Open Collections

UBC Theses and Dissertations

UBC Theses Logo

UBC Theses and Dissertations

Relative susceptibility of interior spruce (Picea glauca x engelmannii engelm.) and lodgepole pine (Pinus… Schulting, Peter John 1987

Your browser doesn't seem to have a PDF viewer, please download the PDF to view this item.

Item Metadata

Download

Media
831-UBC_1987_A6_7 S38.pdf [ 8.24MB ]
Metadata
JSON: 831-1.0097051.json
JSON-LD: 831-1.0097051-ld.json
RDF/XML (Pretty): 831-1.0097051-rdf.xml
RDF/JSON: 831-1.0097051-rdf.json
Turtle: 831-1.0097051-turtle.txt
N-Triples: 831-1.0097051-rdf-ntriples.txt
Original Record: 831-1.0097051-source.json
Full Text
831-1.0097051-fulltext.txt
Citation
831-1.0097051.ris

Full Text

RELATIVE SUSCEPTIBILITY OF INTERIOR ENGELMANNII  SPRUCE (PICEA GLAUCA X  ENGELM.) AND LODGEPOLE PINE  (PINUS CONTORTA  DOUGL. VAR. LATIFOLIA ENGELM.) TO INONOTUS TOMENTOSUS (FR.) TENG IN CENTRAL BRITISH COLUMBIA  by PETER J . SCHULTING A THESIS SUBMITTED IN PARTIAL FULFILMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE s  in THE FACULTY OF GRADUATE STUDIES Department  We a c c e p t  this  t o the  of  Forestry  t h e s i s as c o n f o r m i n g  required  standard  THE UNIVERSITY OF BRITISH COLUMBIA 6 October © Peter  1987  J . Schulting,  1987  In presenting this thesis in partial fulfilment of the requirements for an advanced degree at the University of British Columbia, I agree that the Library shall make it freely available for reference and study. I further agree that permission for extensive copying of this thesis for scholarly purposes may be granted by the head of my department or by his or her representatives.  It is understood that copying or  publication of this thesis for financial gain shall not be allowed without my written permission.  •^o^e.sVcc^  Department of  The University of British Columbia 1956 Main Mall Vancouver, Canada V6T 1Y3 Date  QA.  1 ,  \<\H1  ABSTRACT  Twenty ( F r . ) Teng  sample  plots  c e n t e r s i n mixed  Dougl.  var. l a t i f o l i a  glauca  x engelmannii  Columbia  were  i n host  effect  infection  pine  was  studied  Characteristics isolates  In  of  the  and  to  by of  on  the  and  radial  the  each  the  and  the  often  tree,  symptoms.  from  Possible  cases  also  found.  level Nutt.)  than and  spruce  behaviour  increment  of  fungus  was  found  to  i n n e r xylem  of  the  roots caused spruce,  b e f o r e e x p r e s s i o n of n o t i c a b l e  infected of  spore  crown  spread  thinning  of  the  windfall  of  crown was  commonly  to healthy, c o n t a c t i n g roots. inoculation  In a l l p l o t s ,  pine  spruce. Subalpine Douglas-fir  using  necrosis  decay  roots caused  species studied,  and  behave  p i n e . In p i n e , advanced  In  The  were s t u d i e d plots.  the  and  bores.  study  of  of,  spruce  the  phloem  (Picea  British  of  and  In b o t h h o s t  observed  (Pinus contorta  the pathogen.  in culture  eventual standing mortality.  heartwood  the  increment  fungus  tomentosus  in central  investigate  measuring  spruce  cambium  interior  s p e c i e s a t t a c k e d by  some c a s e s , on  lodgepole pine  Engelm.) s t a n d s  o b t a i n e d from  differently  in Inonotus  Engelm.) and  studied  variation of  located  through  root  showed a h i g h e r  wounds were  infection  f i r (Abies l a s i o c a r p a  (Pseudotsuqa  i i  menziesii  (Hook.)  (Mirb.)  Franco) present infected.  Infection  significantly The  fungus  variation texture. isolates, which  within  the p l o t s  by  t o m e n t o s u s was  the basal  in culture i n growth  area  the i s o l a t e  increment  demonstrated  rate,  Such v a r i a t i o n and c o u l d  were n o t f o u n d  mycelium  occurred  obtained.  iii  a large  or  t o reduce spruce.  amount o f  a n d mat  both within  n o t be r e l a t e d  was  not found  i n pine  color  t o be  color  and  a n d among  to the host  species  from  T a b l e of  Contents  Abstract  ii  List  of T a b l e s  vi  List  of F i g u r e s  viii  Acknowledgement  ix  A.  Introduction  B. Review of  1  the L i t e r a t u r e  2  1 . Taxonomy  2  2. D i s t r i b u t i o n  and Host S p e c i e s  6  3. Methods of Spread  16  4.  20  Ecology  5. Damage  22  a . Windthrow  .  22  b.  Standing M o r t a l i t y  24  c.  Butt  d.  Increment R e d u c t i o n  26  e.  I n d i r e c t Losses  27  Cull  6. V a r i a b i l i t y  . . .25  of  Inonotus  tomentosus  C. Methods  29 31  1.  S e l e c t i o n of  S i t e s and P l o t  2.  Data C o l l e c t i o n  Location  31 35  3. Host S p e c i e s S u s c e p t i b i l i t y  36  4.  Effect  38  5.  I d e n t i f i c a t i o n and D e s c r i p t i o n  of  I n f e c t i o n on Increment  D. R e s u l t s and D i s c u s s i o n  in Culture  39 40  1. E t i o l o g y  40 iv  2.  Spread  3. H o s t 4.  54  Species  Effect  of  Infection  a. L o d g e p o l e b.  Interior  5. C u l t u r a l  Susceptibility on  Increment  54 73  pine  73  Spruce  76  Characteristics  .....79  E. C o n c l u s i o n s  89  Literature  92  Cited  Appendices  97  1. A p p e n d i x  A  ...97  2. A p p e n d i x  B  118  3. A p p e n d i x  C  119  v  List  of  Tables  1. H i s t o r i c a l nomenclature of 2. Host s p e c i e s of 3. Frequency of all 4.  