r e d e p o s i t i o n around C r o n a r t i u m comandrae canker no. 2689, on t h r e e days (0800 t o IcOO'J-hours) i n June 1968.  XX  The average p e r c e n t a g e change o f a e c i o spore d e p o s i t i o n w i t h d i s t a n c e around C r o n a r t i u m comandrae canker no. 2689, on t h r e e days (0800 t o l600 h o u r s ) i n June 1968, p l o t t e d on s e m i - l o g s c a l e . S p o r e - e j e c t o r used f o r l i b e r a t i n g a e c i o spores i n d i s p e r s a l experiments from p o i n t source. Compressor u n i t v w i t h c o n n e c t i n g r u b b e r tubing, f o r regulating a i r flow f o r r e lease o f aeciospores i n d i s p e r s a l experiments. Wind-directional p l a s t i c s l i d e holder spore c o l l e c t o r s ( w i t h o u t m i c r o s c o p e s l i d e s ) used f o r c o l l e c t i n g a e c i o s p o r e s i n d i s p e r s a l experiments and from n a t u r a l sources. P o r t i o n o f spore c o l l e c t o r network, w i t h c o l l e c t o r s on d i f f e r e n t r a d i i a t v a r i o u s d i s t a n c e s from r e l e a s e p o i n t . Wind d i r e c t i o n vane and wind speed anemometer can be seen t o t h e l e f t o f t h e s t a n d f o r the spore-ejector. Note spore c o l l e c t o r s at 5 and 10 f e e t on t h e masts. P o r t i o n o f spore c o l l e c t o r network w i t h c o l l e c t o r s on d i f f e r e n t r a d i i a t v a r i o u s d i s t a n c e s from s p o r e - e j e c t o r a t r e l e a s e p o i n t . Wind speed anemometer a t l e f t of spore-ejector. Diagram o f p o i n t source a e c i o s p o r e d i s p e r s a l experiment sampling g r i d , w i t h 113 sampling c o l l e c t o r s l a i d out f o r southwest w i n d s . Spore c o l l e c t o r s were a l s o l o c a t e d a t t h e 300 and kOO f o o t d i s t a n c e s a l o n g t h e m i d - l i n e . Aeciospore concentration patterns a t three sampling h e i g h t s f o r a spore r e l e a s e one f o o t above ground (Experiment V I I ) . Aeciospore concentration patterns a t three sampling h e i g h t s f o r a spore r e l e a s e f i v e • f e e t above ground (Experiment X ) .  xx i  Composite a e c i o s p o r e c o n c e n t r a t i o n p a t t e r n s at three sampling heights f o r a l l e i g h t spore r e l e a s e s f i v e f e e t above ground (Experiments I I - V I , V I I I - X ) . V e r t i c a l p r o f i l e s o f a e c i o s p o r e concent r a t i o n p a t t e r n s a l o n g the m i d - l i n e o f the s a m p l i n g network f o r two spore r e l e a s e s a t one f o o t (Experiments I and V I I ) , and two spore r e l e a s e s f i v e f e e t above ground (Experiments V and X ) . V e r t i c a l p r o f i l e s o f a e c i o s p o r e concent r a t i o n p a t t e r n s a l o n g the m i d - l i n e and f o r k5° and 90° s e c t o r s o f t h e s a m p l i n g network, f o r one spore r e l e a s e a t f i v e f e e t above ground (Experiment V I I I ) . Composite v e r t i c a l p r o f i l e s o f a e c i o s p o r e c o n c e n t r a t i o n p a t t e r n s a l o n g the m i d - l i n e and f o r ^5° and 90° s e c t o r s o f the samp l i n g network, f o r a l l e i g h t spore r e l e a s e s f i v e f e e t above ground (Experiments I I VI, V I I I - X ) . E f f e c t o f temperature on p e r cent germinat i o n o f t h r e e C r o n a r t i u m comandrae a e c i o spore samples a f t e r 2k hours on water agar (pH 6.8). E f f e c t o f temperature on C r o n a r t i u m comandrae a e c i o s p o r e g e r m i n a t i o n . Average p e r c e n t g e r m i n a t i o n o f f i v e s e r i e s and v a r i o u s numbers o f r e p l i c a s w i t h t h r e e t o f i v e samples each, a f t e r 2k hours on water agar. Average l e n g t h of. germ tubes o f t h r e e C r o n a r t i u m comandrae a e c i o s p o r e samples germinated oh water agar (pH 6.8). a f t er 2k hours a t v a r i o u s t e m p e r a t u r e s . (a) Length o f l o n g e s t germ tube p e r s p o r e , (b) Length o f a l l germ tubes per s p o r e . I n f l u e n c e o f v a r i o u s temperatures on r a t e o f C r o n a r t i u m comandrae a e c i o s p o r e germinat i o n a f t e r v a r i o u s time i n t e r v a l s on water agar.  xxii  83  Average r a t e o f g e r m i n a t i o n o f t h r e e Conartium comandrae a e c i o s p o r e samples a t d i f f e r e n t temp e r a t u r e s and a f t e r v a r i o u s time i n t e r v a l s on water agar.  246  84  Average l e n g t h o f t h e l o n g e s t germ tube from t h r e e t o f i v e C r o n a r t i u m comandrae a e c i o s p o r e samples germinated on..water agar a t d i f f e r e n t temperatures and a f t e r v a r i o u s time i n t e r v a l s .  2 4 8  85  E f f e c t o f hydrogen i o n c o n c e n t r a t i o n on germi n a t i o n o f C r o n a r t i u m comandrae a e c i o s p o r e s a f t e r 24 hours a t 15 and 20°C.  256  86  E f f e c t o f hydrogen i o n c o n c e n t r a t i o n on g e r m i n a t i o n o f C r o n a r t i u m comandrae a e c i o spores a f t e r 2 4 hours a t 5, 15 and 25°C. Spore samples were s t o r e d f o r 4 and 6 weeks at 0°C p r i o r t o u s e .  256  87  Average d a i l y p e r cent g e r m i n a t i o n o f f i v e t o seven C r o n a r t i u m comandrae a e c i o s p o r e samples a f t e r 24 hours on water agar a t 15°C, f o r t h e y e a r s 1965, 1966 and 1967.  273  88  D a i l y p e r cent g e r m i n a t i o n o f a e c i o s p o r e s from two C r o n a r t i u m comandrae cankers f o r t h e y e a r s 1965, 1966 and 1967.  275  89  V a r i a t i o n i n t h e d a i l y p e r cent g e r m i n a t i o n o f a e c i o s p o r e s from seven C r o n a r t i u m comdrae cankers i n I965.  276  90  D a i l y p e r cent g e r m i n a t i o n o f a e c i o s p o r e s from f i v e i n d i v i d u a l a e c i a o f a C r o n a r t i u m comandrae canker d u r i n g t h e 1966 s p o r u l a t i o n period.  277  91  D a i l y p e r cent g e r m i n a t i o n o f a e c i o s p o r e s from f i v e i n d i v i d u a l a e c i a o f a C r o n a r t i u m comandrae canker d u r i n g t h e I967 s p o r u l a t i o n p e r i o d .  278  92  Average p e r cent g e r m i n a t i o n o f t h r e e C r o n a r t i u m comandrae a e c i o s p o r e samples s t o r e d a t t h r e e t e m p e r a t u r e s , and germinated on water agar a t 15°C a f t e r v a r i o u s hours o f s t o r a g e .  288  XX 111  Average p e r cent g e r m i n a t i o n o f f i v e C r o n a r t i u m comandrae a e c i o s p o r e samples s t o r e d i n a wet, d r y o r u l t r a d r y atmosphere, and germinated on water agar a t 15°C a f t e r v a r i o u s i n t e r v a l s o f s t o r a g e . E f f e c t o f exposure o f C r o n a r t i u m comandrae aeciospores t o d i r e c t l i g h t f o r varying l e n g t h s o f time on c l e a r sunny days (June 16, and J u l y 5, 1966; June 23, 1967), and on a c l o u d y day (June 16, 1967).  xxiv  ACKNOWLEDGMENTS  I w i s h t o express my s i n c e r e g r a t i t u d e t o t h e l a t e Dr. J . E. B i e r f o r h i s e n l i g h t e n e d guidance and communicative e n t h u s i a s m much o f t h i s i n v e s t i g a t i o n .  throughout-  I w i s h a l s o t o thank Dr. R. J . Bandoni, D r .  A. L. F a r l e y , Dr. P. G. Haddock, D r . T. M. C. T a y l o r , D r . G. H. N. Towers, and D r . B. J . v a n d e r Kamp, f o r t h e i r c o u n s e l i n g and a s s i s t a n c e i n t h e preparation o f the t h e s i s .  Thanks a r e extended t o t h e Canada Department  o f F i s h e r i e s and F o r e s t r y f o r p e r m i s s i o n t o use, f o r t h i s t h e s i s , gathered  i n t h e course o f work on a Department p r o j e c t .  expressed  data  Appreciation i s  t o Mr. W. Morf and Mr. L. S. S k a l e y , t e c h n i c i a n s o f t h e F o r e s t  Research Laboratory, Calgary, A l b e r t a , f o r assistance i n the c o l l e c t i o n o f data. Thanks a r e due t o D r . B. C. S u t t o n , F o r e s t R e s e a r c h L a b o r a t o r y , Canada Department o f F i s h e r i e s and F o r e s t r y , Winnipeg, M a n i t o b a , f o r t h e i d e n t i f i c a t i o n o f the m a j o r i t y o f the f u n g i .  Other s p e c i a l i s t s who con-  t r i b u t e d t o t h e i d e n t i f i c a t i o n o f v a r i o u s f l o r a l groups a r e :  D r . A. M.  Adams, H o r t i c u l t u r a l E x p e r i m e n t S t a t i o n , O n t a r i o Department o f A g r i c u l t u r e , V i n e l a n d S t a t i o n , O n t a r i o ; D r . F. D. Cook, Department o f S o i l S c i e n c e , U n i v e r s i t y o f A l b e r t a , Edmonton, A l b e r t a ; D r . Y. H i r a t s u k a , F o r e s t R e s e a r c h Laboratory,  Canada Department o f F i s h e r i e s & F o r e s t r y , C a l g a r y , A l b e r t a ;  and D r . D. B. P r e s t , Department o f B a c t e r i o l o g y , U n i v e r s i t y o f W i s c o n s i n , Madison, W i s c o n s i n .  Many i n s t i t u t i o n s p r o v i d e d C r o n a r t i u m  comandrae ma-  t e r i a l f o r t h e d i s t r i b u t i o n a l a s p e c t s o f t h i s study, and I would l i k e t o e s p e c i a l l y thank t h e c u r a t o r s o f t h e f o l l o w i n g h e r b a r i a f o r t h e i r tance:  assis-  A r t h u r Herbarium, Purdue U n i v e r s i t y , L a f a y e t t e , I n d i a n a ; The  xxv  N a t i o n a l Fungus C o l l e c t i o n s , B e l t s v i l l e , M a r y l a n d ; The New Y o r k Garden, New  York; Mac Donald C o l l e g e ,  S t e . A n n e - d e - B e l l e v u e , Quebec; The  W. P. F r a s e r M e m o r i a l Herbarium, U n i v e r s i t y o f Saskatchewan, Saskatchewan; U n i v e r s i t y o f A l b e r t a , Edmonton, A l b e r t a ; Herbarium, Canada Department Mycological  Herbaria,  Botanical  Saskatoon,  Mycological  o f A g r i c u l t u r e , Ottawa; O n t a r i o ;  Canada Department  and the  o f F i s h e r i e s and F o r e s t r y ,  at  C a l g a r y , Quebec, V i c t o r i a and Winnipeg. I am i n d e b t e d t o the f o l l o w i n g taxonomic s p e c i a l i s t s o f the Entomology R e s e a r c h I n s t i t u t e , Ontario,  Canada Department  of A g r i c u l t u r e ,  who made a u t h o r i t a t i v e i d e n t i f i c a t i o n s o f f a u n a l  Dr. E. C. B e c k e r , Dr. D. Brown, Mr. W.  Ottawa,  material:  J . Brown, Dr. J . M.  Campbell,  Dr. T. N. Freeman, Dr. M. Ivanochko, Mr. G. L e w i s , Dr. E. E.  Lindquist,  M i s s M. R. MacKay, Mr. J . E. H. M a r t i n , Dr. W. R. M. Mason, Dr. J . F. M c A l p i n e , Mr. C. D. F. M i l l e r , Dr. A. Mutuura, Dr. 0.  Peck, Dr. B. V.  P e t e r s o n , Dr. W. R. R i c h a r d s , Mr. R. de R u e t t e , Dr. L. K..Smith, Dr. J . R. V o c k e r o t h and Mr. G. S. W a l l e y . the i d e n t i f i c a t i o n o f v a r i o u s  Other s p e c i a l i s t s who  f a u n a l groups a r e :  F o r e s t R e s e a r c h L a b o r a t o r y , Canada Department Winnipeg, M a n i t o b a ; Dr. C. W.  contributed  Dr. G. A.  Bradley,  of Fisheries & Forestry,  O'Brien, Department  o f Entomology,  s i t y o f C a l i f o r n i a , B e r k e l e y , C a l i f o r n i a ; Dr. C. V. G. Morgan, L a b o r a t o r y , Canada Department  to  UniverEntomology  o f A g r i c u l t u r e , Summerland, B r i t i s h Colum-  b i a ; Dr. C. T. P a r s o n s , Manchester Depot, Vermont; Mr. B. A. Sugden, F o r e s t Entomology L a b o r a t o r y , Canada Department  of F i s h e r i e s &  Vernon, B r i t i s h Columbia; Dr. A. L. T u r n b u l l , Department  Forestry,  of B i o l o g i c a l  S c i e n c e , Simon F r a s e r U n i v e r s i t y , Burnaby, B r i t i s h Columbia; and Dr. A. W o o l l e y , Department  o f Zoology, Colorado State U n i v e r s i t y , F o r t  T.  Collins,  xxv i  Colorado. Thanks are extended t o my w i f e , M a r g a r e t , who  p r o o f - r e a d the  m a n u s c r i p t d u r i n g i t s p r e p a r a t i o n , and t o M i s s B a r b a r a Brown who the f i n a l m a n u s c r i p t .  typed  - 1 -  INTRODUCTION  A knowledge o f t h e e f f e c t o f environment that part of i t s l i f e  on a pathogen d u r i n g  c y c l e when i t i s o u t s i d e t h e h o s t i s o f prime im-  p o r t a n c e t o an u n d e r s t a n d i n g o f t h e f a c t o r s i n f l u e n c i n g t h e spread and i n t e n s i f i c a t i o n o f an a i r b o r n e d i s e a s e .  F o r a pathogen t o be a b l e t o  spread and i n t e n s i f y e f f e c t i v e d i s p e r s a l must be a c h i e v e d .  This implies  t h e p r o d u c t i o n o f spores i n p o s i t i o n s from w h i c h t h e y can be t r a n s p o r t e d and d e p o s i t e d a l i v e on h o s t p l a n t s under c o n d i t i o n s and i n q u a n t i t i e s that w i l l lead t o i n f e c t i o n .  The e n v i r o n m e n t a l c o n d i t i o n s under w h i c h  a pathogen can i n t e n s i f y have been f a i r l y a c c u r a t e l y d e f i n e d f o r a few d i s e a s e s , b u t e s t i m a t e s o f t h e d i s t a n c e o f spread o r l o s s t h r o u g h nonv i a b i l i t y under d i f f e r e n t e n v i r o n m e n t a l c o n d i t i o n s have been l a r g e l y guesses.  B e s i d e s a f f e c t i n g d i s p e r s a l , environment  also a f f e c t s the pro-  d u c t i o n o f i n o c u l u m on t h e h o s t , spore g e r m i n a t i o n and subsequent  infec-  t i o n o f t h e h o s t , a l l o f w h i c h s h o u l d t h e r e f o r e be c o n s i d e r e d i n t h i s phase o f d i s e a s e e p i d e m i o l o g y .  F a c t o r s o f t h e environment  are both l i -  m i t i n g and o p t i m a l f o r spore s u r v i v a l and g e r m i n a t i o n and must be known b e f o r e i n f e c t i o n can be p r e d i c t e d . The phase i n t h e l i f e pathogen,  when a spore s t a t e proceeds  cerns t h e f i e l d o f a e r o b i o l o g y .  cycle of a  i n d e p e n d e n t l y o f i t s h o s t s , con-  Gregory (1961) c o n s i d e r e d a e r o b i o l o g y  as "the s t u d y o f p a s s i v e l y airborne-.macro-organisms b e h a v i o u r , movements and s u r v i v a l " .  their  identity,  T h i s phase has been s t u d i e d i n a  few d i s e a s e s a f f e c t i n g a g r i c u l t u r a l crops ( C a r t e r 1963; H i r s t and Stedman 1961, 1962; J a r v i s 1962; M e r e d i t h 1961a, 1961b; Shanmuganathan and  - 2  Arulpragasam  -  1 9 6 6 ; TSireeramulu 1 9 6 2 ; W i l s o n and Baker 1 9 4 6 ) , b u t few-  s t u d i e s have i n v o l v e d a s p e c t s o f t h e a e r o b i o l o g y o f f o r e s t pathogens (Bega i 9 6 0 ;  R i s b e t h 1959;  i ^ r e e r a m u l u 1 9 6 3 ; Van A r s d e l e t a l . 1 9 6 1 ) .  The comandra b l i s t e r r u s t , C r o n a r t i u m comandrae Peck, an imp o r t a n t n a t i v e d i s e a s e damaging s e v e r a l s p e c i e s o f h a r d p i n e i n N o r t h America,  was s e l e c t e d f o r t h i s s t u d y , because o f i t s d i s t i n c t i v e a e c i o -  spore, a unique p y r i f o r m shape, w h i c h i s e a s i l y i d e n t i f i e d a g a i n s t t h e background a i r s p o r a .  Except  i t s history, distribution,  f o r an o u t l i n e o f t h e C_. comandrae r u s t  life  c y c l e , symptoms and damage, t h i s  —  study  i s l i m i t e d t o the a e r o b i o l o g i c a l aspects o f the a e c i a l s t a t e of the r u s t . The r u s t has been known on p i n e f o r n e a r l y 90 y e a r s and was f i r s t recorded i n western P i n u s c o n t o r t a Dougl.,  Canada from A l b e r t a i n 1907 on l o d g e p o l e p i n e ,  and from B r i t i s h Columbia i n 1913 on p o n d e r o s a  p i n e P. p o n d e r o s a Laws. ( A r t h u r and K e r n 1 9 1 4 ) .  Widespread damage by C_.  comandrae i n w e s t e r n N o r t h A m e r i c a was not r e c o r d e d u n t i l t h e m i d - 1 9 5 0 ' s ( M i e l k e 1957), b u t d u r i n g t h e l a s t decade t h e r e have been s e v e r a l r e p o r t s (Andrews and H a r r i s o n 1 9 5 9 ; K r e b i l l 1 9 6 5 ; P e t e r s o n 1 9 6 2 a ) ,  and i t -  has r e c e n t l y become a p l a n t a t i o n p r o b l e m i n o t h e r p a r t s o f N o r t h A m e r i c a ( C o r d e l l et a l . 1967;  Dance and Lynn 1 9 6 5 ; Wolfe e t a l . 1 9 6 8 ) .  The genus  C r o n a r t i u m i n c l u d e s t h e w o r l d ' s most damaging t r e e r u s t f u n g i ( P e t e r s o n 1 9 6 2 b ) ; C. comandrae has been r e p o r t e d t h e most d e s t r u c t i v e stem r u s t on h a r d p i n e s o f w e s t e r n N o r t h A m e r i c a i • ( P e t e r s o n 1 9 6 2 a ; Hawksworth 1 9 6 4 ) . Hawksworth ( 1 9 6 4 )  suggested  t h a t C. comandrae was one o f t h e t h r e e major  d i s e a s e s o f P. c o n t o r t a . The l i f e - c y c l e o f C. comandrae has been summarized b y Hedgcock  -  and Long ( 1 9 1 5 a ) , M i e l k e ( 1 9 6 1 ) ,  3  -  and M i e l k e e t a l . ( 1 9 6 8 ) , but the e n v i r -  onmental c o n d i t i o n s under w h i c h each spore s t a t e develops o r s u r v i v e s are not w e l l known.  C. comandrae i s a h e t e r o e c i o u s r u s t w h i c h grows p e r e n n i a l l y  i n the l i v i n g b a r k o f h a r d p i n e s , and develops h o s t s o f t h e genera Comandra and Geocaulon. the l a t e summer and f a l l ;  a n n u a l l y on the a l t e r n a t e  The p i n e i s i n f e c t e d d u r i n g  t h e p y c n i a appear i n t h e summer 2 o r 3 y e a r s  i n i t i a l i n f e c t i o n , and a e c i a are produced t h e f o l l o w i n g y e a r .  