fungal  Incidence  tomentosus  5  tomentosus  isolation  isolates  of  8 tomentosus as a p e r c e n t  of  i n Canadian decay s t u d i e s  l e v e l s of  tomentosus on  11  interior  spruce  in  Canada  13  5. Stand age,  r e g i o n and  ecological classification  of  sample s i t e s 6.  34  Proportions difference  of p i n e and  spruce  i n f e c t e d and  i n p r o p o r t i o n of p i n e and  spruce  the i n f e c t e d by  site  56  7. P a i r e d - s a m p l e T - t e s t on and  the p r o p o r t i o n of p i n e i n f e c t e d  the p r o p o r t i o n of spruce  infected  58  8. w i l c o x i n M a t c h e d - P a i r s Signed-Ranked t e s t , of p i n e  i n f e c t e d w i t h p r o p o r t i o n of spruce  infected  9. C o r r e l a t i o n of the p r o p o r t i o n of l o d g e p o l e i n f e c t e d with  the p r o p o r t i o n of spruce  10. T - t e s t on the  spruce  60  the  infected  62 of p i n e and  spruce  the d i f f e r e n c e between the p r o p o r t i o n s  spruce  pine  i n f e c t e d w i t h the p r o p o r t i o n of  11. Summary of the p r o p o r t i o n s and  59  infected  s l o p e of the c o r r e l a t i o n of  p r o p o r t i o n of p i n e  proportion  infected  of p i n e  and  infected  12. T - t e s t on geographic  66  the d i f f e r e n c e i n p r o p o r t i o n s region  i n f e c t e d by 67  vi  List  of T a b l e s  (con't)  13. Groupings of e c o l o g i c a l a s s o c i a t i o n s by m o i s t u r e and nutrient  status  6B  14. A n a l y s i s of v a r i a n c e of  the d i f f e r e n c e  in  the  p r o p o r t i o n of pine and spruce i n f e c t e d by e c o l o g i c a l grouping  70  15. Mean i n i t i a l healthy  b a s a l area and b a s a l area  of  and d i s e a s e d p i n e and spruce  16. A n a l y s i s of v a r i a n c e i n r a d i a l 17.  increment  increment of p i n e  75 77  Summary of mean stand ages and t r e e b a s a l a r e a s by  subzone 18. A n a l y s i s of v a r i a n c e  78 in r a d i a l  increment of spruce  19. Common mycelium types found i n c u l t u r e  vii  81 ,87  List  1.  Exposed root tomentosus  2.  ball  of an i n t e r i o r  infection  L o c a t i o n of  of Figures  spruce i n an  I.  center  study s i t e s  23  i n the P r i n c e George F o r e s t  Region 3.  32  L o c a t i o n of  study s i t e s  i n the  P r i n c e Rupert  Forest  Region  33  4.  L i g h t Red S t a i n  5.  Reddish-brown s t a i n i n g and e a r l y tomentosus  i n root  infected  i n f e c t e d with  tomentosus  p i t t e d decay i n  41  I.  roots  ...42  6.  Advanced  tomentosus decay  43  7.  Above ground symptoms of an i n f e c t e d p i n e  47  8.  Red s t a i n i n g e x t e n d i n g  53  9.  Correlation proportion  10.  of  of  Quantitative  from an o l d root wound  the p r o p o r t i o n  of p i n e  i n f e c t e d with  spruce i n f e c t e d differential  variation  61 i n host s p e c i e s  susceptibility 11. Q u a l i t a t i v e  the  64 differential  variation  i n host s p e c i e s  susceptibility  64  12a.  tomentosus  in c u l t u r e  - Type 1  85  12b.  tomentosus  in c u l t u r e  - Type 2  85  12c.  tomentosus  in c u l t u r e  - Type 3  86  13.  Rudimentary  tomentosus f r u i t i n g  viii  bodies  88  Acknowledgement  I  wish to thank my w i f e ,  inspiration.  ix  Maureen,  in a l l  t h i n g s my  I.  Inonotus (Polyporus)  INTRODUCTION  tomentosus ( F r . ) Teng i s a  b a s i d i o m y c e t o u s fungus which a t t a c k s t h e l i v i n g r o o t s of c o n i f e r o u s t r e e s i n N o r t h A m e r i c a and a b r o a d . W h i l e a t t a c k s are b e l i e v e d t o o r i g i n a t e i n the r o o t s , i n f e c t i o n  often  e x t e n d s i n t o t h e b u t t as w e l l . I n f e c t i o n l e a d s t o n e c r o s i s of t r e e t i s s u e . Subsequent d i g e s t i o n of h o s t t i s s u e r e s u l t s in a p i t t e d r o t . Since l i v i n g t i s s u e s are attacked, I . tomentosus q u a l i f i e s a s a r o o t and b u t t r o t t i n g  pathogenic  s p e c i e s . O t h e r r o o t r o t s p e c i e s such as A r m i l l a r i a (Pers.) Herink  and P h e l l i n u s w e i r i i  (Murr.) G i l b e r t s o n have  been e x t e n s i v e l y s t u d i e d and a r e c o n s i d e r e d important  obscura  t o be t h e  r o o t r o t s i n B r i t i s h C o l u m b i a . The i n c i d e n c e and  i m p o r t a n c e o f K. tomentosus i n B r i t i s h C o l u m b i a has y e t t o be d e t e r m i n e d . E a r l y e s t i m a t e s ubiquitous nature  of i t ' s i n c i d e n c e , i t ' s  i n sub-boreal  and b o r e a l f o r e s t s , and i t ' s  wide g e o g r a p h i c range s u g g e s t I ^ tomentosus may be t h e most important  r o o t r o t i n Canada i n terms o f t i m b e r volume l o s t .  C l e a r l y , as northern  interior  f o r e s t s become more  i n t e n s i v e l y managed, t h e r o l e p l a y e d by 1^ tomentosus must be c l a r i f i e d . The i n c i d e n c e , d i s t r i b u t i o n and t r e e a f f e c t e d must be d e t e r m i n e d b e f o r e  l o s s e s t o t h e d i s e a s e and  p o s s i b l e c o n t r o l methods can be c o n s i d e r e d management p l a n s .  1  species  i n timber  2 The  purposes 1.  tomentosus  To  of  this  d e s c r i b e the  infected  interior  e n g e l m a n n i i Engelm.) and latifolia  area  To  To  the  to t h i s  spruce  in I.  (Picea glauca pine  x  (Pinus c o n t o r t a  var.  pathogen.  4.  describe  To  relative  susceptibilities  of  pathogen.  determine  i n the  whether  the  host  effect  of  specificity  infection  is  on  basal  increment. 5.  culture  and  geographic  describe characteristics  region  REVIEW OF  1.  TAXONOMY  The  To  attempt  A.  THE  to  relate  taxonomic  history  (Polyporus)  t o m e n t o s u s was  in  1821  Haddow  in his  tomentosus. F r i e s later  to  were  of  the  the  fungus i n  host  species  or  obtained.  LITERATURE  by  Sweden  these  from which they  some d e t a i l  he  symptoms p r o d u c e d  lodgepole  determine  conifers 3.  present  were:  Dougl.). 2.  northern  study  of  1^  (1941) a n d first  t o m e n t o s u s was Gosselin  reviewed  (1944).  d e s c r i b e d by  Inonotus  Fries  "Systema m y c o l o g i c u m " as  circinatus.  in  Polyporus  a l s o d e s c r i b e d Trametes c i r c i n a t u s  (1863) r e v i s e d t o P o l y p o r u s  in  The  which  3  d i s t i n c t i o n between  these two f u n g i was made on the b a s i s of  gross m o r p h o l o g i c a l f e a t u r e s the c o n t e x t  structure  tomentosus and duplex depth of  the  sporophore; e s s e n t i a l l y  (supposedly homogenous i n P. i n E\ c i r c i n a t u s ) ,  the pore l a y e r  Historically,  of  (greater  and the  relative  i n P_j_ c i r c i n a t u s ) .  t h e r e has e x i s t e d a great d e a l of  c o n f u s i o n r e g a r d i n g the proper taxonomic s t a t u s of fungus.  In Europe the  this  fungus was r e c l a s s i f i e d s e v e r a l  The genus was changed to P o l y s t i c t u s by Cooke ( 1886) Saccardo (1888). K a r s t e n  (1889) p l a c e d the  times. and  species into  the  genus O n n i a . Two s p e c i e s , tomentosus and c i r c i n a t u s were r e c o g n i z e d by most a u t h o r s .  In N o r t h A m e r i c a , a  similar  c o n d i t i o n e x i s t e d w i t h s e v e r a l a u t h o r s r e c l a s s i f y i n g the f u n g u s . E l l i s and E v e r h a r t Mucronoporus, M u r r i l l i n t o P o l y s t i c t u s . In  (1889) p l a c e d i t  i n t o the genus  (1908) i n t o C o l t r i c i a and L l o y d (1908) fact,  some a u t h o r s b e l i e v e d  that  P o l y p o r u s tomentosus d i d not o c c u r i n North America at Instead,  closely related  s p e c i e s , P_j_ d u a l i s P e c k , and  P. c i r c i n a t u s were d e s c r i b e d (Peck  1878). In a c t u a l i t y ,  a u t h o r s gave P_j_ d u a l i s s p e c i e s s t a t u s . relegated  it  to v a r i e t y  all.  Instead  s t a t u s or b e l i e v e d i t  few  they t o be  synonymous w i t h P. tomentosus or P_j_ c i r c i n a t u s  .  Peck h i m s e l f  Polystictus  circinatus  came t o r e g a r d i t  (Haddow  1941). T a b l e  as a v a r i e t y  of  Eventually,  1 p r o v i d e s an overview  of  4 the  nomenclatural  changes under  Haddow a t t r i b u t e d proper  type  sporophore depth  and h a b i t ,  of the pore  layer.  number  of specimens  Haddow  discovered that  the  in early  used  found  stratification  present  setae  in  o f t h e c o n t e x t and  Through an e x a m i n a t i o n by t h e a u t h o r s  the context  descriptions  listed  of a g r e a t i n Table 1  was, i n a l l c a s e s  arose  t o be t h e o n l y d i a g n o s t i c  setae  t o a lack of  d e a l of v a r i a t i o n i n  layers.  feature,  The  with  setae  straight  i n P_;_ t o m e n t o s u s v a r . t o m e n t o s u s a n d c u r v e d  t o m e n t o s u s v a r . c i r c i n a t u s . Haddow  tomentosus, c i r c i n a t u s Polyporus  duplex.  due t o v a r i a t i o n i n  r e l a t i v e t h i c k n e s s o f t h e two c o n t e x t  were  1849 and 1944.  of t h i s c o n f u s i o n  specimens and t o a g r e a t size  Confusion  most  gone b e t w e e n  and d u a l i s  tomentosus with  as a s i n g l e  i t ' s variety  reclassified species:  c i r c i n a t u s . P.  dualis  was deemed  synonymous t o P_j_ t o m e n t o s u s v a r . c i r c i n a t u s .  Gilbertson  (1974) t r a n s f e r r e d  Inonotus the  with  shift  two s p e c i e s ;  t o the genus  some p r o m i n e n t a  variety  authors  the fungus  circinatus  fungus  yet  been  into  has g e n e r a l l y  continue  to classify  t h e genus Onnia  adopted.  t h e genus  and tomentosus.  