The  after  aecio-  spores are d i s p e r s e d and i n f e c t the a l t e r n a t e h o s t s , where s e v e r a l generat i o n s o f u r e d i a may these germinate p l e t i n g the l i f e  be produced.  T e l i a f o l l o w the u r e d i a l s t a t e ,  and  t o produce b a s i d i o s p o r e s w h i c h are a b l e t o i n f e c t p i n e , comc y c l e o f the r u s t .  quent i n t e r v a l s ( K r e b i l l 1965;  Heavy p i n e i n f e c t i o n o c c u r s a t i n f r e -  M i e l k e 1957;  P e t e r s o n 1 9 6 2 a ; Wagener i 9 6 0 )  g i v i n g r i s e t o t h e use o f t h e term "wave" y e a r s o f i n f e c t i o n . gests that s a t i s f a c t o r y environmental e v e r y y e a r , o r even e v e r y decade.  This  sug-  c o n d i t i o n s f o r i n f e c t i o n do not  occur  C h a r a c t e r i s t i c a l l y a n a t i v e pathogen  i s i n s t a b l e b a l a n c e w i t h i t s h o s t s and n a t u r a l environment, b u t i f t h i s b a l a n c e i s u p s e t then l o c a l o r widespread outbreak  —  epidemics w i l l  result.  For  an  an i n t e n s i f i c a t i o n o r spread o f t h e comandra b l i s t e r r u s t t o  o c c u r , e n v i r o n m e n t a l f a c t o r s would have t o be n o n - l i m i t i n g f o r t h r e e s e p a r a t e a e r o b i o l o g i c a l phases d u r i n g the l i f e year.  c y c l e o f t h e r u s t i n any  one  A l t h o u g h the p r e s e n t s t u d y c o n c e n t r a t e s on the f i r s t phase o f a e c i o -  spore s p o r u l a t i o n on p i n e t o d i s p e r s a l and g e r m i n a t i o n on the a l t e r n a t e h o s t , s i m i l a r f a c t o r s would a l s o a f f e c t the r e p e t i t i v e u r e d i a l phase on the a l t e r n a t e h o s t , and the phase between b a s i d i o s p o r e p r o d u c t i o n on t h e a l t e r n a t e h o s t and i n f e c t i o n on s u s c e p t i b l e p i n e .  The b a s i d i o s p o r e phase,  - k  -  i s p r o b a b l y t h e most c r i t i c a l f o r d e t e r m i n i n g o u t b r e a k s or wave y e a r s o f pine i n f e c t i o n .  The b a s i d i o s p o r e phase was  as i t s spore s t a t e was The  not s e l e c t e d f o r s t u d y here,  not v e r y s u i t a b l e f o r d i s t a n c e d i s p e r s a l s t u d i e s .  i n f o r m a t i o n g a i n e d i n t h i s s t u d y can be r e l a t e d t o the p r o b a b l e  b i o l o g y o f t h e o t h e r r u s t spore s t a t e s , and t o t h a t o f o t h e r  aero-  important  C r o n a r t i u m r u s t s , and f o r e s t pathogens i n g e n e r a l . The  a e r o b i o l o g i c a l a s p e c t s w h i c h p l a y a r o l e i n t h e sequence o f  development o f each spore s t a t e o f t h e r u s t and w h i c h must be  considered  i n any a e r o b i o l o g y phase, i n c l u d e b o t h m e t e o r o l o g i c a l and b i o l o g i c a l vironmental factors.  en-  The main a s p e c t s w h i c h a r e c o n s i d e r e d i n the p r e s e n t  s t u d y i n c l u d e , a) the c h a r a c t e r o f the a e c i o s p o r e s , t h e i r p r o d u c t i o n  and  r e l e a s e , b) the boundary l a y e r and t u r b u l e n t f a c t o r s o f the atmosphere as they, a f f e c t escape, t r a n s p o r t , d i s p e r s i o n and d e p o s i t i o n o f a e c i o s p o r e s , c) the e n v i r o n m e n t a l f a c t o r s a f f e c t i n g a e c i o s p o r e g e r m i n a t i o n , d) the e n v i r o n m e n t a l e f f e c t s on a e c i o s p o r e v i a b i l i t y d u r i n g these e v e n t s ,  and  e) b i o l o g i c a l f a c t o r s a f f e c t i n g a e c i o s p o r e p r o d u c t i o n , d i s p e r s a l , germinat i o n and  viability.  - 5 -  THE  ORGANISM  HISTORY OF THE FUNGUS In  the 1870*s C. H. Peck d e s c r i b e d two new r u s t f u n g i .  f i v e y e a r s l a t e r t h e y were connected the same fungus.  Thirty-  as the a e c i a l and t e l i a l s t a t e s o f  I n l875.Peck d e s c r i b e d as a new s p e c i e s , under the name  Periderm!um p y r i f o r m e , a c a u l i c o l o u s o r s t e m - i n h a b i t i n g Peridermium  with  a e c i a h a v i n g obovate, p y r i f o r m , o r o b l o n g - p y r i f o r m s p o r e s , from a s p e c i - . men  c o l l e c t e d b y J . B. E l l i s on p i n e branches a t N e w f i e l d , New J e r s e y .  Peck was  i n f o r m e d b y E l l i s t h a t the c o l l e c t i o n may have been made i n  G e o r g i a , and p l a c e d b y a c c i d e n t among h i s New J e r s e y specimens. and Kern (l9lk),  a f t e r examining  Arthur  the type specimen, b e l i e v e d the i n -  s c r i p t i o n on the o r i g i n a l specimen voucher was c o r r e c t . no r e c o r d s o f any c o l l e c t i o n o f t h i s r u s t i n G e o r g i a .  There are Hedgcock and  Long (1915b) suggested P i n u s r i g i d a M i l l , was the h o s t o f the type men,  as t h i s was the o n l y n a t i v e s p e c i e s o f p i n e i n the type  w h i c h was found i n f e c t e d elsewhere In  still  speci-  locality  (Vermont).  1879 Peck d e s c r i b e d the second fungus, the t e l i a l s t a t e on  the stems o f Comandra p a l l i d a A.DC. C r o n a r t i u m comandrae.  from Colorado, as a new s p e c i e s ,  He s t a t e d t h a t t h i s s p e c i e s appeared t o be a p a l e r  form o f the same s p e c i e s w h i c h o c c u r r e d on l e a v e s o f Comandra u m b e l l a t a (L.) N u t t .  A specimen o f C. comandrae was c o l l e c t e d b y E l l i s on _C.  u m b e l l a t a ( E l l i s and E v e r h a r t , N o r t h American F u n g i , No. 1082) near Newf i e l d , New J e r s e y , i n 1879, w h i c h was used b y Hedgcock and Long (1915a) as more evidence t h a t the type l o c a t i o n f o r P. p y r i f o r m e was c o r r e c t .  An  - 6 -  e a r l y c o l l e c t i o n by T. G-. L e a from Ohio On Thesium umbellatum (C. umbell a t a ) was g i v e n t h e name C r o n a r t i u m a s c l e p i a d e u m Kze. v a r . T h e s i i by Berkeley ( l 8 4 5 ) , deum. In  t o d i s t i n g u i s h t h e r u s t from t h e European C. a s c l e p i a -  B e r k e l e y s t a t e d then t h a t t h i s v a r i e t y might be a d i s t i n c t  species.  1895 Lagerheim, e l e v a t e d i t t o s p e c i e s s t a t u s and termed i t C r o n a r -  tium t h e s i i .  I n t h e meantime Peck (1879) had c o r r e c t l y g i v e n t h e t e l i a l  s t a t e o f t h e r u s t f u l l s p e c i e s s t a t u s as C. comandrae, i t s p r e s e n t v a l i d name. F o l l o w i n g Peck's d e s c r i p t i o n o f P e r i d e r m i u m p y r i f o r m e , cons i d e r a b l e c o n f u s i o n developed i n t h e use o f t h i s b i n o m i a l , w h i c h produced many m i s - i d e n t i f i c a t i o n s o f specimens and erroneous r e p o r t s o f t h e o c c u r rence o f t h e comandra r u s t .  The:specimens  which probably i n i t i a t e d  this  c o n f u s i o n were c o l l e c t i o n s by E l l i s o f P e r i d e r m i u m r u s t s from near t h e type l o c a t i o n o f P. p y r i f o r m e a t N e w f i e l d .  One was c o l l e c t e d on P i n u s  i n o p s A i t (P. v i r g i n i a n a M i l l s ) i n May 1882 ( i s s u e d as P. p y r i f o r m e , Wo. 1021, E l l i s , N o r t h American F u n g i ) , and t h e o t h e r s were c o l l e c t e d on P. v i r g i n i a n a i n A p r i l 1890, and on P. r i g i d a i n May 1890 ( b o t h i n t h e Herbarium, New York B o t a n i c a l Garden).  The l a t t e r two were o r i g i n a l l y  l a b e l l e d P e r i d e r m i u m p i n i W a l l r . , a European Peridermium, b u t were l a t e r r e f e r r e d t o P. p y r i f o r m e . However, t h e s e have now been r e f e r r e d t o P. comptoniae  ( A r t h . ) O r t o n and Adams.  These l a t e r c o l l e c t i o n s o f E l l i s  l e a d A r t h u r and K e r n (1906) and A r t h u r (1907a) t o d e s c r i b e as P. p y r i forme what i s now known as P. comptoniae, c o m p l e t e l y i g n o r i n g Peck's d e s c r i p t i o n s t a t i n g t h a t t h e a e c i o s p o r e s were p y r i f o r m .  C l i n t o n (1908)  l a t e r connected t h i s so c a l l e d P. p y r i f o r m e ( P . comptoniae) w i t h  - 7 -  Cronartium  comptoniae A r t h . , a t e l i a l s t a t e o c c u r r i n g on Comptonia  p e r e g r i n a (L.) C o u l t . and M y r i c a spp.  P r i o r t o t h i s , Underwood and  E a r l e (1896) had c a s t doubt on t h e s t a t u s o f P. p y r i f o r m e  and had sug-  g e s t e d t h a t i t was p o s s i b l y a synonym o f P e r i d e r m i u m cerebrum Peck, a s p e c i e s d e s c r i b e d i n 1873.  They suggested t h i s , because t h e y found  the s t r i k i n g p y r i f o r m spores o f P. p y r i f o r m e P.  cerebrum as t h e y occur i n t h e South".  occurred  " i n b o t h forms o f  T h i s i n d i c a t e s f u r t h e r con-  f u s i o n o f t h e i d e n t i t y o f P e r i d e r m i u m forms on p i n e , as o n l y the t r u e P. p y r i f o r m e has p y r i f o r m  spores.  Hedgcock and Long (1913) d e s c r i b e d a c a u l i c o l o u s s p e c i e s o f Peridermium w i t h ovoid t o p y r i f o r m aeciospores l e c t e d b y E. B e t h e l from Colorado,  on P i n u s  contorta col-  as a new s p e c i e s , P e r i d e r m i u m b e t h e l i .  Unable t o compare t h e specimen o f B e t h e l w i t h type m a t e r i a l o f P. p y r i forme , t h e y r e l i e d (Hedgcock and Long 1915a) on t h e p u b l i s h e d o f A r t h u r and K e r n (1906) w h i c h concluded inaccurate concerning K e r n (1913  5  P. p y r i f o r m e  statement  t h a t Peck's d e s c r i p t i o n was  the p y r i f o r m shape o f t h e a e c i o s p o r e s .  A r t h u r and  1914), however, d i s c a r d e d t h e i r o r i g i n a l i n t e r p r e t a t i o n o f a f t e r s e e i n g a specimen s u b m i t t e d  i n 1913 b y W. P. F r a s e r ,  c o l l e c t e d on P i n u s p o n d e r o s a a t Vernon, B r i t i s h Columbia.  This  specimen  had t h e t y p i c a l p y r i f o r m spores d e s c r i b e d o r i g i n a l l y b y Peck. A r t h u r and K e r n (1914) s p e c u l a t e d t h a t t h e a l t e r n a t e s t a t e s o f t h i s P e r i d e r m i u m would be found on s p e c i e s o f Comandra, as t h i s genus had t h e o n l y u n a t t a c h e d  species o f Cronartium  t h e n known.  A I907  col-  l e c t i o n o f P. p y r i f o r m e on P i n u s c o n t o r t a from D e v i l ' s Lake (now Lake Minnewanka), B a n f f , A l b e r t a , b y E. W. D. Holway, gave support  to this  - 8 -  s p e c u l a t i o n , as Holway wrote on the p a c k e t t h a t the P e r i d e r m i u m on p i n e was undoubtedly Adams (igik) Cronartium  a s s o c i a t e d w i t h a C r o n a r t i u m on Comandra.  Orton and  found t r u e P. p y r i f o r m e on P i n u s pungens Lamb., and comandrae on Comandra u m b e l l a t a w i t h i n ho f e e t o f the i n -  fected pine.  They concluded t h a t t h i s C r o n a r t i u m was t h e a l t e r n a t e s t a t e  o f P. p y r i f o r m e . P. p y r i f o r m e .  They a l s o suggested  Hedgcock and Long (lyik)  t h a t P. b e t h e l i was a synonym o f s u c c e s s f u l l y i n f e c t e d Comandra  u m b e l l a t a w i t h a e c i o s p o r e s o f P. p y r i f o r m e , c o n f i r m i n g the l i f e the r u s t and gave i t t h e new c o m b i n a t i o n  cycle of  Cronartium pyriforme.  I n the  f o l l o w i n g y e a r (1915a) t h e y gave the morphology, l i f e h i s t o r y , and  distri-  b u t i o n o f t h i s r u s t as t h e n known under t h e new c o m b i n a t i o n C. p y r i f o r m e . Kirkwood  (1915) i n o c u l a t e d P i n u s ponderosa w i t h spores from Comandra  ( p a l l i d a ? ) t h r o u g h i n c i s i o n s i n the bark, w h i c h r e s u l t e d i n the ment o f r u s t hyphae w i t h i n the bark and wood of-.--the p i n e .  develop-  T h i s was t h e  f i r s t r e c o r d o f t h e s u c c e s s f u l t r a n s f e r o f C. comandrae from Comandra t o pine.  He a l s o s u c c e s s f u l l y i n o c u l a t e d Comandra p l a n t s w i t h a e c i o s p o r e s  from p i n e .  More r e c e n t l y , Andrews e t a l . (1963) r e p o r t e d s u c c e s s f u l i n -  f e c t i o n o f P i n u s c o n t o r t a w i t h b a s i d i o s p o r e s o f C. comandrae.  Telial  horns were p l a c e d among and above t h e n e e d l e s , w i t h p y c n i a b e i n g a year l a t e r .  observed  Boyce (1916) f i r s t d e s c r i b e d the p y c n i a and p y c n i o s p o r e s  o f C. comandrae from a c o l l e c t i o n on P i n u s p o n d e r o s a made i n J u l y 1916 in California. and p y c n i o s p o r e s  I n the same y e a r , Weir and Hubert (1917) observed  o c c u r r i n g on P. p o n d e r o s a and P. c o n t o r t a , c o n f i r m i n g  the f i n d i n g s o f Boyce. to  pycnia  Meinecke (1929) t r i e d u n s u c c e s s f u l l y f o r 6 y e a r s  i n o c u l a t e d i r e c t l y from p i n e t o p i n e w i t h a e c i o s p o r e s .  This i n d i c a t e s  _ 9 -  t h a t C. comandrae was o n l y h e t e r o e c i o u s and can not be a u t o e c i o u s as i s P e r i d e r m i u m h a r k n e s s i i J.P. Moore (Meinecke 1 9 l 6 ,  1920, I 9 2 9 ;  Hedgcock and Hunt 1920). Many e a r l y h e r b a r i u m specimens o f C. comandrae a r e s t i l l l a b e l l e d w i t h synonyms o r c o m b i n a t i o n s o f them.  Some e a r l y c o l l e c t i o n s  were p l a c e d under t h e form-genus A e c i d i u m ( F a r l o w 1905) i n s t e a d o f P e r i dermium, some o f w h i c h were even n o n - a e c i a l c o l l e c t i o n s ( e g . A e c i d i u m a s c l e p i a d e u m v a r . t h e s i i on Comandra p a l l i d a from Kamloops, B r i t i s h Columbia).  Hedgcock and Long (1915a) gave Caeoma comandrae Peck as a  synonym o f C r o n a r t i u m p y r i f o r m e .  Caeoma comandrae was d e s c r i b e d by  Peck (1884) from a specimen c o l l e c t e d b y M. E. Jones on Comandra p a l l i d a i n Utah., However, a r e c e n t check o f t h e New York B o t a n i c a l Garden Herbarium, where Hedgcock and Long r e p o r t e d t h e specimen d e p o s i t e d , r e v e a l e d no specimen h e l d as Caeoma comandrae, b u t one h e l d as Uredo comandrae b y t h e same c o l l e c t o r .  T h i s specimen p r o v e d t o be P u c c i n i a  comandrae Peck, a s p e c i e s a l s o d e s c r i b e d b y Peck i n 1884 b u t from a c o l l e c t i o n b y T. S. Brandeges i n Washington T e r r i t o r y .  This leaves the  q u e s t i o n o f t h e v a l i d i t y o f Caeoma comandrae as a synonym i n doubt, as i t has not been l i s t e d b y o t h e r s .  - 10 -  DISTRIBUTION AND HOSTS  Comandra b l i s t e r r u s t has been r e p o r t e d on one i n t r o d u c e d and t e n n a t i v e hard p i n e s p e c i e s i n N o r t h A m e r i c a , r a n g i n g f r o m New B r u n s w i c k to New  the Yukon and southward t o Tennessee, n o r t h e r n Alabama, M i s s i s s i p p i , Mexico and C a l i f o r n i a .  I t has not been r e p o r t e d from A l a s k a (Cash  1953; L a u r e n t , p e r s o n a l communication 1966) o r Mexico a l t h o u g h i b l e p i n e s do o c c u r .  suscept-  I t has not been i n t r o d u c e d o u t s i d e N o r t h A m e r i c a  b u t i s a p o t e n t i a l l y dangerous r u s t t o n a t i v e h a r d p i n e s o f Europe and Asia.  The u r e d i a l and t e l i a l s t a t e s o f the r u s t have been r e c o r d e d  over a s i m i l a r range on the a l t e r n a t e h o s t s , Comandra and Geocaulon o f the f a m i l y S a n t a l a c e a e .  The e a r l i e s t l i s t s o f d i s t r i b u t i o n and h o s t s  were g i v e n b y Hedgcock and Long (1915a) and b y A r t h u r (1907b, 1927, 1934).  1925,  More r e c e n t c o l l e c t i o n s have extended the known d i s t r i -  b u t i o n o f the r u s t and r e - i d e n t i f i c a t i o n has e l i m i n a t e d some o f t h e earlier reports. A l l the C r o n a r t i u m  comandrae m a t e r i a l has been seen i n t h e  f o l l o w i n g h e r b a r i a (Herbarium codes are those used i n the " i n d e x Herba r i o r u m " , Lanjouw and S t a f l e u (196U):  CFB, DAOM, DAVFP, PUR, UBC, WIN,  and c r i t i c a l . . ' m a t e r i a l has been seen from the f o l l o w i n g h e r b a r i a : B P I , MISSA, NY.  L i s t s o r i n f o r m a t i o n have been o b t a i n e d from the f o l l o w i n g  h e r b a r i a not i n c l u d e d above:  ALTA, FFB,  MFB, MONT, QFB, QMP, SASK,  WINF, WSP, MacDonald C o l l e g e , S t e . A n n e - d e - B e l l e v u e , and many o f t h e r e g i o n a l l a b o r a t o r i e s o f the U n i t e d S t a t e s F o r e s t S e r v i c e .  - 11 -  D i s t r i b u t i o n o f the Rust on P i n u s P i n u s a t t e n u a t a Lemm. T h i s host was not r e c o r d e d b y A r t h u r (1934) o r more r e c e n t l y , by P e t e r s o n (1962b), b u t t h e r e are specimens from Grant P a s s , Oregon, c o l l e c t e d b y J . R. Weir, Sept. 6, 1916, i n the PUR and BPI h e r b a r i a .  