Inonotus  o f I_j_ t o m e n t o s u s . J a h n  the  into  ( 1978)  been  While  accepted,  I_j_ c i r c i n a t u s a s  proposed  transfer of  but t h i s p r o p o s a l has not  TABLE  Historical  1  Nomenclature of Inonotus  F r i e s 1821 F r i e s 1849 Peck 1878 Cooke 1886 S a c c a r d o 1888 K a r s t e n 1869 E l l i s and Ever h a r t 1689 Hennings 1898 P a t o u l i l a r d 1900 M u r r i l l 1904 Haddow (1908) L l o y d 1908 and 1912 Ames 1913  P_: P. P. P. P. O.  circinatus circinatus dualis circinatus circinatus circinatus  Mucroporus tomentosus P o l y s t l c t u s tomentosus Xanthochrous c i r c i n a t u s  M. P.  circinatus circinatus  C o 1 1 r 1 c i a tomentosa P o l y p o r u s tomentosus P. c i r c i n a t u s P o l y s t i c t u s tomentosus  I . tomentosa P. d u a l l s  tomentosus  (xarthochrous ) circinatus  K i l l e r m a n 1928  Polystlctus  Shope 1931  Polyporus  dualis  O v e r h o l t s 1933  Polyporus  circinatus  tomentosus  Lowe 1934 Jorstad + Juul 1938  P. c i r c i n a t u s P. t o m e n t o s u s P. t o m e n t o s u s var. amerlcanus  Haddow 1941  Polyporus 1974  Polystlctus circinatus P. d u a l i s Polyporus dualls P_. t r l q u e t e r JL* tomentosus var circinatus Fomes  B o u r d o t and G a l z i n 1927  Gilbertson  5  P o l y p o r u s tomentosus P o l y p o r u s tomentosus P o l y p o r u s tomentosus P o l y s t I c t u s tomentosus P o l y s t l c t u s tomentosus Onnla tomentosa  Neuman 1914 B r e s a d o l a 1920  S a r t o r y and M a i r e 1922 Konrad and Maublanc 1926  tomentosus.  Inonotus  tomentosus tomentosus  Polyporus circinatus Polystlctus circinatus var. triqueter  P. c i r c i n a t u s  5ualIs  P. d u a l i s P.dualis  i  P.tomentosus var. c i r c i n a t u s I. c i r c i n a t u s  6 2. DISTRIBUTION  Inonotus  AND HOST SPECIES  tomentosus i s an u b i q u i t o u s  f u n g u s . It  been found in N o r t h A m e r i c a , Europe and A s i a . America,  tomentosus r o o t  rot  B r i t i s h C o l u m b i a , Northern 1985). Inonotus reported  o c c u r s as f a r  n o r t h as  Saskatchewan and A l a s k a  1983,  Ross  circinatus  1983,  Boyce 1963). Most r e p o r t s  (Barnard et  Northwest  and northwards  Partridge  1979,  al  1985,  refer  are of JN_ tomentosus  circinatus occurs.  i n the  Pacific  (Hobbs and  Whitney  and Van  scale  in disease d i s t r i b u t i o n .  tomentosus may e x i s t  exclusively  from W i s c o n s i n , the  Davidson and Redmond 1957,  geographical trend  (Kossuth  Kossuth and Barnard  Groenewoud 1964). T h i s may p o r t e n d a l a r g e  which o n l y  (Shaw  1966).  to I^  I^  Northern  tomentosus or I_j_ c i r c i n a t u s have been  Reports from t h e most southern S t a t e s  of  North  south t o G e o r g i a and the F l o r i d a p e n i n s u l a  and Barnard  limit  In  has  Some southern  southern U . S .  However,  Northern  o c c u r r e n c e s of J N c i r c i n a t u s have been r e p o r t e d above line  (Van Groenewoud 1956). Thus the  beyond  this  ranges of J N c i r c i n a t u s  and I_j_ tomentosus are not m u t u a l l y  e x c l u s i v e . The ranges of  the two s p e c i e s may become c l e a r e r  in future  classification  to  s p e c i e s (or  s t u d i e s becomes more common.  variety)  if  i n surveys and  7 Reported i n c i d e n c e s i n Europe i n c l u d e Sweden 1863,  L l o y d 1908 and M u r r i l l  Germany  (Murrill  1904). Bakshi  be damaging in Northern tomentosus I.  India.  limit  native  rot  high  attacks  wide geographic range  that a l l  native  Marsh.)  on D o u g l a s - f i r  (Table  and t r e m b l i n g  2).  the  predicted  Whitney  (1977)  reported  successful (Betula  aspen (Populus  tremuloides  and Van Groenewoud 1964). As hardwoods  This p a r a l l e l s  the  and Reynolds (1962) w i t h P h e l l i n u s  (Pseudotsuga m e n z i e s i i  case does the v i a b i l i t y indicate  reported  Gosselin  (1964) found t h a t a wide range of  f i n d i n g s of W a l l i s  I.  a great many t r e e s p e c i e s  c o u l d be used as inoculum c a r r i e r s .  carriers  (1927)  The d i s e a s e has not been  (Whitney 1964b, Whitney  w e l l Whitney  neither  reported  to  altitudes.  i n o c u l a t i o n s have been made on white b i r c h  Michx.)  tomentosus  he a l s o  any hardwood s p e c i e s . However,  papyrifera  and  Canadian spruce s p e c i e s and most  p i n e s are a t t a c k e d .  attacking  1920)  appeared t o be  of the s p e c i e s . However,  Tomentosus root  reports  L l o y d (1920),  36th p a r a l l e l  more southern o c c u r r e n c e s at  it's  (1971) found I.  i n China and Japan r e s p e c t i v e l y .  (1944) observed t h a t the  over  (Lloyd  i n A f r i c a . Teng (1932) and S h i r a i  tomentosus  southern  1908), A u s t r i a  (Fries  an a b i l i t y  (Mirb.)  Franco).  of hardwoods as  by the  In  inoculum  fungus t o a t t a c k  or c o l o n i z e dead hardwoods i n n a t u r a l  weirii  conditions.  living  TABLE 2 Conifer a)  In  Hosts  of  8  I.  tomentosus  Canada: amabi1ls (Dougl.) Forb. balsamea (L.) M i l l . l a s i o c a r p a (Hook.) N u t t . l a r i c i n a (Du R o i ) K. K o c h o c c i d e n t a l i s Nutt. abies Karst engelmanni i P a r r y g l a u c a (Moench) V o s s g l a u c a (Moench) V o s s v a r . A l b e r t i a n a (S.Brown) S a r g . P l c e a marlana ( M i l l . ) B.S.P. P i c e a rubens Sarg. P l c e a s l t c h e n s l s (Bong.) c a r r . P l n u s b a n k s l a n a Lamb. Plnus c o n t o r t a Dougl. Pinus c o n t o r t a Dougl. v a r . l a t i f o l i a Engelm. Pinus monticola Dougl. P i n u s p o n d e r o s a Laws. Pinus r e s i n o s a A i t . P i n u s s t r o b u s L. PseHdotsuqa m e n z e i s i i (Mirb.) Franco T h u j a p l i c a t a Donn ex D. Don Tsuga c a n a d e n s l s (L.) C a r r . Tsuga h e t e r o p h y l l a (Raf.) S a r g . Abies Abies Abies Larix Larix Plcea Picea Picea Picea  b)  Host  s p e c i e s abroad  (other than  those  listed  A b i e s p r o c e r a Rehd. C e d r u s d e o d o r a Roxb. Picea abies Karst. P i c e a pungens Engelm. P i c e a rubens Sarg. P i c e a smythiana B o i s s . Plnus e l l i o t i l Engelm. Pinus nigra Arnold P i n u s r a d i a t a D.Don. Plnus r l g l d a M i l l . P i n u s s v l v e s t r l s L. P i n u s t a e d a L. P i n u s c l a u s a v a r . immuginata  From Whitney  1977  and  Whitney  1978.  above):  9 While  these  equally. trees  species  Inoculation experiments  h a v e y i e l d e d an e s t i m a t e  susceptibility  of a small  1976,  Bohaychuk From  are attacked,  these  Whitney  highly  susceptible  1977, W h i t n e y  has  g e n e r a l l y been  species'  potential  impossible to  susceptibility be  site  may be t h a t  is  black  sites.  tomentosus  Lamb.), w h i t e (P^ mariana  Some  idea  species  i n d i c a t o r s of a  i t i s d i f f i c u l t or infection  hazard  the absolute  low i n f e c t i o n  and r o o t  spruce  (F\ glauca  (Mill.)  l e v e l s may interaction  disease  of lodgepole  of t h e i r  i t i s most  these  surveys.  on t h e f u n g u s o r t o an  the predominance  where  from  of host  do n o t a c c u r a t e l y r e f l e c t  due t o a c o i n c i d e n c e  less  A t a n y r a t e , low l e v e l s o f  of a s p e c i e s . Such  spruce  Aside  incidence  to extrapolate  f e a t u r e s on t h e h o s t  (P. b a n k s i a n a and  data  due t o h a b i t a t e f f e c t s  between It  on o t h e r  certainly  are valid  1964b).  sp. ) a r e  somewhat  1962).  from  susceptibility,  t o use such  the species  infection  levels  (Larix  susceptibility  extrapolated  incidence  {Whitney and  1977, W h i t n e y  and tamarack  i n o c u l a t i o n experiments,  high  of s p e c i e s  s u s c e p t i b l e and p i n e s  (Whitney  and o l d e r  of the r e l a t i v e  and Bohaychuk  few  While  on s e e d l i n g s  number  s t u d i e s , spruce  considered  not a l l a r e impacted  pine,  species. jack  (Moench) V o s s )  B.S.P.) a s h o s t  ranges with  pine  species  t h e range  of I .  virulent.  of t h e r e l a t i v e  importance  of  tomentosus  10. to the d i f f e r e n t  conifer  p r o p o r t i o n of t h e i r Canada,  decay a t t r i b u t a b l e  tomentosus i s a major  and l o d g e p o l e p i n e found  (Table  3).  pathogen  i n jack p i n e and f i r s t  results  i n white spruce  Basham and Morowski  (1964)  important  in a limited  red pine  (1978) o b t a i n e d  (P.  similar  f o r white and b l a c k spruce i n n o r t h e r n O n t a r i o . The  low p r o p o r t i o n of I^ listed  In  decay c a u s i n g fungus i n white and b l a c k s p r u c e ,  r e s i n o s a A i t . ) s t u d y . Whitney  tomentosus decay on the other  i n T a b l e 3 may r e f l e c t  their  lesser  due to some r e s i s t a n c e mechanism. It species r e f l e c t  limited  geographic r a n g e s .  species  susceptibility  may a l s o ,  f o r some  o v e r l a p of the h o s t ' s and p a t h o g e n ' s  Inonotus  tomentosus i s c h a r a c t e r i z e d as a  d i s e a s e of s u b - b o r e a l and b o r e a l the  to the d i s e a s e .  tomentosus to be the second most  identified third  s p e c i e s can be d e r i v e d from the  f o r e s t s . T h i s may r e f l e c t  range of the most s u s c e p t i b l e host s p e c i e s or i t may  reflect  the o p t i m a l  and b i o t i c )  growing c o n d i t i o n s ,  of the f u n g u s . If  the l a t t e r  (climatic, is true,  t r e e s p e c i e s most h e a v i l y a t t a c k e d are j u s t where they a r e g r o w i n g . T h i s i d e a limited  o c c u r r e n c e of I^  and P a r t r i d g e  (1979),  higher e l e v a t i o n s disease  abiotic then the  artifacts  i s s u p p o r t e d by the  tomentosus i n Northern  Idaho.  found the d i s e a s e t o be l i m i t e d  (above  of  Hobbs to  1500 M ) . In B r i t i s h C o l u m b i a , the  i s not known i n low e l e v a t i o n  l o d g e p o l e p i n e or c o a s t a l shore p i n e  southern  interior  f o r e s t s . However,  at  TABLE 3  11  F r e q u e n c y of i s o l a t i o n o f t o m e n t o s u s as a p e r c e n t of a l l f u n g a l i s o l a t e s i n C a n a d i a n d e c a y s t u d i e s .  SPECIES  LOCATION  Abies amabl1ls (Dougl.) Forb.  British Columbia  1.0%  Buckland 1949  Abies balsamea (L.) M i l l .  Eastern Canada  1.5%  Basham e t a l 1953  Ontar i o  1.7%  Whitney  Alberta Ontario  85.0% 16.2%  Denyer and R i l e y 1953 W h i t n e y 1978  Ontario  17.9%  Whitney  Pinus contorta Dougl. var l a t i f o l i a Engelm.  Alberta  14.0%  Denyer and 1953  Pinus  Ontario  1.3%  White  Pseudotsuqa menziesii (Mirb) Franco  British Columbia  0.3%  Thomas and Thomas 1954  Tsuga heterophylla (Raf) Sarg.  British Columbia  1.4%  Plcea glauca (Moench) v o s s P i c e a mariana ( M i l l . ) B.S.P.  s t r o b u s L.  % OF TOTAL INFECTIONS  0.8%  From M e r l e r , 1984  AUTHOR  et a l  1978  1978 Riley  1953  Buckland e t a l 1949 F o s t e r and F o s t e r 1951  1 2  higher fir,  elevations,  the author  spruce and p i n e .  average  levels  species, levels level  of  occur and i f data  of  (1962),  In  19.7%  examined were i n f e c t e d w i t h Merler  (1984) r e p o r t e d  Columbia and Whitney  It  were,  in  that  infection  central  interior  incidence l e v e l s  studies fact,  This prior  which d i s q u a l i f i e s  however,  the  infection  More of  28.4 %  British l e v e l s of  up  Table 4  t h a t many of of  these  intrinsic  disease i t s e l f .  