A  r e c e n t p u b l i c a t i o n by P e t e r s o n (1967) does i n c l u d e t h i s p i n e as a h o s t . P i n u s b a n k s i a n a Lamb. A r t h u r (1934) r e p o r t e d C. comandrae o c c u r r e d on t h i s h o s t i n M i c h i g a n , W i s c o n s i n , A l b e r t a and Saskatchewan.  Hedgcock and Long (1915b)  r e p o r t e d s e v e r a l c o l l e c t i o n s from M i c h i g a n and. M i n n e s o t a on P. d i v a r i c a t a (=_P. b a n k s i a n a ) .  I t was a l s o r e p o r t e d from New Brunswick  N e w a l l 1957), Quebec (Pomerleau  1942), O n t a r i o (Conners  ( D a v i d s o n and  1934) and Mani-  t o b a ( R i l e y e t a l . 1952).  P i n u s c o n t o r t a Dougl. Recorded b y A r t h u r (1934) from C o l o r a d o , Idaho, Wyoming and Alberta.  Hedgcock and Long (1915b) r e p o r t e d the o c c u r r e n c e i n M i c h i g a n ,  and P e t e r s o n (1962b) i n C a l i f o r n i a , Montana, Oregon, U t a h and Washington. A l s o r e c o r d e d i n B r i t i s h Columbia ( Z i l l e r 1953) and the Yukon (Molnar i960).  I n I968 two specimens were c o l l e c t e d from the Cypress  Hills,  Saskatchewan (CFB 8406, 8407).  Pinus echinata M i l l . Four r e c e n t r e p o r t s o f the r u s t on t h i s h o s t are from M i s s o u r i ( B e r r y e t a l . 1961), Arkansas  (U.S. F o r e s t S e r v i c e I962; P o o l i n g e t a l .  1964), and Tennessee (Powers e t a l . I967).  - 12 -  P i n u s j e f f r e y i Grev. & B a l f . Peterson  (1962b) r e p o r t e d t h i s s p e c i e s as a h o s t i n C a l i f o r n i a  and Oregon, and I have seen specimens c o l l e c t e d b y L e w i s , from near t h e S i s k i y o u N a t i o n a l F o r e s t , Oregon ( B P I , PUR). P i n u s ponderosa Laws. A r t h u r (193^) r e p o r t e d i t from C a l i f o r n i a ,  C o l o r a d o , Montana,  Oregon, South Dakota, Utah, Washington and B r i t i s h Columbia.  Peterson  (1962b) added t h e s t a t e s o f A r i z o n a , Idaho, Nevada and Wyoming. C o l l e c t i o n s on P. ponderosa v a r . a r i z o n i c a (Engelm.) Shaw a r e i n c l u d e d i n t h i s distribution.  P l a n t a t i o n s o f ponderosa p i n e have r e c e n t l y been r e p o r t e d  i n f e c t e d i n Tennessee  (Powers e t . a l . 1967).  P i n u s pungens Lamb. Orton and Adams (l^lh)  r e p o r t e d t h e r u s t from  Pennsylvania.  Pinus r i g i d a M i l l . There i s a n o n - v e r i f i a b l e specimen from t h i s h o s t , c o l l e c t e d at Mt. C a l v a r y , New J e r s e y i n 1917  (PUR), b u t Hedgcock and Long (1915b)  r e p o r t e d a c o l l e c t i o n from Vermont.  P i n u s s y l v e s t r i s L. The r u s t has been r e p o r t e d on t h i s i n t r o d u c e d p i n e , from Washi n g t o n (Shaw 1958) and C o n n e c t i c u t  ( S p a u l d i n g 1958).  I n Canada i t has  been r e p o r t e d from M a n i t o b a (Thomas 1953), Saskatchewan ( I v e s e t a l . I 9 6 8 ) , and A l b e r t a ( P o w e l l and Morf 1965).  Specimens  f r o m Roscommon,  M i c h i g a n a r e d e p o s i t e d i n t h e BPI and PUR h e r b a r i a , n e i t h e r i s v e r i f i a b l e ,  - 13 -  and i s p r o b a b l y C. comptoniae.  A specimen c o l l e c t e d i n 1887 i n t h e M i s -  s o u r i B o t a n i c a l Garden, S t . L o u i s , M i s s o u r i ( A r t h u r and K e r n I906) and now  i n t h e B P I herbarium,  probably Cronartium  was l a b e l l e d P e r i d e r m i u m p y r i f o r m e , b u t i s '  comptoniae as i t does not have t h e p y r i f o r m s p o r e s .  P i n u s t a e d a L. M i l e s (1934) f i r s t r e p o r t e d t h e r u s t on t h i s host from M i s s i s s i p p i , and more r e c e n t l y i t has been, r e p o r t e d from p l a n t a t i o n s i n Tennessee and t h e n o r t h e a s t e r n corner o f Alabama (Powers e t a l . 1967).  Pinus v i r g i n i a n a  Mill.  There i s a lQlk  c o l l e c t i o n b y Hedgcock from P e t e r b u r g , Penn-  s y l v a n i a , i n t h e PUR herbarium  a t t r i b u t e d t o t h i s host, b u t the c o l l e c t i o n  i s n o t v e r i f i a b l e , and was n o t l i s t e d b y Hedgcock and Long (1915a, or A r t h u r (193*+).  Peterson  . l i k e l y from t h i s p i n e .  1915b)  (1967) r e p o r t e d a P i n u s sp. specimen as  C o r d e l l e t a l . (1967) noted t h a t p l a n t e d and  n a t u r a l P. v i r g i n i a n a were n o t i n f e c t e d i n Tennessee, a l t h o u g h a t s e v e r a l l o c a t i o n s t h e y were i n t e r m i x e d w i t h i n f e c t e d P. t a e d a .  Other r e p o r t s The  i n t r o d u c e d s p e c i e s P. n i g r a A r n o l d and P. p i n a s t e r A i t .  have a l s o been r e p o r t e d as h o s t s o f C. comandrae (Boyce 19^3; 1958;  M i e l k e 196I; V e r r a l l 1964), b u t P e t e r s o n  Spaulding  (1962b) p o i n t s o u t t h a t  these were based on a r e p o r t b y C l i n t o n (1912) a t t h e time when P e r i dermium p y r i f o r m e was o f t e n l i n k e d w i t h C. comptoniae ( A r t h u r and K e r n I906; C l i n t o n 1908).  There i s a l s o a I918 n o n - v e r i f i a b l e specimen l a -  b e l l e d as C. comandrae from Roscommon, M i c h i g a n on P. n i g r a i n t h e BPI  - Ik -  Peterson. (1966b) i n d i c a t e d the erroneous r e p o r t (Gremmen  herbarium.  1964). o f P. r e s i n o s a as a h o s t f o r C. comandrae i n Vermont. i o n upon w h i c h t h i s r e p o r t was based was The  o f the r u s t on  Two comandrae.  collect-  comptoniae.  d i s t r i b u t i o n o f C. comandrae on P i n u s spp.  e a s t e r n Canada i s shown i n F i g s . 1 and Distribution  a g a i n C.  The  i n western  and  2.  Santalaceae  g e n e r a o f the S a n t a l a c e a e  serve as a l t e r n a t e h o s t s f o r C.  The d i s t r i b u t i o n f o r the genus Comandra has r e c e n t l y been  m o d i f i e d as P i e h l  (1965) r e a r r a n g e d  the genus, and p l a c e d what were f i v e  s p e c i e s (C. u m b e l l a t a ( L . ) N u t t . , C. c a l i f o r n i c a Eastw., C. r i c h a r d s i a n a F e r n a l d , C_. p a l l i d a A. DC., s p e c i e s o f one  C. elegans  s p e c i e s (C. u m b e l l a t a ) .  (Rochel) Reichenb.) as f o u r subT h i s i s not too  e s p e c i a l l y i n Canada, as h i s d i s t r i b u t i o n map  satisfactory,  i n d i c a t e s t h a t t h e r e are  l a r g e zones w h i c h have i n t e r m e d i a t e s between two  subspecies.  However,  h i s arrangement has been f o l l o w e d , as i t does e l i m i n a t e the p r e v i o u s p r o b l e m o f d i s t i n g u i s h i n g between C_. u m b e l l a t a and C. r i c h a r d s i a n a , "since t h e y are now  one  subspecies  (ssp. umbellata).  E a r l i e r , Fernald  (1928) had s e p a r a t e d the s p e c i e s C. l i v i d a R i c h a r d s o n Comandra, and formed a new F e r n a l d as the t y p e . c a u l o n has  Piehl  genus, Geocaulon w i t h G.  from the genus  lividum  (Richardson)  (1965) agreed w i t h t h i s s e p a r a t i o n , as Geo-  s e v e r a l f e a t u r e s t h a t s e p a r a t e i t from Comandra.  and Geocaulon are h e m i p a r a s i t e s  (Figs.  3,  k, 5 and 6 ) .  Both Comandra  S e v e r a l papers  have l i s t e d the p l a n t s p e c i e s upon w h i c h Comandra i s p a r a s i t i c 1937;  F i s h e r 1922;  H a r r i n g t o n 19^5;  Hedgcock 1915;  Moss 1926;  (Brooks '.v Piehl.  F i g . 1.  D i s t r i b u t i o n o f C r o n a r t i u m comandrae on P i n u s s p e c i e s i n w e s t e r n Canada.  F i g . 2.  D i s t r i b u t i o n o f Cronartium e a s t e r n Canada.  comandrae on P i n u s s p e c i e s i n  Fig.  3-  F i g . k.  Fig.  5.  F i g . 6.  A e r i a l stems o f Geocaulon l i v i d u m , w i t h a x i l l a r y g r e e n i s h flowers.  A e r i a l stems o f Comandra u m b e l l a t a s s p . p a l l i d a , w i t h m i n a l w h i t e f l o w e r s about t o b r e a k open.  ter-  S e r i e s o f a e r i a l stems o f Comandra u m b e l l a t a s s p . p a l l i d a , w h i c h b r a n c h j u s t below t h e s o i l and a r e a t t a c h e d t o an . underground-rhizome. Some a e r i a l stems dead from p r e v i o u s year.  Large h a u s t o r i u m o f Comandra u m b e l l a t a s s p . p a l l i d a a t t a ched t o Populus t r e m u l o i d e s Michx. r o o t , w i t h s m a l l e r h a u s t o r i u m on s m a l l e r r o o t .  17  -  - 18 -  1965)5  1926,  b u t I am aware o f o n l y one p u b l i s h e d r e p o r t f o r Geocaulon (Moss as C. l i v i d a ) .  Geocaulon i s a genus w i t h a d i s t i n c t l y n o r t h e r n  d i s t r i b u t i o n i n N o r t h America,  e x t e n d i n g from Newfoundland t o A l a s k a ,  and o n l y as f a r south as n o r t h e r n New England, W i s c o n s i n , M i c h i g a n , Minnesota,  Idaho and Washington ( F e r n a l d 1928; H a r r i s 1965; H i t c h c o c k  e t a l . 1964; Raup 1947 p i . X X I I ) .  Comandra u m b e l l a t a ( L . ) N u t t . ssp. u m b e l l a t a P i e h l  (=C. u m b e l l a t a ; CJ.  richardsiana). A r t h u r (1934) r e p o r t e d C. comandrae o c c u r r e d on t h i s h o s t i n C a l i f o r n i a , Delaware, I l l i n o i s , I n d i a n a , Iowa, M a s s a c h u s e t t s , Mississippi,  Michigan,  New Hampshire, New J e r s e y , New York, Ohio, Oregon, P e n n s y l -  v a n i a , Washington, W i s c o n s i n , O n t a r i o and Quebec.  P i e h l ' s (1965')  dis-  t r i b u t i o n map suggests t h a t t h e r e c o r d s f o r C a l i f o r n i a , Oregon, and Washington were on t h e ssp. c a l i f o r n i c a and n o t u m b e l l a t a .  Hedgcock  and Long (1915a) r e p o r t e d a. c o l l e c t i o n from Vermont, and Powers e t a l . (1967) r e c o r d i t from Alabama and Tennessee. t r i c t o f Columbia and M i n n e s o t a lections  Specimens from t h e D i s -  a r e d e p o s i t e d i n t h e BPI h e r b a r i u m .  Col-  l i s t e d as C. u m b e l l a t a o r C. r i c h a r d s i a n a from M a n i t o b a and  Saskatchewan a r e i n t h e UBC and WINF h e r b a r i a , and a r e p r o b a b l y t r u e C. u m b e l l a t a spp. u m b e l l a t a , o r an i n t e r m e d i a t e between t h i s and s s p . p a l lida. Comandra u m b e l l a t a (L.) N u t t . ssp. p a l l i d a (A.DC.) P i e h l  ( =C. p a l l i d a )  A r t h u r (1934) r e p o r t e d i t from A r i z o n a , C o l o r a d o , Idaho, Minnesota, M i s s o u r i , Montana, New Mexico, N o r t h Dakota,  Oregon,  Iowa,  South  - 19 -  Dakota, Utah, Washington, Wyoming, A l b e r t a , B r i t i s h Columbia, Northwest Territories lections  and Saskatchewan.  Hedgcock and Long ( 1 9 1 5 a ) r e p o r t e d  col-  f r o m Nebraska, and a c o l l e c t i o n from M i c h i g a n i s d e p o s i t e d i n  the PUR h e r b a r i u m .  There i s a specimen o f t h i s h o s t from Texas i n t h e  BPI h e r b a r i u m , and one r e p o r t e d as Comandra sp. (NY herbarium) w h i c h s h o u l d p r o b a b l y be r e f e r r e d t o ssp. p a l l i d a . o c c u r r e n c e i n M a n i t o b a , and Conners  Bisby (1938) reported i t s  (1934) i n Ontario.  There i s some  doubt about t h e specimen upon w h i c h A r t h u r ( 1 9 3 4 ) based h i s Northwest T e r r i t o r i e s record.  J . A. Parmelee  ( p e r s o n a l communication 1 9 6 6 ) b e -  l i e v e d t h i s r e c o r d was based on the DAOM specimen, No. l 8 6 l , M a r t i n C a b i n , S l a v e Lake A l b e r t a , c o l l e c t e d interpreted  i n 1 9 2 9 , and t h a t A r t h u r  'Slave Lake' as Great S l a v e Lake, Northwest  i n s t e a d o f L e s s e r S l a v e Lake, A l b e r t a . the Northwest T e r r i t o r i e s  f r o m near  Territories,  However, A r t h u r ( 1 9 2 7 )  included  i n h i s d i s t r i b u t i o n l i s t p r i o r t o the  o f t h e above DAOM specimen.  collection  There i s no r e c o r d o f Comandra from t h e  Northwest T e r r i t o r i e s o r t h e Yukon i n the DAO o r CAN h e r b a r i a , o r more r e g i o n a l h e r b a r i a .(ALTA, CAFB, UAC, UBC), n o r t h o f about 59°N. l a t i t u d e  i n Alberta.  and i t has not been  collected  P o r s i l d and Cody ( 1 9 6 8 ) , i n  t h e i r c h e c k l i s t o f v a s c u l a r p l a n t s i n t h e c o n t i n e n t a l Northwest T e r r i t o r i e s , i n d i c a t e d t h a t C_. p a l l i d a was e x p e c t e d t o o c c u r , b u t a t p r e s e n t t h e r e i s no r e c o r d from the. a r e a .  Comandra u m b e l l a t a ( L . ) N u t t , ssp. c a l i f o r n i c a (Eastw.) P i e h l . I am aware o f no h e r b a r i u m specimens under t h i s s u b s p e c i e s name, b u t a c c o r d i n g t o P i e h l ' s ( 1 9 6 5 ) d i s t r i b u t i o n map, t h e specimens r e c o r d e d b y Hedgcock and Long ( 1 9 1 5 a ) and A r t h u r ( 1 9 3 4 ) under C.  - 20 -  u m b e l l a t a from C a l i f o r n i a , Oregon, and Washington, s h o u l d be s s p .  califor-  nica.  Geocaulon  l i v i d u m (Richardson) F e r n a l d A r t h u r (1934) gave t h e f o l l o w i n g d i s t r i b u t i o n o f t h e r u s t on  t h i s h o s t (as Comandra l i v i d a ) , W i s c o n s i n , M a n i t o b a and Quebec. and Molnar  (1953) f i r s t r e p o r t e d the o c c u r r e n c e i n B r i t i s h  and Molnar  (i960) i n t h e Yukon.  Baranyay  r e p o r t e d t o be on Comandra.  Territories,  and B o u r c h i e r  though i n a c c u r a t e l y  A c o l l e c t i o n made i n 1937 b y D. V. B a x t e r  (DAOM 5558 and PUR 48514) from G r e a t S l a v e Lake, Northwest and l a b e l l e d Comandra sp. p r o v e d t o be G. l i v i d u m . h e r b a r i u m specimens,  Columbia,  e t a l . (1962) f i r s t r e p o r t e d  t h e i n f e c t e d h o s t i n A l b e r t a , and a y e a r l a t e r Baranyay (1963) r e p o r t e d i t i n the Northwest  Ziller  some l a b e l l e d  Territories,  Other u n r e p o r t e d  as C. l i v i d a , extend the known d i s -  t r i b u t i o n t o Idaho (NY), Washington (DAOM) and O n t a r i o (DAOM).  Other r e p o r t s A r t h u r (1934) r e f e r r e d t o B u c k l e y a d i s t i o c h o p h y l l a ( N u t t . ) T o r r . as a h o s t o f C_. comandrae i n Tennessee.  T h i s s p e c i e s i s no  l o n g e r r e g a r d e d as a h o s t o f C. comandrae, as H e p t i n g (1957) made t h e c o n n e c t i o n o f the C r o n a r t i u m o c c u r r i n g on B u c k l e y a w i t h  Peridermium  appalachianum  H e p t i n g and Cummins, g i v i n g the c o m b i n a t i o n  appalachianum  H e p t i n g , a r u s t w h i c h o c c u r s i n N o r t h C a r o l i n a , Tennessee  and  Cronartium  Virginia. The d i s t r i b u t i o n o f C_. comandrae on Comandra and Geocaulon  w e s t e r n and e a s t e r n Canada i s shown i n F i g s . 7 and 8.  in  Very r e c e n t l y a  - 21 -  d i s t r i b u t i o n map f o r C. comandrae on P i n u s been p u b l i s h e d b y K r e b i l l (1968b).  spp. and Comandra spp.has  Generally h i s northern  distribution  l i n e f o r Canada does n o t extend f a r enough n o r t h , e s p e c i a l l y i n t h e Yukon and Northwest T e r r i t o r i e s ,  and i n M a n i t o b a and Quebec.  l e c t i o n f r o m Great Whale R i v e r , has been m i s p l a c e d o f Quebec.  i n Ontario  The  col-  instead  F i g . 7.  D i s t r i b u t i o n o f C r o n a r t i u m comandrae on t h e S a n t a l a c e a e , Comandra u m b e l l a t a and Geocaulon l i v i d u m , i n w e s t e r n Canada.  F i g . 8.  D i s t r i b u t i o n o f C r o n a r t i u m comandrae on t h e S a n t a l a c e a e , Comandra u m b e l l a t a and Geocaulon l i v i d u m , i n e a s t e r n Canada.  - 2k  -  LIFE CYCLE P i n e i n f e c t i o n by C r o n a r t i u m comandrae u s u a l l y t a k e s p l a c e d u r i n g summer and autumn t h r o u g h the n e e d l e s on branches o r main stems (Andrews e t a l . I963).  I t i s p o s s i b l e t h a t the fungus i n f e c t s young  t r e e s d i r e c t l y t h r o u g h the b a r k o r t h r o u g h wounds (Kirkwood 1915)5 t h i s mode o f e n t r y i s o f minor importance.  but  P e n e t r a t i o n o f the n e e d l e  by the germ tube o f the b a s i d i o s p o r e o r s p o r i d i u m may be t h r o u g h the s t o m a t a l pore or d i r e c t l y t h r o u g h the e p i d e r m a l c e l l s .  Following;pene-  t r a t i o n a mass o f hyphae d e v e l o p s w i t h i n the n e e d l e t i s s u e near the p o i n t o f i n f e c t i o n , t h e n h y p h a l growth p e n e t r a t e s downwards t h r o u g h the v a s c u l a r bundle o f the needle and i n t o t h e b a r k w h i c h becomes d i s c o l o r e d The l e n g t h o f the p e r i o d between needle i n f e c t i o n and b a r k d i s c o l o r a t i o n the i n c u b a t i o n p e r i o d , v a r i e s a c c o r d i n g t o t h e l e n g t h o f growing  season,  the s e a s o n a l weather, and the d i s t a n c e t h e hyphae must grow between p o i n t o f i n f e c t i o n and the bark.  The mycelium o f the fungus, w h i c h  branches i r r e g u l a r l y , r a m i f i e s i n t e r c e l l u l a r l y t h r o u g h a l l l i v e p i n e b a r k t i s s u e s , and extends i n t o the o u t e r sapwood t i s s u e s , m a i n l y i n the m e d u l l a r y r a y s (Adams 1919;  K r e b i l l 1968a).  K r e b i l l found the a d v a n c i n g  hyphae i n the phloem, never more t h a n 2.cm.beyond t h e v i s i b l e o f the i n f e c t e d stem.  swelling  The i n f e c t e d b a r k i n c r e a s e d i n t h i c k n e s s due t o  the i n t e r c e l l u l a r hyphae and expanding parenchyma c e l l s .  W i t h the i n -  crease i n t h i c k n e s s the c o n d u c t i n g s i e v e c e l l s i n t h e phloem c o l l a p s e . One or more u n i c e l l u l a r h a u s t o r i a may p e n e t r a t e i n t o a s i n g l e parenchyma c e l l from the i n t e r c e l l u l a r hyphae.  K r e b i l l also reported haustoria  sometimes p r e s e n t i n s i e v e elements and x y l e m r a y t r a c h e i d s , and noted  - 25 -  t h a t t h e h a u s t o r i a v a r y i n s i z e and morphology depending tion.  on t h e i r p o s i -  P e t e r s o n (1966a) r e p o r t e d t h e i n t e r e s t i n g o c c u r r e n c e o f C.  comandrae hyphae i n t i s s u e s o f t h e dwarf m i s t l e t o e , Arceuthobium  ameri-  canum N u t t . ex Engelm., on P i n u s c o n t o r t a , w i t h h a u s t o r i a p e n e t r a t i n g i n t o Arceuthobium parenchyma c e l l s .  The r e a c t i o n o f t h e sapwood t o i n -  f e c t i o n i s a decrease o r c e s s a t i o n o f growth i n t h e canker a r e a ( F i g s . 9 and 10).  R e s i n o s u s a l s o o c c u r s and t h e r e s i n may t h o r o u g h l y  t h e b a r k and o u t e r sapwood ( F i g s . 9? 10  a n (  i  impregnate  ll) •  The p y c n i a l o r spermogonium s t a t e appears  as a d i s t i n c t  crust-  l i k e l a y e r o f p a r a l l e l p y c n i o p h o r e s o r spermatiophores p u s h i n g up between t h e o u t e r l a y e r o f c o r t i c a l parenchyma c e l l s and t h e o v e r l y i n g t h i n w a l l e d c e l l s o f t h e p h e l l o g e n (Adams 1919).  The p y c n i o p h o r e s a r i s e  from  a zone o f i n t e r t w i n e d u n i n u c l e a t e hyphae w h i c h make up t h e base o f each pycnium.  The p y c n i o p h o r e s  c o n t a i n a s i n g l e n u c l e u s which d i v i d e s , one  n u c l e u s m i g r a t i n g i n t o t h e immature p y c n i o s p o r e o r spermatium which a r e a b s t r a c t e d from t h e f r e e ends o f t h e p y c n i o p h o r e s a t m a t u r i t y .  The pycn-  i o s p o r e s appear t o escape t h r o u g h t h e i r r e g u l a r c r a c k i n g o f t h e o v e r l y i n g p h e H u m t i s s u e , and a r e exuded w i t h a t h i n g e l a t i n o u s f l u i d t o form o r a n g e - c o l o r e d d r o p l e t s ( F i g . 1 2 ) , w h i c h e v e n t u a l l y d r y on t h e s u r f a c e o f t h e b a r k t o form d a r k - c o l o r e d s p o t s c a l l e d " p y c n i a l s c a r s " . I n s e c t s a r e a t t r a c t e d t o t h e d r o p l e t s and h e l p d i s t r i b u t e t h e p y c n i o s p o r e s . The p y c n i o s p o r e s a r e 3 t o k b y 3 t o 7M-, p y r i f o r m i n shape ( F i g . 1 3 ) , and p r o b a b l y have a s e x u a l f u n c t i o n s i m i l a r t o t h a t shown b y P i e r s o n (1933) f o r pycniospores of Cronartium r i b i c o l a Fischer.  The p y c n i a appear one  t o t h r e e y e a r s a f t e r t h e i n i t i a l i n f e c t i o n and precede t h e a e c i a l s t a t e  Fig.  9-  F i g . 10.  F i g . 11.  T r a n s v e r s e s e c t i o n s t a k e n from the c e n t r a l p o r t i o n o f ' f o u r C r o n a r t i u m comandrae cankers on l o d g e p o l e p i n e showing e c c e n t r i c growth caused by t h e r u s t . Note t h e r e s i n i m p r e g n a t i o n o f t h e o u t e r sapwood.  T r a n s v e r s e s e c t i o n s t a k e n from t h e c e n t r a l p o r t i o n o f f i v e C r o n a r t i u m comandrae cankers on l o d g e p o l e p i n e • showing e c c e n t r i c growth caused by t h e r u s t .  T r a n s v e r s e s e c t i o n t a k e n t h r o u g h t h e a e c i a l zone o f a C r o n a r t i u m comandrae canker showing t h e a e c i a s i t u a t e d i n t h e b a r k t i s s u e s . Note t h e r e s i n i m p r e g n a t i o n o f t h e b a r k between a e c i a , and t h e s w e l l i n g o f t h e o u t e r r i n g o f sapwood caused by t h e fungus.  - 26 -  F i g . 12.  F i g . 13.  P y c n i a l drops on the p y c n i a l zone o f a C r o n a r t i u m comandrae canker on the stem and branches o f a l o d g e p o l e p i n e . Note the s w e l l i n g o f the stem a s s o c i a t e d w i t h the canker.  Group o f p y r i f o r m p y c n i o s p o r e s o f C r o n a r t i u m comandrae.  x 2500.  F i g . Ik.  P y r i f o r m a e c i o s p o r e s o f C r o n a r t i u m comandrae showing s i z e and shape v a r i a t i o n o f t h e - s p o r e t a i l . X 700.  F i g . 15.  P y r i f o r m a e c i o s p o r e s o f C r o n a r t i u m comandrae showing the spore w a l l o r n a m e n t a t i o n . X 700.  - 28 -  on t h e i d e n t i c a l a r e a b y a t l e a s t one season. sues j u s t b e n e a t h t h e p e r i d e r m  (Adams 19195  The p y c n i a occur i n t i s -  K r e b i l l 1968a) and o v e r l i e  the a e c i a w h i c h d e v e l o p s e v e r a l c e l l s deeper i n t h e c o r t e x ( F i g . l l ) . H i r a t s u k a and Cummins (1963) found t h a t t h e C r o n a r t i u m  s p e c i e s were t h e  o n l y r u s t s t o d e v e l o p an i n t r a c o r t i c a l p y c n i a , and one o f t h e few r u s t g e n e r a i n w h i c h p y c n i a l growth was i n d e t e r m i n a t e . ion  u s u a l l y begins  Pycniospore  i n June, o r s h o r t l y a f t e r t h e main a e c i a l  productproduction  p e r i o d , and i n some seasons t h e y may c o n t i n u e t o be produced i n t o October. The p r i m o r d i a o f t h e a e c i a appear as a mass o f u n i n u c l e a t e h y p h a l c e l l s , o r i g i n a t i n g i n t h e c o r t i c a l parenchyma o f t h e h o s t some 3 t o 8 c e l l s below t h e p h e l l o g e n  (Adams 1919)-  The c e l l s o f t h e c e n t r a l  p o r t i o n o f t h e matured primorddfa become d i f f e r e n t i a t e d f i r s t t o f o r m b i nucleate b a s a l c e l l s .  Each o f t h e f i r s t b a s a l c e l l s formed c u t s o f f an  i n t e r c a l a r y c e l l and a c e l l w h i c h becomes metamorphosed t o f o r m a p e r i d i a l c e l l o f the c e n t r a l arch. continuous  The p e r i d i u m so formed c o n s i s t s o f a  membrane one t o t h r e e c e l l s t h i c k , and t h e aecium i s d e l i m i t e d  by t h e p e r i p h e r a l p e r i d i a l c h a i n s .  The b i n u c l e a t e b a s a l c e l l s cut o f f  chains o f c e l l s w h i c h f u r t h e r d i v i d e i n t o an i n t e r c a l a r y c e l l and an aeciospore.  The c e l l w a l l s o f t h e a e c i o s p o r e s b e g i n t o t h i c k e n when  t h e y a r e about t h e t h i r d o r f o u r t h spore from t h e b a s a l c e l l . spores t h e n become acuminate above (Adams 1919?  The a e c i o -  A r t h u r 193^) or below  ( A r t h u r and K e r n l^lk; Hedgcock and Long 1915a) t o g i v e a t y p i c a l p y r i form shape, a r e q u i t e v a r i a b l e i n s i z e , 17 t o 37 by 38 t o 77u,  with  c o l o r l e s s 2 t o 3u t h i c k w a l l s , w h i c h a r e f i n e l y v e r r u c o s e w i t h s m a l l i r r e g u l a r t u b e r c l e s ( F i g s . Ik and 1 5 ) .  As t h e a e c i a c o n t i n u e t o mature t h e  F i g , 16.  Canker o f C r o n a r t i u m comandrae on a young "branch o f P i n u s c o n t o r t a , w i t h the p e r i d i a o f t h e a e c i a s t i l l u n r u p t u r e d and f o r m i n g t y p i c a l b l i s t e r s p u s h i n g t h r o u g h t h e b a r k .  Fig.  17.  Canker o f C r o n a r t i u m comandrae w i t h abundant r u p t u r e d a e c i a c o v e r i n g the a e c i a l zone.  Fig.  18.  B r i g h t o r a n g e - c o l o r e d a e c i o s p o r e s o f C r o n a r t i u m comandrae c o v e r i n g t h e exposed a e c i a o f the canker.  - 29 -  - 30 -  o v e r l y i n g h o s t t i s s u e becomes r a i s e d and g r a d u a l l y r u p t u r e s e x p o s i n g t h e w h i t e b l i s t e r s c o n s i s t i n g o f t h e p e r i d i u m ( F i g . 16) e n c l o s i n g a powdery mass o f p y r i f o r m o r a n g e - y e l l o w  aeciospores ( F i g . 18).  The a e c i a ( F i g s .  17 and 18) a r e produced over t h e a r e a w h i c h p r e v i o u s l y produced p y c n i a , and occur over o r between p y c n i a l s c a r s , u s u a l l y i n t h e y e a r  following  p y c n i a l p r o d u c t i o n , a l t h o u g h a e c i a may n o t occur u n t i l t h e second year o r even l a t e r .  Aeciospore production v a r i e s widely with seasonal  d i t i o n s , l o c a l i t y , and b a r k t h i c k n e s s .  con-  I n t h e s t u d y a r e a t h e main p e r i o d  o f p r o d u c t i o n began i n l a t e May and c o n t i n u e d i n t o J u l y and up t o t h e end o f August a t a reduced r a t e .  I n some y e a r s t h e cankers may produce  a second crop o f a e c i o s p o r e s l a t e i n t h e season, b u t i n t h e c u r r e n t y e a r p y c n i a l zone.  There may be a g r e a t v a r i a t i o n between i n d i v i d u a l  a e c i a from one canker which can p r o l o n g t h e p r o d u c t i o n p e r i o d , b u t u s u a l l y p r o d u c t i o n from an i n d i v i d u a l aecium o n l y l a s t s f o r two t o f i v e weeks. F r e q u e n t l y t h e i n d i v i d u a l a e c i a a r e so c l o s e t o g e t h e r t h a t s e v e r a l app e a r t o c o a l e s c e t o form one l a r g e i r r e g u l a r aecium.  After the aecio-  spores a r e r e l e a s e d , t h e p e r i d i u m d i s i n t e g r a t e s , and b a r k n e c r o s i s o c curs around t h e a e c i a l r u p t u r e s .  Adams (1919) r e p o r t e d t h a t a new cork  l a y e r i s formed below t h e a e c i a , and t h e dead t i s s u e i s sloughed o f f o r remains as a rough, d e e p l y - c r a c k e d bark ( F i g s . 19, 20 and 2 l ) . (1968a) found no e v i d e n c e o f t h e development o f a secondary  Krebill  cork l a y e r  beneath a e c i a , b u t a l a y e r occurred a t times beneath p y c n i a . The mature a e c i o s p o r e s , w h i c h a r e g e n e r a l l y d i s p e r s e d b y t h e wind t o s u s c e p t i b l e Comandra o r Geocaulon p l a n t s , a r e b i n u c l e a t e ( F i g s . 22 and 2 3 ) .  Upon g e r m i n a t i n g one t o s i x germ tubes may emerge t h r o u g h  F i g . 19.  Young canker o f C r o n a r t i u m comandrae on P i n u s c o n t o r t a w i t h rough b a r k c o v e r i n g the o l d a e c i a l zone around t h e b r a n c h stub t h a t a c t e d as e n t r y p o i n t t o t h e stem. Abundant a e c i a were produced around the o l d a e c i a l zone.  F i g . 20.  S l i g h t h y p e r t r o p h y o f the young P i n u s c o n t o r t a stem caused by C r o n a r t i u m comandrae, w i t h c h a r a c t e r i s t i c rough, c r a c k e d b a r k , i n the o l d e r canker a r e a caused by a e c i a r u p t u r i n g .  F i g . 21.  T y p i c a l b a s a l stem canker o f C r o n a r t i u m comandrae showing o l d e r rough b a r k zone and c u r r e n t y e a r a e c i a l zone w i t h many d i s p e r s e d a e c i o s p o r e s caught i n t h e c r e v i c e s o f the b a r k .  - 32  the spore w a l l ( F i g . 2h),  -  but u s u a l l y the two n u c l e i m i g r a t e  o f the d e v e l o p i n g germ tubes ( F i g . 25) t o e l o n g a t e t o any e x t e n t . w h i c h may 26 and 27).  The  into  and o n l y t h i s germ tube  one  continues  germ tubes have m u l t i p l e s h o r t branches  occur i r r e g u l a r l y t h r o u g h o u t the l e n g t h o f the germ tube The  germ tubes are non-septate,  w i t h the two n u c l e i c o n t i n u -  i n g t o m i g r a t e toward the t i p o f the tube d u r i n g e l o n g a t i o n . t i o n s are s u i t a b l e an a p p r e s s o r i u m f e c t i o n peg may  develop  forms ( F i g . 28)  ( F i g s . 29 and 30)  the stomata t o i n f e c t and develop  (Figs.  I f condi-  from w h i c h an i n -  and p e n e t r a t e the h o s t  a b i n u c l e a t e mycelium.  through  T h i s type  of  a e c i o s p o r e g e r m i n a t i o n i s t y p i c a l o f the h e t e r o e c i o u s r u s t s , and i s i n marked c o n t r a s t t o t h a t o f the a u t o e c i o u s r u s t s ( H i r a t s u k a 1968; suka e t a l . 1966;  P o w e l l and Morf 1966).  The  Hirat-  stem, l e a f p e t i o l e s ,  f l o r a l b r a c t s and f r u i t s o f the a l t e r n a t e h o s t may  leaves,  a l s o be i n f e c t e d .  One  t o t h r e e weeks a f t e r a e c i o s p o r e i n f e c t i o n , the u r e d i a appear on the s u r f a c e o f the l e a v e s as s m a l l y e l l o w p u s t u l e s ( F i g . 31)•  The u r e d i a de-  v e l o p f r o m a h y p h a l p l e x u s o f t e n b e n e a t h a stoma o f the h o s t l e a f (Moss I928).  The v e r t i c a l hyphae a r i s e i n a p a l i s a d e f a s h i o n and become d i -  v i d e d i n t o t h r e e c e l l s ; the p e r i d i a l , i n t e r c a l a r y and sporogenous  cells.  A t a l a t e r stage the i n t e r c a l a r y c e l l s d i s o r g a n i z e , the b a s a l sporogenous c e l l s g i v e r i s e t o s p o r e - i n i t i a l s , and the p e r i d i a l c e l l s g i v e r i s e t o the p e r i d i u m o f the u r e d i a l s o r u s .  The p e r i d i u m o f the u r e d i a l  sorus  q u i c k l y r u p t u r e s e x p o s i n g a, mass o f e l l i p t i c a l t o g l o b o i d ,  orange-yellow  u r e d i o s p o r e s , 16 t o 23 by 19~28u, w i t h n e a r l y c o l o r l e s s 1.5  t o 2\x t h i c k  w a l l s , which are s p a r s e l y e c h i n u l a t e .  easily  The u r e d i o s p o r e s  are  d i s s e m i n a t e d by wind or i n s e c t s and are a b l e t o germinate and  re-infect  F i g . 22.  A e c i o s p o r e s o f C r o n a r t i u m comandrae, s t a i n e d w i t h HC1Giemsa, each w i t h two n u c l e i . X 700.  F i g . 23.  A e c i o s p o r e o f C r o n a r t i u m comandrae, s t a i n e d w i t h HC1C-iemsa, w i t h two n u c l e i . X 1200.  F i g . 2k.  I n i t i a t i o n o f a e c i o s p o r e g e r m i n a t i o n , w i t h f o u r germ tubes d e v e l o p i n g t h r o u g h t h e spore w a l l , a f t e r 1 hour. X 1200.  F i g . 25.  C r o n a r t i u m comandrae a e c i o s p o r e w i t h s e v e r a l germ t u b e s , b u t o n l y one germ tube w e l l developed a f t e r 6 hours. Note t h e s h o r t m u l t i p l e branches on t h e n o n - s e p t a t e germ t u b e . X 380.  Fig.  26.  Group o f germinated a e c i o s p o r e s w i t h t y p i c a l b r a n c h i n g o f germ tubes a f t e r 2k h o u r s . X 125.  - 33 -  F i g . 27.  A e c i o s p o r e o f C r o n a r t i u m comandrae w i t h one developed germ tube, a f t e r 3 hours. Note t h e n o n - s e p t a t e c o n d i t i o n o f t h e germ tube and t h a t t h e two n u c l e i have m i g r a t e d towards t h e t i p o f t h e germ tube. X 830.  F i g . 28.  Germ tubes o f two C r o n a r t i u m comandrae a e c i o s p o r e s a f t e r 3 hours, showing development o f a g l o b o s e and club-shaped a p p r e s s o r i u m on t h e developed germ tube, i n t o w h i c h t h e two n u c l e i have m i g r a t e d . X 700.  F i g . 29.  Germ tube development o f a C r o n a r t i u m comandrae a e c i o spore a f t e r 3 hours, showing a globose a p p r e s s o r i u m w i t h two n u c l e i and an i n f e c t i o n peg. X 1080.  Fig.  30.  Germ tube development o f two C r o n a r t i u m comandrae a e c i o spores a f t e r 3 hours, showing two t y p e s o f a p p r e s s o r i a each w i t h two n u c l e i and an i n f e c t i o n peg. X 6l0.  F i g . 31.  U r e d i a o f C r o n a r t i u m comandrae on c e n t r a l p o r t i o n o f a Comandra u m b e l l a t a s s p . p a l l i d a , l e a f . Dark s t r u c t u r e s on b o t t o m a r e young t e l i a .  F i g . 32.  T e l i a o f C r o n a r t i u m comandrae on b o t h s u r f a c e s o f a p o r t i o n o f Comandra u m b e l l a t a s s p . p a l l i d a l e a f .  - 35 -  - 36 -  t h e same a l t e r n a t e h o s t , thus s p r e a d i n g  and  i n t e n s i f y i n g the  infection  d u r i n g the summer, o f t e n over a d i s t a n c e o f a hundred m i l e s or more. I n the p r e s e n t  study u r e d i a l i n f e c t i o n s were found hundreds o f m i l e s  out on the P r a i r i e s away from any p i n e s o u r c e .  A l l p a r t s o f Comandra  p l a n t s were h i g h l y s u s c e p t i b l e t o i n f e c t i o n at any stage i n t h e i r d e v e l o p ment.  Four t o seven g e n e r a t i o n s  o f the u r e d i a l s t a t e may  be produced i n  a s i n g l e season, i f weather c o n d i t i o n s p e r m i t , w h i c h g r e a t l y i n c r e a s e s the p o s s i b i l i t i e s o f p i n e T e l i a may  form about two t o t h r e e weeks a f t e r the u r e d i a  pear on the p l a n t s and may new  infection.  o c c u r on l e s i o n s i n i t i a t e d by u r e d i a or  l e s i o n s l a t e i n t h e season.  