knowledge of  these  spruce  found i n Canadian spruce s t a n d s .  i n c l u d e an element  s t u d i e s of  be  levels of  in  incidence  of a l l  stands known to be i n f e c t e d were s e l e c t e d , surveyed.  incidence  tomentosus.  (1986) r e p o r t e d  s h o u l d be n o t e d ,  incidence  incidence  fact,  27% i n e a s t e r n Canadian spruce f o r e s t s .  summarizes  tree  stands may not  reported  i n spruce i n stands i n the  to  damaging  the  s c a r c e f o r most host s p e c i e s . However,  20% or more i n a f f e c t e d  recently,  these  spruce, studies indicate  uncommon. Whitney trees  in stands of whether  Douglas-  surveys have been performed on spruce stands  North A m e r i c a . In levels  indicate  attacking  says n o t h i n g of  s o , how f r e q u e n t l y .  is fairly  a number of  This information  the d i s e a s e  nor does i t  has found i t  infection  b i a s . Some  Accordingly,  s t u d i e d and assures bias  i n c i d e n c e l e v e l s as i n d i c a t o r s  of  Table  Incidence  Levels  Interior  of  4  13  I ^ tomentosus  Spruce  on  i n Canada  Species  Location  Percent Infected  Author  W.  Pr.Rupert  14%  Kondo  Spruce  Taylor, Spruce  Sask .  19.7%  Spruce  Pr.  Georqe  28.4%  Spruce  Pr.  Rupert  5%  Unger,  W.  Ontar i o  27%  Whitney,  BC/Yukon  35%  Yukon  33%  Spruce  Spruce  McLeod,  B.C.  spruce  N.  Ontario  18%  w.  spruce  N.  Ontario  14%*  various  aaes.  infection  Whitney, *  *  1962  Merler,  1964 1984 1986  Wood, Van and  Sickle  Shore,  1984  55%  B.  * Represents averaqe  1965  *  levels  Whitney  1978  Whitney  1978  calculated  t o r s t a n d s of  1 4  average  infection  regions.  As w e l l ,  levels  f o r stands i n t h e i r  severely  infected  stands are more  to c a t c h the a t t e n t i o n of those p e r f o r m i n g rate,  a certain  amount  d i s e a s e behaviour indications they  of the average  idea of the p r o p o r t i o n infection  stands i n f e c t e d , accurately Sickle  the s t u d y . At any  of d i s e a s e i s r e q u i r e d  studies e f f i c i e n t .  likely  to make  Thus these  f i g u r e s are  of the l e v e l s which can be found i n s t a n d s , but  say n o t h i n g  average  respective  i n c i d e n c e . They a l s o g i v e no  of stands i n f e c t e d .  level  of  tomentosus cannot  Kondo and T a y l o r  (1984) are r e p o r t s  (1985) and Van  of the Canadian F o r e s t  F o r e s t I n s e c t and D i s e a s e S u r v e y . As such they i n c i d e n c e s but say n o t h i n g They a l s o f a i l  to report  an  f o r a l l s t a n d s , or the p r o p o r t i o n of  the importance  be a s s e s s e d .  Without  Service  report  of the stand s e l e c t i o n  how many stands  (if  process.  any) were  surveyed and found t o be f r e e of the d i s e a s e . Without information,  a g a i n no statements  r e g a r d i n g average  this  infection  l e v e l s can be made.  More severe problems e x i s t incidence studies seems p i n e the  for pine  i n p i n e s are more r a r e .  s p e c i e s are i n c l u d e d i f  surveys s i n c e Most o f t e n ,  it  they occur with spruce in  stands s t u d i e d . The e x c e p t i o n s t o t h i s would f o r the  most p a r t be s t u d i e s on I^ southern U n i t e d S t a t e s .  circinatus  on hard p i n e s i n the  Unger and .Stewart  ( 1986) surveyed  15 l o d g e p o l e p i n e stands i n the c e n t r a l Columbia f o r  tomentosus root  rot.  high i n f e c t i o n l e v e l s reported  interior  However,  of  the  British  extremely  ( a v e r a g i n g 87% i n pure pine  s t a n d s and 90% i n mixed pine and spruce s t a n d s ) , c o u p l e d w i t h v e r y low a s s o c i a t e d m o r t a l i t y q u e s t i o n the  rates  survey methods used and the  assessment and pathogen i d e n t i f i c a t i o n Details  of  l e a d the author infection  techniques  these methods were not s u p p l i e d i n the  status utilized. report.  F u r t h e r q u e s t i o n of t h e s e r e s u l t s i s i n t r o d u c e d by the t h a t the e t i o l o g y Phellinus pini  for  the pathogen appears t o  (ThorerFr.)  P i l . more than i t  to  fact  resemble does  tomentosus.  The o v e r a l l e f f e c t susceptibility It  to  of t r e e v i g o u r on host  tomentosus i s not c l e a r l y u n d e r s t o o d .  has been shown t h a t low t r e e v i g o u r r e s u l t s i n  susceptibility Gosselin  t o r o o t d i s e a s e (Whitney 1977).  (1944),  i n f e c t i o n rates  found f a s t e r than t h e i r  growing t r e e s  However,  t o have h i g h e r  slower growing c o u n t e r p a r t s . A  " " p o s i t i v e c o r r e l a t i o n between  i n f e c t i o n r a t e and host  r a t e may be due to more e x t e n s i v e r o o t systems i n growing  (and thus l a r g e r )  greater  probability  heightened  trees.  Larger root  growth  faster  systems have a  of c o n t a c t i n g an inoculum s o u r c e .  