The  t e l i a are r e d d i s h - b r o w n ,  h a i r - l i k e s t r u c t u r e s , 80 t o 120u  o f t e n curved,  slender,  t o 16 by 28 t o kk\i,  with i n a gela-  I n t i m e , u s u a l l y under m o i s t c o n d i t i o n s , each t e l i o s p o r e  g e r m i n a t e s i n s i t u by means o f a f i v e - c e l l e d promycelium, each o f f o u r upper c e l l s b e a r i n g one  the 2  s m a l l t h i n - w a l l e d globose b a s i d i o s p o r e ,  t o 6 by 2 t o 6\i, on the t i p o f a s t o u t s t e r i g m a . disseminated  in  They are com-  n e a r l y c o l o r l e s s , 2 t o 3M- t h i c k , smooth w a l l s , h e l d t o g e t h e r tinous matrix.  on  t h i c k and about 1 mm  l e n g t h , w h i c h stand up from the h o s t t i s s u e ( F i g . 32). posed o f columnar rows o f t e l i o s p o r e s , 12  ap-  by the wind, and  i f d e p o s i t e d on s u s c e p t i b l e p i n e  w h i l e s t i l l v i a b l e , can g e r m i n a t e under f a v o u r a b l e f e c t the p i n e , thus c o m p l e t i n g  The b a s i d i o s p o r e s  are  needles  c o n d i t i o n s and i n -  the l i f e c y c l e o f the  fungus.  - 37 -  SYMPTOMS The f i r s t v i s i b l e s i g n o f comandra b l i s t e r r u s t i n f e c t i o n on a p i n e i s a y e l l o w - b r o w n spot on a n e e d l e .  This i s at the p o i n t of i n -  f e c t i o n , b u t i t i s d i f f i c u l t t o d i s t i n g u i s h m a c r o s c o p i c a l l y from s p o t s caused by o t h e r a g e n c i e s .  I n the f i r s t ,  second o r t h i r d  similar  season  f o l l o w i n g i n f e c t i o n an a r e a o f t h e bark a t the base o f t h e n e e d l e  fasc-  i c l e becomes d i s c o l o r e d , and a s m a l l s p i n d l e - s h a p e d s w e l l i n g may d e v e l o p . T h i s i s t h e b e g i n n i n g o f a canker, w h i c h m o s t l y b e g i n s on n e e d l e - b e a r i n g branches and stems.  I n f e c t i o n s on main stems u s u a l l y e n t e r v i a s m a l l  ( F i g . 33).  M y c e l i a l growth i n t h e h o s t t i s s u e i s more r a p i d  branches  l o n g i t u d i n a l l y t h a n l a t e r a l l y , and more r a p i d towards t h e base t h a n t h e t i p o f t h e b r a n c h o r stem. Reddish-orange years a f t e r i n f e c t i o n .  p y c n i a l drops appear on t h e b a r k two o r more I n subsequent  y e a r s t h e p y c n i a form i n a zone  b e h i n d t h e p e r i p h e r y o f t h e i n f e c t e d t i s s u e and o u t s i d e t h e a e c i a l  zone.  P y c n i a l drops a r e not v i s i b l e l o n g ; t h e y may be e a t e n by l a r v a l and a d u l t i n s e c t s , d r y up, o r be washed b y r a i n .  " P y c n i a l s c a r s " may s t i l l be seen  i f t h e b a r k i s r e l a t i v e l y smooth. A e c i a appear as w h i t e b l i s t e r s p u s h i n g t h r o u g h the b a r k over p y c n i a l zones o f t h e p r e v i o u s y e a r s . ing  masses o f orange a e c i o s p o r e s .  The a e c i a l p e r i d i a r u p t u r e r e l e a s -  A e c i a a r e o n l y p r e s e n t i n one a r e a f o r  one season as t h e p y c n i a l and a e c i a l zones c o n t i n u a l l y form around t h e p e r i p h e r y o f t h e canker. bark.  Not a l l a e c i a appear as b l i s t e r s t h r o u g h t h e  I n t h i c k - b a r k e d t r e e s t h e a e c i a a r e o f t e n b u r i e d and t h e a e c i o s p o r e s  F i g . 33-  Branch i n f e c t i o n o f C r o n a r t i u m comandrae on P i n u s c o n t o r t a , w i t h mycelium c a u s i n g s w e l l i n g i n the main stem around the b r a n c h s t u b .  Fig,. 35-.  Heavy r e s i n o s i s f o r m i n g a d r i e d c r u s t over a p o r t i o n o f a C r o n a r t i u m comandrae stem canker. Note rough b a r k o f o l d b r a n c h canker (upper l e f t ) w h i c h s e r v e d as e n t r y f o r the r u s t t o the stem.  F i g . 3^.  Tree w i t h dead s p i k e - t o p 'above a canker o f C r o n a r t i u m comandrae g i r d l i n g the t r e e ; lower branches a r e p r o g r e s s i v e l y k i l l e d by downward growth o f r u s t .  - 39 -  appear t h r o u g h c r a c k s i n t h e b a r k . a e c i a w i t h a e c i o s p o r e s a r e exposed.  On removing t h e b a r k l a r g e areas o f Under n a t u r a l c o n d i t i o n s o n l y a  s m a l l p r o p o r t i o n o f these a e c i o s p o r e s would be d i s p e r s e d .  Some t r e e s ,  e s p e c i a l l y t h o s e w i t h marked r e s i n o s i s , seldom produce a e c i o s p o r e s , and i t i s d i f f i c u l t t o e s t a b l i s h t h e l i m i t s o f t h e canker m i c r o s c o p i c a l l y . Where a e c i a have been produced, t h e b a r k c r a c k s and t h e cambium and u n d e r l y i n g wood d i e .  As t h e fungus grows t h e b r a n c h o r stem  t u a l l y i s g i r d l e d and p a r t s beyond t h e canker d i e .  even-  Spike-topped trees  3*0 and f l a g g e d branches a r e conspicuous symptoms o f t h e d i s e a s e .  (Fig.  When cankers d e v e l o p on t h e main stem, t h e canker zone i s o c c a s i o n a l l y c o n s t r i c t e d i n s t e a d o f s w o l l e n , as i n c r e a s e d d i a m e t e r growth o f t h e h e a l t h y stem c o n t i n u e s above and below t h e canker d u r i n g canker g i r d ling.  R e s i n e x u d a t i o n i s a conspicuous symptom o f d i s e a s e , e s p e c i a l l y  on P i n u s c o n t o r t a ( F i g . 35).  Much o f t h e r e s i n o s i s i s caused by t h e  fungus b u t some i s caused b y r o d e n t s chewing t h e s u c c u l e n t i n f e c t e d b a r k of  t h e canker.  Rodent damage i s u s u a l l y a good i n d i c a t i o n o f t h e p r e -  sence o f t h e d i s e a s e .  ( F i g s . 36 and 37).  The f i r s t symptom o f t h e r u s t on t h e a l t e r n a t e h o s t i s t h e y e l low s w e l l i n g s o f t h e d e v e l o p i n g u r e d i a l s o r i , w h i c h r u p t u r e t o r e l e a s e the  u r e d i o s p o r e s . A few weeks l a t e r t h e t e l i a l "horns" a r e e a s i l y v i s i -  ble  d e v e l o p i n g on t h e same a r e a as t h e u r e d i a l s o r i o r from new l e s i o n s .  U r e d i a and t e l i a may be p r e s e n t over t h e whole p l a n t .  There i s some  c h l o r o s i s o f the leaves i n h e a v i l y i n f e c t e d p l a n t s , s l i g h t evidence o f reduced l e a f growth and d e f o l i a t i o n i s u s u a l l y premature.  F i g . 36.  A n n u a l r o d e n t damage on a l a r g e C r o n a r t i u m comandrae stem canker on P i n u s c o n t o r t a . Note t h e s t r i p o f d r i e d dead b a r k n o t removed on each a n n u a l v i s i t , and t h e abundant e x u d a t i o n o f r e s i n .  F i g . 37.  Rodent damage on a C r o n a r t i u m comandrae stem canker around a b r a n c h w h i c h a c t e d as an e n t r y p o i n t f o r t h e r u s t i n t o t h e stem. S p o r u l a t i o n o f t h e a e c i a l zone can be seen o u t s i d e t h e chewed a r e a .  - ko -  - 1+1 -  DAMAGE  Comandra b l i s t e r r u s t damage i n n a t u r a l p i n e stands i s n o t u s u a l l y s p e c t a c u l a r , i n f e c t i o n b e i n g l i m i t e d t o some e x t e n t b y t h e d i s t r i b u t i o n o f i t s a l t e r n a t e h o s t s , which, a l t h o u g h r a n g i n g over much o f N o r t h America,  are r e s t r i c t e d l o c a l l y t o s m a l l areas.  Comandra umbel-  l a t a i s r a r e l y found w i t h i n p i n e stands as i t f a v o u r s d r y open a r e a s , b u t Geocaulon l i v i d u m o c c u r s on w e t t e r areas w i t h i n s p r u c e - p i n e s t a n d s . Heavy p i n e i n f e c t i o n i s u s u a l l y found a d j a c e n t t o areas o f i n f e c t e d a l ternate hosts.  The r u s t a t t a c k s t r e e s o f a l l ages and s i z e s and causes  m o r t a l i t y by b a s a l o r stem g i r d l i n g .  The p e r i o d between i n i t i a l  infec-  t i o n and d e a t h o f t h e t r e e may be as much.as t h i r t y y e a r s , o r more, and t h e n o n l y a few t r e e s d i e each y e a r .  S e e d l i n g s may be k i l l e d w i t h i n a  few y e a r s f o l l o w i n g i n f e c t i o n , because t h e r u s t can e n t e r t h r o u g h on t h e main stem and t h e fungus soon g i r d l e s t h e s m a l l stem.  needles  The fungus  c h a r a c t e r i s t i c a l l y e n t e r s t h e main stem o f t h e o l d e r t r e e v i a b r a n c h i n fections.  The fungus o n l y grows one to. t h r e e i n c h e s down t h e b r a n c h  each y e a r so t h a t t h e rust.may be p r e s e n t i n t h e b r a n c h many y e a r s bef o r e i t reaches t h e main stem.  I f the branch i s k i l l e d before the r u s t  reaches t h e main stem, t h e main stem w i l l not u s u a l l y become i n f e c t e d . Dead tops o r s p i k e - t o p s were common damage f e a t u r e s i n mature stands where t h e fungus had g i r d l e d t h e stem t h e r e b y k i l l i n g t h e upper stem and branches. C h i l d s (1968) showed t h a t downward growth on t h e stem by t h e r u s t i s l e s s t h a n 6 i n c h e s p e r y e a r , thus t h e lower p o r t i o n o f t h e t r e e may l i v e f o r many y e a r s i f v i g o r o u s lower branches  remain.  I n f e c t i o n w i t h i n a stand  may have o c c u r r e d i n o n l y a few y e a r s , o r even i n one y e a r , when c o n d i t i o n s  - h2 -  f o r i n f e c t i o n were f a v o u r a b l e and a s u p p l y o f b a s i d i o s p o r e s was a v a i l able. S e v e r a l r e p o r t s show t h a t p i n e i n f e c t i o n b y : t h e r u s t i s i n frequent.  Observations  were made f o r over 50 y e a r s i n an i n f e c t e d P i n u s  p o n d e r o s a stand i n S i s k i y o u County, C a l i f o r n i a , and i t appeared t h a t c u r r e n t damage was f r o m i n f e c t i o n kO y e a r s ago w i t h l i t t l e o r no r e c e n t i n f e c t i o n ( C a l i f o r n i a F o r e s t P e s t C o n t r o l A c t i o n C o u n c i l 1 9 5 8 ; U. S. F o r e s t S e r v i c e 1 9 5 5 ; Wagener i 9 6 0 ) .  I n f e c t i o n o f P. p o n d e r o s a i n s o u t h -  e r n Nevada was r e p o r t e d as s p o r a d i c b y Wagener ( i 9 6 0 ) w i t h o f i n f e c t i o n b y t h e r u s t i n .1932 o r 1 9 3 3 " and f e c t i o n on p i n e s i n c e  a wave  almost no new i n -  Wagener ( 1 9 5 0 ) r e p o r t e d t h e r u s t on P.  ponderosa i n a southeastern  Idaho p l a n t a t i o n " . . . . c h i e f l y on t h e 19^0  and 19^1 growth o f branches  and no e v i d e n c e was found i n I 9 6 2 t o  i n d i c a t e any more r e c e n t i n f e c t i o n ( K r e b i l l , p e r s o n a l communication 1 9 6 2 ) . L i t t l e o r no r e c e n t i n f e c t i o n was found on P. p o n d e r o s a i n t h e B l a c k H i l l s , South Dakota, and areas o f C o l o r a d o i n stands where damage was reported 25-35 years p r e v i o u s l y (Peterson 1962a).  Peterson  (1962a)  pre-  sented e v i d e n c e t h a t i n f e c t i o n o c c u r r e d two t o s e v e r a l decades ago on P. c o n t o r t a i n t h e B i g h o r n and Shoshone N a t i o n a l F o r e s t s , Wyoming, w i t h almost no r e c e n t One  infection.  o f t h e f i r s t r e p o r t s o f damage i n t h e U n i t e d S t a t e s was  made b y Boyce (Hedgcock and Long 1 9 1 5 a ) i n Klamath N a t i o n a l F o r e s t ,  Cali-  f o r n i a , where he found 34% o f t h e young P. p o n d e r o s a dead and a f u r t h e r 17% i n f e c t e d i n a stand r e p r e s e n t i n g average r a t h e r t h a n heavy i n f e c t i o n . Meinecke ( 1 9 2 8 ) r e p o r t e d 35% o f P. p o n d e r o s a dead o r doomed i n an a r e a  - k3 -  near Mt. Lassen,  C a l i f o r n i a , and c a l c u l a t e d t h a t t h i s r e s u l t e d i n a 21$  r e d u c t i o n o f f o r e s t cover.  Other r e p o r t s o f damage t o P. ponderosa have  been noted from o t h e r p a r t s o f C a l i f o r n i a (U.S. F o r e s t S e r v i c e 195'5, 1956), a 30-year o l d p l a n t a t i o n i n C a r i b o u N a t i o n a l F o r e s t , Idaho (Wegener 1950), C h a r l e s t o n Mountains, s o u t h e r n Nevada (Wegener i 9 6 0 ) ,  the Black  H i l l s o f South D a k o t a (Hedgcock and Long 1915a; L u c k i n b i l l 1935)> and t h e e a s t s i d e o f t h e Cascade Range i n Oregon and Washington ( C h i l d s 1968). Damage t o young P. pungens i n P e n n s y l v a n i a was noted b y Adams and O r t o n (191^) and Hedgcock and Long (1915a); t h e l a t t e r r e p o r t e d t h a t 58$ o f the p i n e w h i c h were p r o d u c i n g  a e c i a a t t h e end o f June were dead b y  autumn from t h e g i r d l i n g e f f e c t o f t h e fungus. U n t i l r e c e n t l y , P. c o n t o r t a was c o n s i d e r e d o n l y an o c c a s i o n a l h o s t f o r t h i s pathogen ( M i e l k e 1957) a l t h o u g h P e t e r s o n unpublished  (1962a) quotes an  I925 r e p o r t b y B e t h e l o f damage t o t h i s h o s t i n C o l o r a d o .  M i e l k e (1957) r e p o r t e d mature stands  c o v e r i n g s e v e r a l square m i l e s where  50 - 98$ o f t h e t r e e s were i n f e c t e d i n areas o f Idaho, U t a h and Wyoming. Andrews and H a r r i s o n (1959) a l s o r e p o r t e d s e v e r e l y i n f e c t e d P. c o n t o r t a stands i n Wyoming a d j a c e n t t o sagebrush areas s u p p o r t i n g t h e a l t e r n a t e host.  Peterson  (1962a) s t u d i e d damage on 2k p l o t s i n i n f e c t e d stands i n  n o r t h e r n Wyoming and found 21$ o f t h e t r e e s i n f e c t e d .  The stands were  l a r g e l y 50 - 100 y e a r s o l d w i t h i n f e c t i o n c e n t e r e d i n t i s s u e u s u a l l y over 20 y e a r s o l d , w i t h t h e cankers  a t an average h e i g h t o f 12 f e e t .  K r e b i l l (1965) s t u d i e d i n f e c t e d stands i n 12 N a t i o n a l F o r e s t s l o c a t e d from c e n t r a l Montana t o n o r t h e r n U t a h and found t h a t some cankers were 100 y e a r s o l d , and t h a t t h e r e had been a b u i l d up between 1910 t o 19^5,  - 44 -  w i t h o n l y sparse r e c e n t i n f e c t i o n .  I n most cases over 90% of- the sampled  cankers were i n t r u n k s and from o n e - f o u r t h caused s p i k e t o p s .  t o o n e - h a l f o f these had  The c e n t e r s o f cankers ranged from 1.5 t o 69 f e e t  above ground w i t h the mean h e i g h t around 26 f e e t . cankers had been s c a r r e d b y r o d e n t chewing.  About 90% o f the  I n one sample p l o t 26% o f  a l l l i v e l o d g e p o l e p i n e s were i n f e c t e d , and t h e r e was c o n s i d e r a b l e  evi-  dence o f r e c e n t m o r t a l i t y b y r u s t g i r d l i n g . R e c e n t l y C. comandrae has become a problem i n young p l a n t a t i o n s o f P. ponderosa and P. t a e d a i n the Cumberland P l a t e a u a r e a o f e a s t e r n Tennessee and n o r t h e a s t e r n Alabama (Powers e t a l . 1967).  Powers e t a l .  r e p o r t p l a n t a t i o n s w i t h over 9°% o f the l e s s t h a n 10 y e a r - o l d t r e e s i n f e c t e d , and one case, w i t h 57% o f the 2 y e a r - o l d P. t a e d a t r e e s i n f e c t e d . More r e c e n t l y , C o r d e l l e t a l . (1967) r e p o r t e d t h a t 40% o f the P. t a e d a p l a n t a t i o n s l e s s t h a n 10 y e a r s o l d were i n f e c t e d on the Cumberland P l a t e a u , and on these p l a n t a t i o n s 6% o f the t r e e s were i n f e c t e d . also detected  They  severe i n f e c t i o n and m o r t a l i t y i n a P. e c h i n a t a p l a n t a t i o n  i n the same a r e a .  C. comandrae i s a l s o p r o v i n g a p l a n t a t i o n p r o b l e m on  P. e c h i n a t a i n the C l a r k N a t i o n a l F o r e s t , M i s s o u r i ( B e r r y e t a l . 1961), and Ozark N a t i o n a l F o r e s t , Arkansas ( D o o l i n g e t a l . 1964).  I n the  latter  a r e a 44% o f the t r e e s on t h r e e p l o t s p l a n t e d i n 1958 i n the B u f f a l o D i s t r i c t were dead b y 1967 ( D o o l i n g 1967).  Wolfe e t a l . (1968) r e c e n t l y  found t h a t the r u s t was now p r e s e n t on a l l 8 d i s t r i c t s o f the Ozark N a t i o n a l F o r e s t , b e i n g observed i n 19% o f the P. e c h i n a t a stands examined. I n Canada t h e r e have been few r e c o r d e d r e p o r t s o f e x t e n s i v e damage, a l t h o u g h the r u s t has been r e p o r t e d as f a i r l y common t h r o u g h o u t  - 45 -  the P r a i r i e P r o v i n c e s .  I n 1959 two a r e a r e p o r t s o f m o r t a l i t y were r e -  corded i n P. c o n t o r t a stands:  one i n young t r e e s near Robb, A l b e r t a  (Thomas e t a l . i 9 6 0 ) , and the o t h e r i n the T e s l i n - W h i t e h o r s e a r e a o f the Yukon w i t h 5 - 10$ o f the s a p l i n g s i z e t r e e s i n f e c t e d (Molnar i 9 6 0 ) . L a t e r , Baranyay and Stevenson (1964) r e p o r t e d  3$ o f the l i v i n g t r e e s on  seven 0.05-acre p l o t s i n 20 y e a r - o l d r e g e n e r a t i o n  near Robb were i n f e c t e d  by C. comandrae, the range between p l o t s v a r y i n g from 1.4 t o 10.8$. eastern Bay,  In  Canada, 22$ o f the t r e e s i n a P. b a n k s i a n a p l a n t a t i o n a t Thunder  Ontario,  were i n f e c t e d (Dance and Lynn 1965), and more t h a n 10$ o f  young P. b a n k s i a n a i n a s m a l l a r e a a t S a i n t U r b a i n , Quebec, were i n f e c t e d ( M a r t i n e a u and O u e l l e t t e 1966). Meinecke (1928), H o r t o n (1955), and M i e l k e e t a l . (1968) ment i o n the v a l u e o f the r u s t as a b i o t i c t h i n n i n g agent c a u s i n g m o r t a l i t y i n o v e r s t o c k e d young p i n e s t a n d s .  