E v i d e n c e r e g a r d i n g p r e d i s p o s i t i o n through s t r e s s t o tomentosus may be c o n f u s e d by the d i r e c t  effects  of  I. the  16 stress  i n d u c i n g agent  collar  weevil,  this.  While  whether  H y l o b i u s warreni  tomentosus and a t t a c k (Krebill  1962,  this correlation  t o the p r o v i s i o n of correlation  Whitney  (through  by the  by H_;_ w a r r e n i  1961),  it  at  i s not  i s due t o weakening of  the  clear  the t r e e or  confounded by the q u e s t i o n of which  pest a r r i v e d on the scene f i r s t .  attack  between  i n f e c t i o n c o u r t s f o r b a s i d i o s p o r e s . The  is further  predisposing  The root  Wood i s a good example of  s t u d i e s have shown a c o r r e l a t i o n  i n c i d e n c e of root c o l l a r  on i n f e c t i o n p r o b a b i l i t y .  stress)  E i t h e r p e s t c o u l d be  the host f o r more  successful  other.  3 . METHODS OF SPREAD  The p r i n c i p a l method of d i s e a s e spread o c c u r s v i a c o n t a c t s between h e a l t h y  r o o t s and i n f e c t e d r o o t s or stumps  (Whitney 1977). T h i s spread mechanism a l s o a l l o w s of  infection  i n t o the next  rotation.  Such t r a n s f e r  transfer occurs  r e a d i l y where f u n g a l mycelium i s p r e s e n t on or i n the of the  In for  i n f e c t e d root  artificial  (Myren and P a t t o n  bark  1970).  i n o c u l a t i o n s , r o o t wounds were n e c e s s a r y  successful transfer  of mycelium (Whitney 1962,  1964). T h i s does not appear to be the case i n conditions.  root  1963,  natural  The n e c e s s i t y of wounds i n a r t i f i c i a l  conditions  17 may  be  due  to the  pathogen  i n these  Although  not  promote  inferior  experiments  strictly  infection areas  infection  (Krebill  the  of  the  site  o f most  roots are in  the  early  fall,  abundance. evidence I.  them  sufficient  sporophores Sporophores  Infection The slim  chances given  likely  by  to occur  production  be  of  sporophores,  facilitating  stipitate  or  habit  Inonotus  while  two  result  spread  an  has  late  sessile. may  differ  is  between  circinatus  not  spore  been  tends  produces  demonstrated.  however, a r e induced  i n a g i v e n s t a n d . The the  There  1977).  event,  o f new  of  1961).  tomentosus  (Whitney  in nature  a year  Where  in  sporophores  over  c o n t a c t s are  wound c a n  Whitney  to  least  b a s i d i o s p o r e s are produced  sporophore  s m a l l number  at  to  of o b s e r v i n g such the  be  summer  sporophores  spores  susceptible  root  i n the  to  providing  transfers.  1957,  circinatus.  sessile  stipitate  can  that  t o m e n t o s u s and  mainly  and  believed  T h i s may  appressed  precipitation  1970).  wounds a r e  1961).  further  the  Patton  a compression  (Redmond  of  which are  inoculum  appressed  to suggest  to produce  tightly  of c o n t a c t , thus  between  Given  why  root  tissue  Whitney  successful  tightly  area  mycelium  reason  and  c o n d i t i o n s by  of d e c l i n i n g 1962,  potential  (Myren  necessary,  in natural  localized  part  inoculum  ubiquitous nature  very  infections continued of  this  18 fungus and the presence of new i n f e c t i o n c e n t e r s b e a r i n g no apparent  historical  infection However, event  infection  sources,  suggest t h a t some  c e n t e r s must be i n i t i a t i e d by b a s i d i o s p o r e s . the  frequency  (and thus the  i s o f t e n q u e s t i o n e d . There  importance)  of  i s some i n d i r e c t  this  evidence  which suggests spread by s p o r e s may be a common e v e n t . Inonotus  tomentosus has been r e p o r t e d  plantations cultivated  in Wisconsin a l l fields  reported  in t h i s  of which were l o c a t e d on o l d  (Barnard et  i n f e c t i o n and the number of study,  al  infection  i n the absence of  b a s i d i o p o r e s p l a y an important  question.  centers  infected  stumps or  s t r o n g l y suggest t h a t and f r e q u e n t  role  i n spread  i n f e c t i o n by b a s i d i o s p o r e s i s a l s o  Cut stumps have not been shown t o be  (Barnard e_t al_ 1985,  Whitney  i n o c u l a t i o n s t u d i e s , wounding of deeper was found n e c e s s a r y f o r basidiospores  (Whitney 1963,  of these r e s u l t s to n a t u r a l  1966).  In  Animal t r a m p l i n g ,  in  infection artificial  r o o t s to the cambium or  successful  Whitney  inoculation  1966). If  with  extrapolation  c o n d i t i o n s i s v a l i d , a source of  wounding would be r e q u i r e d f o r b a s i d i o s p o r e  root  of  fungus.  The avenue of  courts  1985). The l e v e l s  individual  r o o t s from a p r e v i o u s r o t a t i o n ,  of t h i s  i n a number of  infection.  compression wounds, e x c e s s i v e m o i s t u r e ,  f e e d i n g o r g a n i s m s , b a s a l c a n k e r s of  Cronartium  19 fusiforme  (Cumm.)  Hedge, and Hunt and root deformation  to improper p l a n t i n g  p r o c e d u r e s have been suggested as  s u i t a b l e wound c a u s i n g agents Whitney  1961,  Boyce  I n f e c t i o n of  (Redmond 1957,  to be washed or o t h e r w i s e  subterranean  feeding  soil is clearly  H e t e r o b a s i d i o n annosum ( F r . ) viable  i n the s o i l  insects  Bref.  