P e t e r s o n (1962a), however, s t a t e s  t h a t m o r t a l i t y caused b y the r u s t " c o u l d seldom o r never be w r i t t e n o f f as harmless o r b e n e f i c i a l n a t u r a l t h i n n i n g " .  He found t h a t the  infected  t r e e s were l a r g e r , and t h a t suppressed t r e e s d i d not respond t o r e l e a s e by the r u s t .  He a l s o found t h a t over h a l f the i n f e c t e d l i v e  lodgepole  p i n e t r e e s had a lower v i g o r r a t i n g t h a n t h e y would have i f r u s t - f r e e . C h i l d s (1968) found t h a t t o p - k i l l i n g o f P. p o n d e r o s a b y the r u s t was n o t i n d i c a t i v e o f h i g h - r i s k t r e e s , u n l e s s the r e m a i n i n g crown was s m a l l and o f poor v i g o r .  N o r d i n (195*0 r e p o r t e d  the r u s t was more p r e v a l e n t i n  pure t h a n i n mixed stands i n A l b e r t a . Damage a p p r a i s a l surveys c a r r i e d out i n a s s o c i a t i o n w i t h  the  p r e s e n t s t u d i e s , i n d i c a t e d t h a t 44$ o f t t h e 725 t r e e s on a 0.1 a c r e p l o t  - k6 -  i n a 20 y e a r - o l d P. b a n k s i a n a s t a n d s o u t h o f Rae, N.W.T., were i n f e c t e d i n I965 w i t h an average o f 1.6 i n f e c t i o n s p e r tree;..  By 1966, 7.9% o f  the i n f e c t e d t r e e s had d i e d and many o t h e r s had dead tops ( J . A. Baranyay, u n p u b l i s h e d d a t a I965, I966).  N e a r l y a t h i r d o f t h e cankers noted i n  I965 were i n a c t i v e i n 1966, b u t many new a c t i v e cankers were r e c o r d e d . A 1966 s u r v e y on a 0 . 1 a c r e p l o t i n a 25 y e a r - o l d P. c o n t o r t a s t a n d near Saskatchewan  R i v e r C r o s s i n g , B a n f f N a t i o n a l Park, A l b e r t a ,  indicated  32.7% o f t h e t r e e s i n f e c t e d b y t h e r u s t , Peridermium s t a l a c t i f o r m e A r t h . . & K e r n , and 10% b y C. comandrae ( J . A. Baranyay, u n p u b l i s h e d d a t a 1966). Over h a l f t h e C_. comandrae i n f e c t e d t r e e s were dead b y 1968.  A I92U  p l a n t a t i o n o f P i n u s s y l v e s t r i s near Beaver Mines, A l b e r t a , was r e p o r t e d t o have e x t e n s i v e r o d e n t damage, some w i t h f r e s h chewing which l e a d me t o suspect a r u s t i n f e c t i o n .  A f t e r c o n s i d e r a b l e s e a r c h , one s t r i p o f a c t i v e  canker w i t h spores was found a t t h e edge o f a chewed a r e a ( P o w e l l and Morf 1965).  Of 50 t r e e s t a l l i e d a l l had r o d e n t damage, s u g g e s t i n g r u s t  i n f e c t i o n was once w i d e s p r e a d throughout t h e p l a n t a t i o n .  Heavilyi n -  f e c t e d C. u m b e l l a t a ssp. p a l l i d a grew w i t h i n a few f e e t i n an open meadow. About 500 cankers were measured and observed on t r e e s v a r y i n g i n age from 3 t o 125 y e a r s from 23 areas i n t h e Rocky Mountains and F o o t h i l l s regions o f Alberta.  The average h e i g h t o f t h e c e n t e r o f t h e can-  k e r s was l.k f e e t , w i t h t h e h i g h e s t c e n t e r e d a t 15 f e e t .  The average  l e n g t h o f t h e cankers was 1.2 f e e t , w i t h t h e l o n g e s t 15 f e e t .  I n 1966  and 1967, kkO l i v e cankers were tagged (103 i n 1967), b u t b y 1968 101 o f o f t h e s e were dead, o f w h i c h o n l y 13 were b r a n c h c a n k e r s .  Canker  mortality  at one l o c a t i o n amounted t o 82%, and a t another 55% b o t h o c c u r r e d i n 10 5  - kl -  t o 20 y e a r - o l d s t a n d s .  I n a regeneration  a r e a where few t r e e s were over  5 y e a r s o l d , kk% o f the t r e e s w i t h cankers d i e d i n one y e a r . i n f e c t i o n i n t h i s a r e a amounted t o 23% on a 0.22  Total tree  acre p l o t i n 19°7 b u t 5  i t was u n l i k e l y t h a t any o f t h e i n f e c t e d t r e e s would s u r v i v e f o r more than a few y e a r s a t t h e c u r r e n t m o r t a l i t y r a t e .  The r e g e n e r a t i o n o c -  c u r r e d i n an a r e a o f Comandra and chances f o r f u r t h e r i n f e c t i o n i n t h e s e young age t r e e s were h i g h .  - hd -  AREA OF STUDY  LOCATION, PHYSIOGRAPHY AND GEOLOGY The main a r e a o f study was  r e s t r i c t e d t o a p o r t i o n o f the  K a n a n a s k i s R i v e r v a l l e y c e n t e r e d on the K a n a n a s k i s F o r e s t E x p e r i m e n t S t a t i o n some 5 m i l e s south o f Seebe, A l b e r t a ( F i g . 38), where the f o o t h i l l s merge w i t h the f r o n t range o f the Rocky Mountains.  The  Kananaskis  F o r e s t Experiment S t a t i o n , an a r e a o f about 2k square m i l e s , i s bounded t o the n o r t h b y the Stoney I n d i a n R e s e r v a t i o n , and on a l l o t h e r s i d e s b o r d e r s on the Bow R i v e r F o r e s t , o f the A l b e r t a Rocky Mountains F o r e s t Reserve.  The K a n a n a s k i s R i v e r , w h i c h f l o w s n o r t h t o j o i n the Bow R i v e r  a t Seebe, l i e s i n a n o r t h - s o u t h v a l l e y s i t u a t e d between two o f the e a s t e r n s e r i e s o f ranges o f the Rocky M o u n t a i n s .  I n the n o r t h e r n p a r t o f  the v a l l e y the K a n a n a s k i s Range l i e s t o the west and the F i s h e r Range t o the e a s t .  The a l t i t u d e o f the a r e a v a r i e s between U,200 and 9 500 5  f e e t , w i t h t i m b e r l i n e between 6,500 and 7 , 0 0 0 f e e t .  B a r r i e r Lake i s  a man-made water body over f o u r m i l e s l o n g , f i l l i n g a p o r t i o n o f the  flat-  bottomed U-shaped v a l l e y between the n o r t h e r n end o f the K a n a n a s k i s F o r e s t E x p e r i m e n t S t a t i o n and a p o i n t about a m i l e south o f the Evans Thomas Creek.  The v a l l e y bottom, w h i c h a t some p o i n t s i s !§• m i l e s wide,  i s f l a n k e d at many p l a c e s b y t e r r a c e s o f g l a c i a l t i l l and m o r a i n i c mat e r i a l , and l a c u s t r i n e and a l l u v i a l d e p o s i t s ; evidence g l a c i a t i o n s and r e t r e a t s t o w h i c h the a r e a was  of previous  subjected.  From a p p r o x i -  m a t e l y the 5,000 f o o t l e v e l the s l o p e s r i s e g r a d u a l l y o r p r e c i p i t o u s l y t o masses o f u p t h r u s t and f o l d e d sedimentary  rocks deposited  from  Fig.  38-  Map o f t h e l o w e r K a n a n a s k i s R i v e r V a l l e y , A l b e r t a , i n g the l o c a t i o n s o f the main study areas.  show-  - k9 -  - 50 -  Cambrian t o Cretaceous t i m e s , w h i c h form the mountains and r i d g e tops o f the ranges.  The g e o l o g i c a l f o r m a t i o n s , o f the a r e a were s t u d i e d b y B a w l i n g  (1905) and Beach (19U3).  SOILS  A p r e l i m i n a r y s u r v e y o f the s o i l s o f t h e a r e a was made b y C r o s s l e y (1951), who found w e l l d r a i n e d a l l u v i a l s o i l s and eroded  soil  and r o c k areas dominant i n the f l o o d p l a i n s o f the K a n a n a s k i s R i v e r and i t s major e a s t bank t r i b u t a r i e s .  S h a l l o w stomy/ l i t h o s o l s o i l s are p r o -  minent on moderate t o steep s l o p e s where e r o s i o n i s q u i t e r a p i d .  The  dominant s o i l s on the lower s l o p e s o f the v a l l e y are g r e y p o d z o l s o f c a l c a r e o u s o r n o n - c a l c a r e o u s o r i g i n , w i t h brown p o d z o l s and p o d z o l s gene r a l l y a t higher elevations.  L o c a l i z e d areas s u p p o r t hangmoor p e a t ,  half-  bog and sod s o i l s , which r e s u l t from e x c e s s i v e m o i s t u r e , poor d r a i n a g e , or s o i l s a t h i g h a l t i t u d e s under g r a s s o r open f o r e s t .  Brown f o r e s t  s o i l s , and l o c a l i z e d chernozem and r e n d z i n a s o i l s are found around  the  n o r t h e r n end o f B a r r i e r Lake and t o the n o r t h , where the F o o t h i l l s  abut  a g a i n s t the Rocky Mountains.  A f o r e s t l a n d c l a s s i f i c a t i o n map,  s u r f i c i a l m a t e r i a l , was r e c e n t l y p r e p a r e d f o r some o f the R e s e a r c h F o r e s t ( D u f f y and England  based on  Kananaskis  I967), and a g e n e r a l i z e d s o i l s map  was p r e p a r e d f o r t h e Marmot Creek b a s i n ( J e f f r e y 1965).  CLIMATE The main c l i m a t i c c h a r a c t e r i s t i c o f the K a n a n a s k i s V a l l e y i s  - 51  -  i t s v a r i a b i l i t y , t y p i c a l o f most mountain r e g i o n s i n c o n t i n e n t a l l o c a t i o n s . T h i s v a r i a b i l i t y i s o n l y now becoming more a c c u r a t e l y d e t e r m i n e d t h r o u g h the i n c r e a s e i n the number o f c l i m a t o l o g i c a l s t a t i o n s i n the v a l l e y ,  and  e s p e c i a l l y t h r o u g h the i n t e n s i v e i n s t r u m e n t a t i o n a s s o c i a t e d w i t h the A l b e r t a Watershed R e s e a r c h Program i n the b a s i n o f Marmot Creek (Munn and S t o r r 1967), and o t h e r s p e c i a l i z e d p r o j e c t s (MacHattie K a n a n a s k i s ( e l e v a t i o n k,560 f e e t MSL, W)  1966,  1967).  l a t i t u d e 51°02'N, l o n g i t u d e 1 1 5 ° 0 3 '  i s the o n l y s t a t i o n w i t h a r e l a t i v e l y l o n g c l i m a t o l o g i c a l r e c o r d upon  which acceptable  average c o n d i t i o n s can be based, but even some o f the  parameters r e c o r d e d  at t h i s s t a t i o n are based on s h o r t term, and  times i r r e g u l a r p e r i o d s .  T h i s s t a t i o n i s l o c a t e d 300  o f study l o c a t i o n 1 ( F i g s . 38 and grassed  39)j  some-  f e e t t o the  east  on a k n o l l on the edge o f a l a r g e  c l e a r i n g , open t o the south and e a s t , w i t h k-0 f o o t t r e e s  im-  m e d i a t e l y t o the n o r t h , t h u s the s i t e i s not p a r t i c u l a r l y t y p i c a l o f this forested region.  A l l o t h e r c l i m a t o l o g i c a l s t a t i o n s i n the  lower  K a n a n a s k i s v a l l e y have been e s t a b l i s h e d d u r i n g the l a s t decade, and  again  r e c o r d s f o r some parameters a r e • i n t e r m i t t e n t , and i n the case o f P i g e o n Lookout ( e l e v . 6,000 f t MSL,  l a t . 51°03'N, long. 1 1 5 ° 0 V w ) depend on  the d u r a t i o n o f the summer f o r e s t f i r e season.  Tables  I , I I and I I I ,  g i v e the c l i m a t i c summaries f o r the m e t e o r o l o g i c a l parameters r e c o r d e d Kananaskis,  K a n a n a s k i s Boundary Ranger S t a t i o n ( e l e v . k,800 f t MSL,  lat.  50°55'W, l o n g . 115°08'W), P i g e o n Lookout, and f o r a few s t a t i o n s i n the Marmot Creek B a s i n ( e l e v . 5,200 t o 9,200 f t MSL, 115°10'W).  Data f o r Kananaskis,  l a t . 50°57%  at  long.  K a n a n a s k i s Boundary and P i g e o n Lookout  s t a t i o n s were e x t r a c t e d from the "Monthly Record", Canada Department o f  F i g . 39-  Study l o c a t i o n 1, on the K a n a n a s k i s F o r e s t E x p e r i m e n t S t a t i o n , near the n o r t h e a s t shore o f B a r r i e r Lake, showing the l o c a t i o n o f i n f e c t e d l o d g e p o l e p i n e , Comandra p l a n t p l o t s , and i n s t r u m e n t s used d u r i n g the s t u d y .  Table I.  Monthly and annual climatic summaries for Kananaskis, l a t . 51°02'N, long. 115 03'W, 4,560 f t MSL, for the period of record (1939 - 1968).  elev.  JAN  FEB  MAR.  APR  MAY  JUN  JUL  AUG  SEP  OCT  NOV  DEC  YEAI  15.2 59 -50  20.4 61  24.8  51.4 88 23  57-6 93 23  55.7 92 29  86 15  40.9 80 - 8  27.2 66 -32  21.8  64 -41  45.0 82 - 7  48.7  -42  34.3 75 -24  64  36. c 93 -50  1.05 10.24  1.41 14.07  i.4o 12.89  2.49 20.18  3.06  4.08  2.49 0.00  2.78 0.16  2.28 0.34  1.44 8.88  1.15 10.52  1.17 11.24  24.7S 97.5:  69 26 1  138  150  41 2  214 45 2  236  48 4  308 62 5  254 57 5  163 43 5  121 37 2  71 27 1  61 25 1  2031 43  2  246 59 l  6.8  6.8  5.9  5-9  5-3  5-4  5.0  4.8  5-1  7.2  7-2  7.7  6.1  9 9 14 6 5 "27 20 5 9 13  10 9 ll 6 6 • 28 21 10 6 12  8 9 14 8 6 23 26 8 4 17  8 10 13 10 5 25 20 7 3 20  7 7 14 .10 8 26 17 9 2 21  8 9 14 10 10 25 15 7 3 22  ' 5 9 13 10 7 29 16 7 5 14  .  TEMPERATURE " F Daily mean Extreme maximum Extreme minimum  1  -44  1 PRECIPITATION inches Total Snowfall SUNSHINE DURATION hours Average Per cent of possible Years of data WIND SPEED mph Mean  7.48 1.53  2  3  WIND DIRECTION FREQUENCY North Northeast East Southeast South Southwest West Northwest Calm Years of wind data  <j? 8 8 12 9 4 29 17 5 9 12  .'8  5 10 3 4 39 18 9 4 10  Data f o r period August 1939 to December 1968 Data for 1939-19^1, 3 D a t a for 1939-1941, 1946-1968 (summers only 1946-1954).  4 6 9 9 5 40 16 6 5 15  6 7 9 8 4 38 18 5 6 11  6 8 9 10 5 37 17 5 6 12  7 8 12 8 • 6 30 18 7 5  -  1946-1947 and 1968, largely summers on  T a b l e I I . Monthly and a n n u a l c l i m a t i c summaries f o r s e l e c t e d s t a t i o n s i n the Marmot Creek Research l a t . 50°57'N, l o n g . 115°10'W, e l e v . 5,300 - 8,000 f t MSL ( 1 9 6 2 . - 1967). Station  Basin,  Years , Record  Oct  Nov  Dec  Yr  MEAN DAILY TEMPERATURE °F ko.2 4 9 . 2 56.9 56.2 48.4 38.3 47-3 54.3 53-5 46.2 34.2 43.2 53.0 51.4 42.3  39-6 36.9 32.7  23.4 22.1 19.4  17.2 15.3 14.7  35-3 33.4 29.8  5 5 2-4  MEAN 38.9 4l.O 30.8  DAILY 50.0 48.7 43.9  MAXIMUM TEMPERATURE °F 59-9 6 8 . 9 68.0 59-6 58.8 6 7 . 8 6 6 . 8 57-7 5 2 . 8 66.0 6 2 . 8 51-9  48.1 46.9 4o.6  31.0 30-3 26.6  24.2 23.4 21.3  44.5 44.0 39.0  5 5 2-4  l4.7 10.6 6.2  MEAN 21.5 20.9 12.9  DAILY 30.4 28.0 24.5  MINIMUM TEMPERATURE °F 38.5 kk.$ 4 4 . 3 137-2 ; 35-8 4 0 . 9 4 0 . 1 34.6 33-6 39-9 39-9 32.6  .30.3..15.8 27.1 14.0 24.8 12.2  10.2 7-3 8.0  25.9 23.0 20.7  5 5 2-4  -1.26 --  -1.34 --  -2.54 --  27.1  6 4-6 4  5-2  5-3 , W,  MEAN WIND SPEED mph and DIRECTION 4.7 5-2 4.8 5-2 5-1 k.8 SE NW NW, , W „ w 'w  4.9 NW/W  2-3 2-3  Elevation (feet)  Jan  Feb  Mar  Apr  'May  Confluence 1 Confluence 5 Cabin 5  5300 5800 6800  14.9 12.8 12.6  24.1 22.6 17.4  23.0 21.2 15.9  30.6 31.0 21.9  Confluence 1 Confluence 5 Cabin 5  5300 5800 6800  22.4 21.4 19.5  32.5 33-0 26.5  31-2 31-7 25.6  Confluence 1 Confluence 5 Cabin 5  5300 5800 6800  7.4 4.2 5-6  15.7 12.1 8.3  Confluence 1 Confluence 5 Cabin 5  5300 5800 6800  -1.26 --  Confluence 4  5600  4.3 W  NW (W)  S  (SE)  --  --  . --  --  Confluence 5  5750  --  --  —  —  Aug  Sep  /  (W/SE)  8000  Jul  TOTAL PRECIPITATION i n c h e s -4 . 2 6 2.17 2.29 1-843.06 4.55 2.50 2.53 2.04 -3-47 2.26 3.09 2.33  /  Upper Ridge  Jun  --  (SE)  ---  (NW/SE)  10.7 NW/SW  N  (NW/SE)  8.1 SW/W  MEAN EVAPORATION i n c h e s — 3-29 4.86 3-88  lAnemometer moved t o 150 f t tower a t 5,900 f t i n J u l y 1967,  2.17  9.7 SW  m  5.9 s  2.25  ,  4.2 W  (SW)  (NW)  11.2 sw/w  p r e v i o u s l y on 33 f t tower.  k.9  s  12.4 SW  2.70  1.55  13.5 w  x  T,of  2-4 1-4 4-6  Table I I I .  M o n t h l y t e m p e r a t u r e and p r e c i p i t a t i o n averages a t K a n a n a s k i s Boundary Ranger S t a t i o n , l a t . 50°55'N, l o n g . 115°08'W, e l e v . 4 , 8 0 0 f t MSL ( 1 9 6 2 - 1 9 6 8 ) , and P i g e o n M o u n t a i n Lookout, l a t . 51°03'N, l o n g . 1 1 5 ° 0 V W , e l e v . 6 , 0 0 0 f t MSL ( 1 9 6 0 - 1 9 6 8 ) % compared w i t h t h o s e a t K a n a n a s k i s f o r a s i m i l a r p e r i o d (1963-I968). 1  Station  JAW  FEB  MAR  APR  K a n a n a s k i s Boundary Pigeon Lookout Kananaskis  12.9 -15.I  19.2 -26.7  23-5 -25.6  3 0 . 8 1+2.5 --34.5 43.6  K a n a n a s k i s Boundary Pigeon Lookout Kananaskis  1.45 I . 8 7 1 . 6 l --l . l 4 0 . 8 2 1.57  2  2  1.72 -3-39  MAY  JUW  JUL  AUG  SEP  MEAN TEMPERATURE °F 4 9 . 5 5 6 . 3 5 4 . 8 48.8 47*3 5 6 . 1 5 5 - 1 ^ 5 - 3 5 0 . 6 57-4 55-8 4 9 . 3  T e m p e r a t u r e d a t a 1963, I965-I968;  WOV  DEC  42.2  24.6  12.3  34.8  40.9  26.3  18.1  36.2  1.71  2.59  26.09  1-33  1-88  26,98  TOTAL PRECIPITATION i n c h e s 3-02 ' 4 . 7 4 1.68 2 . 0 5 2 . 0 3 1.62 -5.98 2.71 2.29 3.09 I . 9 9 3 - 2 2 5-12 I . 8 3 2 . 4 7 2 . 7 8 1 . 4 3  •^-Temperature d a t a began 1963. 2  OCT  P r e c i p i t a t i o n data i960,  I962-I968.  YR  -  56  -  T r a n s p o r t ( v e r y r e c e n t months from t h e C l i m a t o l o g i c a l S t a t i o n R e p o r t , Forms 230k  and 2 3 0 6 ) ,  and t h o s e from Marmot Creek s t a t i o n s . from t h e  " C o m p i l a t i o n o f H y d r o m e t e o r o l o g i c a l R e c o r d , Marmot Creek B a s i n " , V o l ume  I - I I I , Water S u r v e y o f Canada, Department o f E n e r g y Mines  and  Resources, Calgary, A l b e r t a . The w i n t e r c l i m a t e i s c h a r a c t e r i z e d by an a l t e r n a t i o n o f c o l d , d r y , r a t h e r s t i l l p e r i o d s , w i t h p e r i o d s o f c o m p a r a t i v e l y warm, d r y , windy, chinook a i r , w h i c h g i v e s t o t h e g e n e r a l a r e a o f s o u t h w e s t e r n A l b e r t a o f the g r e a t e s t w i n t e r temperature ranges. e x p e r i e n c e d a 108°F t e m p e r a t u r e range.  I n December 1968,  one  Kananaskis  The v a r i a t i o n i n w i n t e r weather  i s d e t e r m i n e d l a r g e l y by t y p e and c i r c u l a t i o n o f a i r masses. c i r c u l a t i o n i s from t h e n o r t h and west w h i c h r e s u l t s i n t h e  The main predominance  o f m a r i t i m e P o l a r (mP) and c o n t i n e n t a l A r c t i c (cA) a i r masses, w i t h t h e o c c a s i o n a l o c c u r r e n c e o f u n s t a b l e and v e r y c o l d m a r i t i m e A r c t i c a i r masses (Penner 1 9 5 5 ) .  M a r i t i m e T r o p i c a l (mT)  (mA)  a i r masses may o c c a s i o n -  a l l y e n t e r t h e r e g i o n a l o f t from t h e s o u t h i n w i n t e r a s c e n d i n g t h e P o l a r a i r m a s s e s b u t never r e a c h i n g t h e s u r f a c e (Anonymous 1 9 5 6 ) . p e r i o d s o f t h a w i n g may  o c c u r i n a l l w i n t e r months when mP  r e g i o n , w i t h t e m p e r a t u r e s i n t h e 50's not b e i n g uncommon.  Week-long a i r enters the LongLey  ( 1 9 6 7 b ) found t h a t K a n a n a s k i s , on average, had 29 Chinook days  (above  !+0°F) d u r i n g t h e w i n t e r months December t o F e b r u a r y , w h i c h was  two more  t h a n C a l g a r y , and 19 more t h a n B a n f f .  I n c o n t r a s t , p e r i o d s o f sub-zero  t e m p e r a t u r e s o f a d u r a t i o n o f a week or more are c o m p a r a t i v e l y r a r e .  Ex-  treme low t e m p e r a t u r e s o c c u r when s t a b l e cA a i r s t a g n a t e s over t h e e a s t e r n slopes  o f the R o c k i e s and w e s t e r n P r a i r i e s .  O f t e n twjr.  c 1. _  :'• '..*;:  - 57 -  temperatures i n the v a l l e y are lower t h a n on the h i g h e r s l o p e s as c o o l a i r c o l l e c t s i n the v a l l e y under i n v e r s i o n c o n d i t i o n s , and Chinook a i r , o r warm a i r masses under subsidence are o n l y e x p e r i e n c e d a t the h i g h e r levels.  The Chinook i s c h a r a c t e r i z e d by a s t r o n g w e s t e r l y f l o w o f mP  a i r w i t h l e e waves f o r m i n g troughs and c r e s t s r o u g h l y p a r a l l e l t o the mountain r a n g e s , w h i c h r e s u l t , when warm d r y a i r i s drawn i n t o a midl a t i t u d e low p r e s s u r e c e n t e r i n the l e e o f the Rocky Mountains. descends t h e leeward s i d e o f the mountains  The a i r  a t the d r y a d i a b a t i c l a p s e  r a t e w h i c h b r i n g s h i g h temperatures and low h u m i d i t i e s t o the areas where t h e Chinook reaches the ground.  Condensation and c l o u d s form near  t h e c r e s t o f each s t a n d i n g wave w i t h the a s c e n d i n g o f a i r , g i v i n g t h e w e l l known Chinook A r c h .  Much o f the r e d b e l t c o n i f e r f o l i a g e  injury  observed on v a l l e y s l o p e s , which i s v e r y prominent i n some seasons,  has  been a t t r i b u t e d t o the abrupt a l t e r n a t i o n s o f c o o l a r c t i c a i r and warm chinook a i r (Henson 1952;  M a c H a t t i e 1963).  The w i n t e r s are r e l a t i v e l y  d r y , o n l y about 30% o f the a n n u a l p r e c i p i t a t i o n a t K a n a n a s k i s ( T a b l e I ) o c c u r s d u r i n g t h e s i x w i n t e r months, October t o March.  Snow accounts  f o r n e a r l y 40% o f the a n n u a l p r e c i p i t a t i o n o f about 25 i n c h e s a t Kanana s k i s , b u t much o f t h i s f a l l s i n A p r i l , which i s the h i g h e s t s n o w f a l l month. June 1951  The h e a v i e s t s i n g l e s n o w f a l l on r e c o r d a t K a n a n a s k i s o c c u r r e d i n when 33 i n c h e s f e l l i n two days.  v a l l e y , S t o r r (1967) found t h a t 70 t o 75%  A t the h i g h e r l e v e l s o f the o f the a n n u a l p r e c i p i t a t i o n  o c c u r r e d as snow o r a m i x t u r e o f r a i n and snow.  June i s t h e w e t t e s t month  o f the y e a r , and August the month o f g r e a t e s t p r e c i p i t a t i o n v a r i a b i l i t y . The v a r i a t i o n o f summer p r e c i p i t a t i o n and temperature depends upon the  - 58 -  d u r a t i o n and f r e q u e n c y o f s u b s i d i n g d r y mP a i r from t h e west, mT a i r from t h e G u l f o f Mexico w h i c h may b r i n g c o n s i d e r a b l e m o i s t u r e t o t h e F r o n t Ranges, and i n l a t e summer and autumn t h e c o n t i n e n t a l T r o p i c a l (cT) a i r which may p e n e t r a t e from t h e s o u t h and g i v e h o t , d r y s p e l l s .  In  s p r i n g and autumn, mA a i r may e n t e r t h e r e g i o n f o r a few days b u t i t i s absent i n summer.  V a l l e y bottom s t a t i o n s r e c e i v e l e s s p r e c i p i t a t i o n  than v a l l e y slope s t a t i o n s .  S t o r r (1967)  found t h a t on t h e average sum-  mer r a i n f a l l i n c r e a s e d about 1 . 5 i n c h e s p e r thousand f e e t i n t h e e a s t f a c i n g Marmot Creek b a s i n w i t h a l e v e l l i n g o f f a t about 7 , 6 0 0 f e e t MSL. A l t h o u g h t h e y e a r s o f r e c o r d o f sunshine a r e s h o r t , t h e low t o t a l hour v a l u e s f o r May and June, when l e s s t h a n 50 p e r cent o f t h e p o s s i b l e d u r a t i o n was r e c o r d e d ( T a b l e I ) , ' r e f l e c t t h e passage o f lows j u s t t o the s o u t h o f t h e a r e a w h i c h b r i n g c l o u d y , m o i s t a i r t o t h e r e g i o n . There i s g e n e r a l l y l e s s v a r i a t i o n i n t h e summer month temperatures than i n w i n t e r temperatures.  Temperatures  above 8o°F a r e e x p e r i e n c e d  i n s e v e r a l months b u t t h e h i g h e l e v a t i o n o f t h e a r e a , a l l above 4 , 2 0 0 f e e t , i s r e s p o n s i b l e f o r c o o l summer n i g h t s and lower daytime t u r e s t h a n o c c u r on t h e h o t , d r y P r a i r i e s t o t h e e a s t . t u r e s u s u a l l y o c c u r near t h e end o f J u l y .  tempera-  Maximum tempera-  D i f f e r e n c e s i n t h e average  d a i l y maximum and minimum temperatures d u r i n g t h e summer months a r e a p p r o x i m a t e l y 10 t o 15°F between t h e upper and lower areas o f t h e v a l l e y (Munn and S t o r r 1 9 6 7 ) .  F r o s t s can o c c u r i n any month, and t h e  average f r o s t - f r e e p e r i o d f o r t h e y e a r s 1 9 5 1 t o 1964 a t K a n a n a s k i s was 62 days, w i t h t h e average date o f t h e l a s t s p r i n g f r o s t , June 2 1 , and the f i r s t autumn f r o s t , August 22 ( L o n g l e y 1 9 6 7 a ) .  D u r i n g t h e summer  -  59  -  months t h e r e were many o c c u r r e n c e s o f low n i g h t r e l a t i v e h u m i d i t i e s i n the Kananaskis v a l l e y ( M a c H a t t i e 1966)  when c h i n o o k - t y p e winds o c c u r r e d .  Webb ( 1 9 6 5 ) showed t h a t t h e s e low h u m i d i t y n i g h t s were a s s o c i a t e d w i t h a i r subsidence under c o n d i t i o n s f a v o u r i n g l e e wave f o r m a t i o n . (.1966) found t h a t d a i l y minimum h u m i d i t i e s were' r e m a r k a b l y o f v e g e t a t i o n cover, s i t e and topography,  independent  and t h a t the i n c r e a s e w i t h  e l e v a t i o n was v e r y s l i g h t from v a l l e y bottom t o 1 , 0 0 0 the v a l l e y s i d e s .  f e e t e l e v a t i o n up  N i g h t l y maximum h u m i d i t i e s were more v a r i a b l e  f r e q u e n t l y d e c r e a s e d a b r u p t l y w i t h e l e v a t i o n j u s t above v a l l e y (11%  MacHattie  bottom  i n 300 f e e t ) w i t h a more g r a d u a l d e c r e a s e above t h i s l e v e l  t h a n 1% p e r 100  and  (less  feet).  The mean monthly wind speeds a t K a n a n a s k i s were h i g h e r i n the w i n t e r months, when chinooks are f r e q u e n t , t h a n i n the summer months ( T a b l e I ) . December had the h i g h e s t mean wind speed and J u l y , f o l l o w e d by August, the l o w e s t .  closely  The s t r o n g e s t winds come from the s o u t h -  west o r west a t a l l times o f the y e a r .  The dominant wind d i r e c t i o n i n  p r a c t i c a l l y a l l months o f the year was  from the southwest, a l t h o u g h i n  c e r t a i n y e a r s winds from the s o u t h e a s t or e a s t were dominant i n some o f t h e summer months.  M a c H a t t i e ( 1 9 6 7 ) i n d i c a t e d t h a t t h i s s o u t h e a s t wind  component was dominant a t n i g h t , a l t h o u g h o f o n l y low speed, and  was  t y p i c a l o f a d o w n v a l l e y wind coming from the Lusk Creek s u b - v a l l e y .  He  showed t h a t the wind components a c r o s s the main v a l l e y had a more p r o nounced d a y - n i g h t c y c l e t h a n t h e wind components a l o n g the v a l l e y , b o t h a t K a n a n a s k i s and i n Marmot Creek a t 5680 f e e t .  No a p p r e c i a b l e d i u r n a l  o s c i l l a t i o n o f winds o c c u r r e d up and down the v a l l e y a t K a n a n a s k i s i n  - 6o -  summer, b u t he found t h a t t h e southwest component c o u l d be dominant f o r most o f t h e day, o r under c e r t a i n c o n d i t i o n s , d u r i n g o n l y t h e d a y l i g h t hours.  I n t h e summer months t h e r e was u s u a l l y a marked maximum wind  speed i n mid- and l a t e - a f t e r n o o n , w i t h a minimum o c c u r r i n g around sunrise.  A t a v a l l e y bottom s t a t i o n near t h e c o n f l u e n c e o f Marmot Creek  and t h e K a n a n a s k i s R i v e r t h e r e was a morning-evening s l o p e wind  cycle,  w i t h t h e wind b l o w i n g toward t h e more i n t e n s e l y i n s o l a t e d s l o p e .  Munn  and S t o r r (1967) a l s o showed t h e prominence o f t h e n o r t h w e s t - s o u t h e a s t sub v a l l e y winds i n August a t t h e 5,600 f o o t l e v e l on Marmot Creek.  They  found a wind speed maximum j u s t b e f o r e s u n r i s e w i t h a d o w n v a l l e y wind, and another maximum i n t h e e a r l y a f t e r n o o n w i t h an u p v a l l e y wind.  The  minima, about 0800 and 1800 h o u r s , were a s s o c i a t e d w i t h wind s h i f t s from d o w n v a l l e y t o u p v a l l e y and v i c e v e r s a .  A t the Confluence h s t a t i o n a t  5,600 f e e t ( T a b l e I I ) , v e r y c l o s e t o t h e l o c a t i o n used by Munn and S t o r r (1967),  t h e r e was a l s o a h i g h o c c u r r e n c e o f d o w n v a l l e y n o r t h w e s t winds  i n many months, and o f u p v a l l e y s o u t h e a s t winds i n t h e months March t o J u l y , b u t a t t h e r i d g e s t a t i o n a t 8000 f e e t , t h e winds were p r e d o m i n a n t l y from t h e southwest, and a t a monthly mean v e l o c i t y two o r more t i m e s t h o s e o f t h e lower s t a t i o n . A comparison o f monthly mean t e m p e r a t u r e s and monthly p r e c i p i t a t i o n f o r t h e p e r i o d 1963 t o 1968 ( T a b l e I I I ) ,  showed t h a t K a n a n a s k i s  had warmer summer months t h a n K a n a n a s k i s Boundary by about 1 ° , and warmer w i n t e r months by as much as 7 ° .  D u r i n g t h e w i n t e r months K a n a n a s k i s r e -  c e i v e d l e s s p r e c i p i t a t i o n , b u t more d u r i n g t h e summer months t h a n Kanana s k i s Boundary.  Temperatures a t P i g e o n Lookout were 1° c o o l e r i n J u l y  -  and August,  and 3 t o 5°  61  -  c o o l e r i n June and September t h a n a t K a n a n a s k i s ,  and P i g e o n Lookout g e n e r a l l y r e c e i v e d more p r e c i p i t a t i o n .  M o n t h l y temp-  e r a t u r e s were a l s o g e n e r a l l y lower i n the Marmot Creek B a s i n ( T a b l e I I ) than a t K a n a n a s k i s .  P r e c i p i t a t i o n from A p r i l t o June was h i g h e r a t  K a n a n a s k i s t h a n i n Marmot Creek.  VEGETATION  The  area f a l l s  l a r g e l y w i t h i n the S ub-alpine F o r e s t R e g i o n r  (Rowe 1 9 5 9 ) ; w h i c h i s found between the approximate  l i m i t s of  5,000  and 6 , 5 0 0 f e e t , where the c l i m a x s p e c i e s a r e Engelmann s p r u c e , P i c e a engelmanni  P a r r y , and w e s t e r n w h i t e s p r u c e , P i c e a g l a u c a (Moench) Voss  v a r . a l b e r t i a n a (S. Brown) S a r g . , a l t h o u g h l a r g e a r e a s are covered by the s u b - c l i m a x s p e c i e s , l o d g e p o l e p i n e .  A t h i g h e r e l e v a t i o n s , between  6, 500 f e e t and the t i m b e r l i n e , the A l p i n e F o r e s t s are c h a r a c t e r i z e d by t h e dominance o f a l p i n e f i r , A b i e s l a s i o c a r p a (Hook.) N u t t . , and larch, Larix l y a l l i i Pari. Douglas  A t the n o r t h e r n end o f the K a n a n a s k i s  alpine valley,  f i r , Pseudotsuga m e n z i e s i i ( M i r b . ) F r a n c o , a c l i m a x s p e c i e s o f  the Montane F o r e s t R e g i o n i s p r e s e n t on warm, d r y s l o p e s a t lower v a t i o n s (up t o 4 , 7 5 0 f e e t ) .  ele-  Balsam p o p l a r , Populus b a l s a m i f e r a L. o c c u r s  on the a l l u v i a l s o i l s o f t h e v a l l e y , and t r e m b l i n g aspen, P. t r e m u l o i d e s Michx.,  competes w i t h l o d g e p o l e p i n e , as a p i o n e e r s p e c i e s f o l l o w i n g  fire,  on the lower s l o p e s o f the v a l l e y where brown f o r e s t s o i l s  predomin-  ate.  F i r e has been the most i m p o r t a n t f a c t o r i n f o r e s t s t a n d development  i n the a r e a , and throughout the S u b - a l p i n e and F o o t h i l l s F o r e s t Regions of A l b e r t a (Horton I 9 5 6 ; Smithers 1962).  Smithers (1956) records a  - 62 -  major f i r e i n t h e a r e a around 1865, i n 1910,  1920 and. 1936.  and o t h e r i m p o r t a n t f i r e s o c c u r r e d  T h i s l e a d t o l o d g e p o l e p i n e stands o f even age,  sometimes o v e r s t o c k e d w i t h v e r y h i g h d e n s i t i e s p e r a c r e .  A number o f  workers have s t u d i e d t h e s i l v i c a l and e c o l o g i c a l c h a r a c t e r i s t i c s o f l o d g e p o l e p i n e i n t h e a r e a and much o f t h i s work has been summarized by Smithers  (1962). F o r e s t cover maps have r e c e n t l y been p u b l i s h e d f o r t h e Kanan-  a s k i s F o r e s t Experiment Development, I 9 6 7 ) ,  S t a t i o n (Canada Department o f F o r e s t r y and R u r a l  and Marmot Creek Watershed R e s e a r c h B a s i n ( D e p a r t -  ment o f F o r e s t r y o f Canada, I 9 6 5 ) , w h i c h i n d i c a t e t h e major s p e c i e s p r e s e n t , t h e i r average h e i g h t c l a s s , and crown c l o s u r e d e n s i t y , as w e l l as showing p o o r l y d r a i n e d and r o c k o u t c r o p a r e a s .  The f o r e s t and a l p i n e  cover t y p e s and h a b i t a t t y p e s o f t h e Marmot Creek Watershed B a s i n , w i l l be p u b l i s h e d by K i r b y and O g i l v i e i n 1969.  Research  This w i l l r e -  p r e s e n t t h e f i r s t d e t a i l e d e c o l o g i c a l v e g e t a t i o n r e p o r t f o r any.area i n the K a n a n a s k i s v a l l e y .  The r e p o r t d e s c r i b e s e l e v e n f o r e s t h a b i t a t t y p e s  and t e n a l p i n e h a b i t a t t y p e s , and i n d i c a t e s t h a t kQfo o f t h e Marmot Creek Basin c o n s i s t s of non-productive and r o c k .  forest area —  a l p i n e f o r e s t , meadow  The r e s t o f t h e a r e a i s l a r g e l y o f t h e s p r u c e - f i r - p i n e  t y p e , w i t h areas o f p i n e and p i n e - a s p e n  cover types a t lower  where t h e y became e s t a b l i s h e d a f t e r t h e 1936  cover  elevations,  fire.  STUDY LOCATIONS  The  s t u d y l o c a t i o n s ( F i g . 38) o c c u r r e d i n b o t h mixed and even  aged stands o f l o d g e p o l e p i n e , except l o c a t i o n 8 w h i c h was i n a g r a s s  - 63 -  clearing.  The even aged stands were a p p r o x i m a t e l y  kO y e a r s - o l d , and became e s t a b l i s h e d a f t e r f i r e . regeneration mixed s t a n d  10, 2k, 28,  32 and  There was a l s o one  a r e a w i t h few t r e e s over 5 y e a r s o l d ( l o c a t i o n 9).  The  ( l o c a t i o n l ) had t r e e s r a n g i n g i n age from 7 t o 60 y e a r s ,  w i t h most t r e e s grouped i n t h e 21 o r 52 y e a r age c l a s s .  Location 3  was s i t u a t e d i n t h e lower l e v e l s o f t h e Marmot Creek Watershed R e s e a r c h B a s i n near t h e c l i m a t o l o g i c a l s t a t i o n , C o n f l u e n c e 1 (Table I I ) . Laboratory minations  experiments w i t h a e c i o s p o r e s ,  d a i l y counts and g e r -  t e s t s , were carried;.'out i n t h e l a b o r a t o r y a t t h e K a n a n a s k i s  Forest Experiment S t a t i o n , or at the Forest Research Laboratory, gary.  The spore d i s p e r s a l and m i c r o c l i m a t i c s t u d i e s were c a r r i e d o u t  at l o c a t i o n s 1, 2, d a i l y germination  3 and 8 ( F i g . 38).  The a e c i o s p o r e  t e s t s came from l o c a t i o n 1.  collections for  Spore c o l l e c t i o n s f o r ex-  p e r i m e n t a l purposes came from most o f t h e l o c a t i o n s .  Collections of the  a s s o c i a t e d m i c r o f l o r a and m i c r o f a u n a came from l o c a t i o n s 1, 2, 7.  Cal-  Measurements and o b s e r v a t i o n s  3, 5 and  o f canker growth and a c t i v i t y , and t h e  i n c i d e n c e o f a s s o c i a t e d m i c r o f l o r a , m i c r o f a u n a and r o d e n t damage, were made a t a l l . l o c a t i o n s e x c e p t l o c a t i o n 8. Besides obtained  t h e main a r e a o f study,  a d d i t i o n a l i n f o r m a t i o n was  from many p o i n t s w i t h i n t h e S u b - a l p i n e ,  Montane and F o o t h i l l s  F o r e s t Regions between Robb (53°13'N, ll6°58'W) and W a t e r t o n Lakes Nat i o n a l P a r k (49°03'N, 113°55'W"). measurements and o b s e r v a t i o n s  The i n f o r m a t i o n m a i n l y  consisted of  o f canker growth and a c t i v i t y , and i n c i -  dence o f a s s o c i a t e d m i c r o f l o r a and m i c r o f a u n a .  C o l l e c t i o n s of the  a s s o c i a t e d m i c r o f l o r a were made from a number o f p l a c e s .  Material for  - 6k  -  the r e a r i n g o f a s s o c i a t e d m i c r o f a u n a from P i n u s c o n t o r t a was o b t a i n e d from B a r i l Creek, Saskatchewan R i v e r C r o s s i n g , C l i n e R i v e r , and near Robb, and from P. b a n k s i a n a , kO m i l e s south o f Rae, N.W.T. E x t e n s i v e s u r v e y f o r d i s t r i b u t i o n a l d a t a were c a r r i e d - out over the s o u t h e r n p o r t i o n o f the P r o v i n c e south o f 56°N.  Additional d i s t r i -  b u t i o n d a t a and m a t e r i a l f o r r e a r i n g a s s o c i a t e d m i c r o f a u n a was  gathered  by p e r s o n n e l o f the F o r e s t I n s e c t and D i s e a s e Survey i n the course o f their regular duties.  - 65 -  AECIOSPORE PRODUCTION  PERIODS OF AECIOSPORE PRODUCTION AND  ENVIRONMENTAL  FACTORS AFFECTING SPORULATION  " • '  Methods  :  A e c i a l cankers were observed  a t r e g u l a r i n t e r v a l s from e a r l y  May t o October a t l o c a t i o n s 1 and 2 ( F i g . 38) i n t h e y e a r s 196k t o .1968, at  l o c a t i o n 3 from 1966 t o 1968 and a t l o c a t i o n 9 i n 1968.  At location  1 o b s e r v a t i o n s were made 2 o r .3 times a week i n t h e y e a r s 196k t o I967 and w e e k l y i n 1968.  A t l o c a t i o n 2 o b s e r v a t i o n s were made s e v e r a l t i m e s  a week i n 196k and once o r t w i c e a week from 1965 onwards. and 9 were v i s i t e d a t l e a s t once a week.  Locations 3  I n some y e a r s o b s e r v a t i o n s were  made on a d a i l y b a s i s on s e l e c t e d t r e e s a t l o c a t i o n 1 t o e s t a b l i s h t i a t i o n o f s p o r u l a t i o n and d u r a t i o n o f 'production from i n d i v i d u a l and i n d i v i d u a l a e c i a l p u s t u l e s .  inicankers  S p o r u l a t i o n was c o n s i d e r e d t o have com-  menced w i t h t h e r u p t u r e o f t h e p e r i d i u m o f t h e aecium and t h e r e l e a s e o f the f i r s t a e c i o s p o r e s , and t o have ceased when no f u r t h e r a e c i o s p o r e s were p r e s e n t i n t h e r u p t u r e d aecium.  Three t o 30 i n d i v i d u a l p u s t u l e s  were marked on i n d i v i d u a l a e c i a l cankers w i t h l e t t e r e d o r numbered p i n s p r i o r to r u p t u r i n g o f the p e r i d i a .  E f f o r t s were made t o s e l e c t  p u s t u l e s which, would n o t c o a l e s c e w i t h nearby p u s t u l e s .  isolated  Observations o f  a e c i o s p o r e s p o r u l a t i o n were a l s o made a t o t h e r l o c a t i o n s a t i r r e g u l a r i n t e r v a l s i n t h e v a r i o u s y e a r s and p r o v i d e g e n e r a l i n f o r m a t i o n on t h e sporulation period.  D a i l y temperatures,  r e l a t i v e humidity,  rainfall,  wind and r a d i a t i o n d a t a were r e c o r d e d on hygrothermographs, r a i n gauges and r a i n f a l l r e c o r d e r , wind speed and d i r e c t i o n r e c o r d e r and a c t i n o g r a p h  -  66 -  p l a c e d a t l o c a t i o n 1 from mid May t o l a t e August. ments and i n f e c t e d t r e e s i s shown i n F i g . 39 j  a n d  Location of instru-  - additional details of  the i n s t r u m e n t s a r e g i v e n i n t h e a e c i o s p o r e d i s p e r s a l s e c t i o n . F o r A p r i l and t h e f i r s t h a l f o f May i n each year weather d a t a was o b t a i n e d from t h e K a n a n a s k i s ' c l i m a t o l o g i c a l s t a t i o n , 330 f e e t from t h e n e a r e s t i n f e c t e d t r e e o f the study area.  Weather d a t a was a l s o r e c o r d e d a t  l o c a t i o n s 2 and 3 f o r t h e p e r i o d o f study. In  a d d i t i o n t o o b s e r v i n g t h e phenology o f a e c i o s p o r e produc-  t i o n i n t h e d i f f e r e n t y e a r s a t t h e v a r i o u s l o c a t i o n s , t h e phenology o f the o t h e r spore s t a t e s o f t h e fungus was observed. Comandra p l a n t s were observed  A t l o c a t i o n 1,  two o r t h r e e t i m e s a week f o r t h e presence  o f t h e u r e d i a l and t e l i a l s t a t e s on f o u r p l o t s , each one square meter i n a r e a ( F i g . 3 9 ) - Some o b s e r v a t i o n s were a l s o made on t h e phenology o f l o d g e p o l e p i n e and Comandra p l a n t s a t t h i s  r''' . '.V . y v  location.  Results  i;  T a b l e I V summarizes t h e b e g i n n i n g and end o f a e c i o s p o r e  sporu-  l a t i o n a t l o c a t i o n s 1, 2,t:3 and 9 f o r 1964 t o 1 9 6 8 , and i n d i c a t e s t h e main spore p r o d u c t i o n p e r i o d .  Aeciospore production v a r i e d widely w i t h  s e a s o n a l c o n d i t i o n s and l o c a l i t y . mid-May t o e a r l y June.  Aeciospore  s p o r u l a t i o n began f r o m  A t h i g h e r e l e v a t i o n s t h e p e r i o d was one t o two  weeks l a t e r , and t r e e s on s i t e s w i t h a n o r t h o r west a s p e c t were l a t e r i n i n i t i a t i n g s p o r u l a t i o n than those w i t h a south o r e a s t a s p e c t .  The  main p e r i o d o f a e c i o s p o r e s p o r u l a t i o n l a s t e d t h r e e t o f i v e weeks, u s 1  u a l l y commencing a t t h e b e g i n n i n g o f June, a l t h o u g h i n I 9 6 5 and 1 9 6 8 spore p r o d u c t i o n was c o n c e n t r a t e d i n a two week p e r i o d .  For three years,  s p o r u l a t i o n ended i n t h e second -half o f J u l y , b u t i n t h e o t h e r two y e a r s 1  - 67  Table I V .  Year  -  Date o f b e g i n n i n g and end o f C r o n a r t i u m comandrae a e c i o s p o r e p r o d u c t i o n , and main s p o r u l a t i o n p e r i o d a t a number o f l o c a t i o n s i n the y e a r s 1964 - 1968 i n c l u s i v e . L o c a t i o n no.  Began  Ended  Main P e r i o d  1964  1 2  June 2 June 1  July July  I965  1 2  May May  24 18  J u l y 30 J u l y 24  May 30 - June  1966  1 2 3  May May  13 26  Aug. Aug. Aug.  May 26 - J u l y June 1 - J u l y  1967  1 2 3  May May May  31 30 31  Sept. 1 Aug. 3+ Aug. 17  June 5 - J u l y 8 June 10 - J u l y 10 June 12 - J u l y 10  1 2 3 9  June May JJune June  J u l y 18 J u l y 22+ J u l y 31 J u l y 30  June June June June  1968  6 31  2 2  19 18  18 10+ l6  June 3 - June 30 June 8 - J u l y 7 13  #  18 13 18 13  -  3 4  28 23 28 28  June June June June  * I n f o r m a t i o n not complete  spore p r o d u c t i o n c o n t i n u e d t o t h e m i d d l e o r end o f A u g u s t . s p o r a d i c and v e r y l i g h t r e s u m p t i o n  of aeciospore production,  I n I966 a induced  p o s s i b l y by warm weather, o c c u r r e d a t l o c a t i o n 2 i n September and  October.  The s c a t t e r e d a e c i a l p u s t u l e s , r a r e l y more t h a n 2 mm i n d i a m e t e r ,  occurred  i n t h e p y c n i a l zone o f t h e same y e a r . T a b l e V g i v e s t h e date o f b e g i n n i n g and end o f spore p r o d u c t i o n f o r a number o f cankers a t l o c a t i o n 1 f o r the y e a r s 1965 t o I 9 6 8 .  Figs.  40 and 4 l summarize t h e weather and phenology d a t a f o r t h e v a r i o u s r u s t s t a t e s a t l o c a t i o n 1 i n t h e same years.'  Most a e c i a l cankers i n any one  year began s p o r u l a t i o n a t a p p r o x i m a t e l y the same time a l t h o u g h some cank e r s were c o n s i s t e n t l y e a r l y , e.g. # 2 7 2 1 , b u t o t h e r s were l a t e , #2712.  e.g.  From o b s e r v a t i o n s t h e r e was a tendency f o r s p o r u l a t i o n t o o c c u r  Table V'.,  Date of beginning and end o f spore production, and t o t a l number of days o f production from i n d i v i d u a l cankers at location 1 from 1965 to 1968 i n c l u s i v e . Main spore production period i s included for 3 to 6 cankers for the years 1965 to 1967.  1965  Tree No.  1966  1968  1967 Main period  Total no. of days  Main period  Total no. of days  Main period  Total no. of days  July 30  June 5-11 July 2-10  57+  No production  May 10  July 2k  May 30June 11  60  June 1 June 15 (1 pustule only)  1U  May 28  Aug. 25  2711  May 29  June 26  29  May 2k  June 19  27  May 31  June 23  2712  June 1+  July 26  June 24-27  56+  May 30  Aug. 18  June 2-, July k  81  June 5  Aug. k  2713  May 27  July 26  June 5-13  61  May 20  Aug. 15  May 25July 3  June 2  Sept. 1  2715  June 8+  July 17  40+  May 26  Aug  77  June 6  Aug. 7  63  Canker area dead  2716  June h  July 20  kl  Tree dead  2717  No data  2719  May 2k  July 16  2720  May 30+  June 20  2721  May 22  July 12  2722  May 2k  2723 2728  Began  Ended  2516  June 1*+  2710  June 5-17  Began  Ended  Began  Ended  Began  Ended  90  June 6  July 6  2k  No production  June 11-July 18  60  June 15  June 5Aug. 7  92  T o t a l no. of days  Tree dead  10  June 2June 25  July 8  June' 8 June 30 (tree dead July 22)  31 0 2k  23 —  June 2+  July 30  59+  June 2  Aug. 15  75  June 10  J u l y 16  37  5k  May 26  Aug. 15  82  May 30  Aug. 15  78  June 10  J u l y 18  39  22+  Tree dead  52  May 13  July 27  July 5  1+3  May 22  July 8  kd  May 31  July 7  38  June 10  July 3  21+  May 2k  July k  k2  Tree dead  June 2+  July 22  51+  May 25  July 8  k5  May 31  Aug. 10  72  June 6  J u l y 18  >*3  May 26June 11  May 22July 3  76  May 30 June 27 (tree dead J u l y 18)  29  CA  co  F i g . hO.  Summary o f the d a i l y maximum and minimum t e m p e r a t u r e s : and r e l a t i v e h u m i d i t y t a k e n from hygrothermograph r e c o r d s , the d a i l y r a i n f a l l , and the phenology, o f the a e c i a l , pyc n i a l , u r e d i a l and t e l i a l s t a t e s o f C r o n a r t i u m comandrae, " May t o August i n 1965 and 1966.  - 69 -  MAXIMUM  30  10  15  25 30  10  15 20 JULY  20  25  30  10  10  15 20 JUNE  25  30  Moderate  Heavy  10  15  20  15 30  10  15  K>  15 20 MAY  25 30  10  10  15  10  25 30  10  15  13 20 MAY  25 30  20  25  20  15 30  10  15  10  20  AND MINIMUM TEMPERATURE °C  -10 RELATIVE HUMIDITY 16 NO.OF HOURS 80 A N D 100% * 0 2  4  2.0 1.5 RAINFALL INCHES  1.0 0.5 0  AECIOSPORES PYCNIOSPORES UREDIOSPORES TELIOSPORES  MAXIMUM  15 20 JUNE  15  20  25  30  5  10 15 20 AUGUST  5  10 15 20 AUGUST  25 30  jo  AND MINIMUM T E M P E R A T U R E "> °C  RELATIVE HUMIDITY 16 NO. O F H O U R S 80 A N D 100%  RAINFALL INCHES  1.0  AECIOSPORES PYCNIOSPORES UREDIOSPORES TELIOSPORES 10  80n HUMIDITY 100| %  Light  15 20 25 JULY  30  F i g . 1+1.  Summary o f the d a i l y maximum and minimum t e m p e r a t u r e s and r e l a t i v e h u m i d i t y t a k e n from hygrothermograph r e c o r d s , the d a i l y r a i n f a l l , and the phenology o f the a e c i a l , pyc n i a l , u r e d i a l and t e l i a l s t a t e s o f C r o n a r t i u m comandrae, May t o August i n 1967 and 1968.  - 70 -  10  MAXIMUM AND  15  20  25 30  10  IS  15 20 25 30 MAY  10  15 20 25 JUNE  10  15 20 25 30  20 25 30  10  15  20 25 30  10  15 20 JULY  25 30  10  15 20  25 30  10  15 20  20  MINIMUM TEMPERATURE  1 0  °C  -10 24  r  RELATIVE HUMIDITY NO. O F H O U R S 8 80 A N D 100% o 1 6  2.0 1.5 RAINFALL INCHES  1.0 0.5  AECIOSPORES PYCNIOSPORES UREDIOSPORES TELIOSPORES  l_  5  10  10  15  20 25 30  30  5  1  5  10 15 20 AUGUST  5  10 15 20 AUGUST  1  MAXIMUM AND MINIMUM TEMPERATURE °C  RELATIVE HUMIDITY NO. O F H O U R S 8 80 A N D 100% o  RAINFALL INCHES  AECIOSPORES PYCNIOSPORES UREDIOSPORES TELIOSPORES L_  10  15 20 25 30 10 15 20 25 30 MAY JUNE 80n HUMIDITY , 100i % , Moderate Heavy U  g  h  10  15 20 25 30 JULY  NR = No Record  - 71  -  f i r s t from cankers on the s m a l l e r b r a n c h e s , s e e d l i n g s and young t r e e s , a l l h a v i n g t h i n smooth b a r k , somewhat l a t e r on l a r g e branches w i t h t h i c k e r bark, and l a t e r s t i l l on stems o f o l d e r t r e e s w i t h a t h i c k e r rough b a r k .  T h i s was p r o b a b l y because the d e v e l o p i n g a e c i a were deeper  s e a t e d i n t h e t h i c k e r roughened b a r k t h a n i n t h i n smoother bark, because o f the m e c h a n i c a l problems  of r u p t u r i n g t h i c k bark.  and  Usually  the a e c i a l b l i s t e r s o f t h i c k b a r k e d cankers e r u p t e d i n the b a r k  fissures,  b u t r e m o v a l o f b a r k o f t e n exposed a l a r g e a e c i a l a r e a underneath where spores remain unexposed f o r much o f the season.  Cankers on t r e e s i n t h e  open or on a s o u t h or e a s t a s p e c t o f a t r e e a l s o began s p o r u l a t i o n s l i g h t l y e a r l i e r t h a n those in a dense s t a n d or l o c a t e d w i t h a d o m i n a n t l y west o r n o r t h a s p e c t .  Even on t r e e s which were n e a r l y g i r d l e d by t h e  fungus, s p o r u l a t i o n began from p u s t u l e s on the s o u t h e a s t o r s o u t h a s p e c t and was  l a s t from those on the n o r t h .  Cankers a few f e e t up the stem  a l s o s p o r u l a t e d b e f o r e b a s a l cankers s i t u a t e d i n the t h i c k e r b a r k zone of  a tree.  I n i t i a t i o n o f s p o r u l a t i o n was  about two t o t h r e e weeks a f t e r  e v i d e n c e o f shoot bud b r e a k on t h e l o d g e p o l e p i n e w h i c h u s u a l l y commenced i n l a t e A p r i l or e a r l y May.  S i m i l a r l y , growth o f Comandra p l a n t s began  before aeciospore sporulation.  Growth amounted .to an i n c h or more o f  a e r i a l p l a n t development by May  12 i n 1965  and May  10 i n I966, and the  Comandra p l a n t s were f l o w e r i n g by the l a s t week i n May sites.  Development was  l a t e r i n 1967  and 1968.  i n favourable  I n 1963  and I96U Comandra  p l a n t s were f i r s t noted as p r o d u c i n g u r e d i o s p o r e s a t l o c a t i o n 1 on June 18 and 28 r e s p e c t i v e l y .  I n l a t e r y e a r s the i n c i d e n c e o f i n f e c t i o n  and  u r e d i a l and t e l i a l development was r e c o r d e d on Comandra p l a n t s on t h e f o u r p l o t s a t .-.location 1 ( F i g . 39).  Each p l o t h a v i n g an average o f 53  - 72  -  These p l o t s were l a i d out w i t h i n 100  a e r i a l shoots each y e a r .  i n f e c t e d t r e e s , and i n most cases were w i t h i n 1+0 f e e t . f e c t i o n s were.recorded  I n 1966  i n t e r v a l o c c u r r e d i n 1965  ( F i g . 1+0),  p r e s e n t 12 days (on June 17)  after  A s i m i l a r long  but i n I968 ( F i g . l+l) u r e d i a were  a f t e r the f i r s t a e c i o s p o r e s were r e c o r d e d .  E a r l y s p r i n g s may  advance and l a t e s p r i n g s r e t a r d somewhat  the advent o f a e c i a l p r o d u c t i o n . than i n 1968.  no i n -  on the p l a n t s u n t i l 39 days (on June 28)  a e c i o s p o r e r e l e a s e from the a d j a c e n t t r e e s ( F i g . 1+0).  1966  feet of  E a r l y May  S p o r u l a t i o n was  (May  1-15)  was  t h r e e weeks e a r l i e r i n  u n u s u a l l y warm i n 1966  there  b e i n g t w i c e the number o f degree days above 0°C than i n the same p e r i o d i n 1967 May 1967  1966  and 1968,  and one and a h a l f times the number i n 1965.  the r e g i o n was  and 1968,  May was  e x p e r i e n c e d i n May  i n f l u e n c e d by a p r e v a l e n c e o f mP a i r , b u t i n dominated by mA  p e r a t u r e s t o the r e g i o n .  The presence temperatures,  and cA a i r masses.  1966", but t h i s a i r mass was  i n the f i r s t h a l f o f May  as o c c u r r e d i n 1967  1967 May  and 1968,  and 1968, 1965  and 1966  Wo  cA a i r was  p r e s e n t f o r a few days  b r i n g i n g w e l l below n o r m a l tem-  d i d not e x p e r i e n c e the c o l d cA a i r ,  b u t was  dominated by mA  a i r d u r i n g t h e month.  o f warmer a i r masses, w i t h t h e i r c o r r e s p o n d i n g warmer s u r f a c e c o u l d w e l l account  compared w i t h 1967  and 1968.  delayed to a l e s s e r  extent  A p r i l had a l s o been warmer i n I965  than i n the l a t t e r two y e a r s .  I n i t i a t i o n o f s p o r u l a t i o n i n most  y e a r s seemed t o s t a r t on a day o f , o r a day f o l l o w i n g , r a i n , t h a t m o i s t u r e was  1966  for sporulation being e a r l i e r i n  than the o t h e r y e a r s , and~.'why s p o r u l a t i o n was i n 1965  During  r e q u i r e d t o r u p t u r e the p e r i d i a .  The  suggesting  end o f the main  spore p r o d u c t i o n p e r i o d o f t e n c o i n c i d e d w i t h a heavy r a i n f a l l w h i c h washed the spores from the a e c i a .  T h i s was most n o t i