Spore  r a i n w a t e r . The  survival evident.  soil.  (especially  s q u i r r e l s , v o l e s or j u s t  c a p a c i t y of these b a s i d i o s p o r e s f o r i n t o the  during B a s i d i o s p o r e s of  can s u r v i v e and remain  (Kuhlman 1969). Whitney  (1966),  reported  t h a t s p o r e s of I_j_ tomentosus remained v i a b l e a f t e r at  -18  d e g . C and a f t e r  thawing.  s e v e r a l c y c l e s of  R e s i s t a n c e t o c o l d temperatures  g e r m i n a t i o n and i n f e c t i o n dispersal  i n the  i n the  f a l l and o v e r w i n t e r i n g  s e c o n d ; the c r e a t i o n  1961),  of  i n the  first;  the t r e e ' s  low v i g o u r host t i s s u e  of the wound. K r e b i l l  suggest t h a t the  would a l l o w  for  spore  soil.  spread mechanism would  seem t o be a combination of two f a c t o r s : p r o v i s i o n of a p h y s i c a l avenue i n t o  storage  f r e e z i n g and  spring following  The r o l e of wounding i n e i t h e r  vicinity  1962,  r o o t s r e q u i r e s spores  c a r r i e d through the  c a r r i e r s c o u l d i n c l u d e root  transport  Krebill  1963).  injured  Hylobius warreni),  due  the t i s s u e and in  (1962) and Whitney  the (1960  and  r o l e of wounds i n p r o v i d i n g avenues  20 of  infection  the  seems t o l i e i n t h e l a t t e r  factor  rather  that  former.  4. ECOLOGY  Data from  regarding  observations  studies  of t h e fungus  i n the f i e l d .  w h i c h must  Both  be e x t r a p o l a t e d  Important are  the autecology  circinatus (Whitney  h a v e been  1962b). Whitney  temperature mycelial  humus  s t u d i e s have  sandy  form),  indirect  ecology  mycelium  evidence  i n nature.  growth  in culture  tomentosus and I .  t o grow b e s t  and Bohaychuk  on a c i d i c (1977),  media  found  that a  20 d e g r e e s C. was b e s t f o r  Thick,  linked  a number o f s i t e  o f I_j_ t o m e n t o s u s soils  root  disease.  moist  complex  features to Generally,  w h i c h h a v e a t h i c k humus l a y e r (mor  low pH, a r e e x c e s s i v e l y d r a i n e d  to p e r i o d i c drought  Hylobius  to fungal  of a p p r o x i m a t e l y  incidences  shallow,  provide  growth.  Field high  sources  In c u l t u r e ,  found  comes  i n c u l t u r e and i n c i d e n c e  factors affecting  pH a n d t e m p e r a t u r e .  o f I_j_ t o m e n t o s u s  are high  (mor) humus (Warren  risk  areas  forms  (Whitney  favour  1956). Thus,  and a r e s u b j e c t 1962).  b e e t l e s of the  a mor humus m i g h t  21 therefore  affect  tomentosus  on the number of avenues of humus forms are o f t e n  i n c i d e n c e through  infection.  indicative  of  its  effects  Alternatively,  slow n u t r i e n t  mor  turnover  and c o n s e q u e n t l y poor s o i l n u t r i e n t c o n d i t i o n s . Such c o n d i t i o n s c o u l d cause t r e e s t r e s s which may p r e d i s p o s e them to  infection.  s t r e s s the  In  fact,  a t t a c k by  t r e e and cause p r e d i s p o s i t i o n . Thus the mechanism  behind t h i s  association is unclear.  type and water d e f i c i e n t  fungal  ecology.  s h a l l o w n e s s p r o b a b l y has i t s main e f f e c t  providing a higher proportion (especially of  F a c t o r s such as humus  s o i l s may r e l a t e more t o  s t r e s s p r e d i s p o s i t i o n than t o soil  warreni c o u l d a l s o  the  of  compression wounds)  fungus  (Myren and Patton  root  the  found t h a t I^ temperatures  damage  were most  culture. heavily  distribution  demonstrated.  advantage  has been p o s t u l a t e d as an the  fungus'  (Hobbs and P a r t r i d g e  effects While  of  showed  (Van Groenewoud 1956). Whitney  below 20 degrees C . T h i s a b i l i t y  in determining  behind the  fungus i n  tomentosus had a c o m p e t i t i v e  temperatures  spread  findings  w h i l e those w i t h a s o i l pH of 7 or h i g h e r  no s i g n i f i c a n t  factor  through  c o n t a c t s and wounds  1971). The  Stands w i t h low s o i l pH (down to 4.5)  lower  Similarly,  and thus f a c i l i t a t i n g  r e g a r d i n g pH agree w i t h s t u d i e s of  afflicted  tree  (1962), at  to t h r i v e  at  important  geographic and  elevational  1979). The mechanisms  s o i l t e x t u r e have not been  sandy s o i l s r e p r e s e n t  clearly  h i g h hazard  the  22 reason or  for this  availability  the  associated  their  tree  a i r or  nutrient  s t r e s s or  It could  other  level. on  be  pore Such  fungal  microbial  related to  size  drainage  effects including  factors could vitality  or  competition  exert  indirectly  effects.  D A M A G E  Direct  damage by  live  windthrow,  butt  cull.  a.  of  influence directly  through  5.  i s unclear.  I ^ tomentosus can  standing  mortality,  take  four  increment  forms:  reduction  and  WINDTHROW  Live  windthrow  of  tomentosus  results roots  offer  little  falling, (Figure  decay  field. of  i n d i c a t o r of  W i n d t h r o w due  major  supporting  little  s t r u c t u r a l support.  of  tree  the  wind. T h i s  w h i c h have  predominant  i n the  from the  instability  gale  i s the  fallen  along  results in  eventually  The  the  to  presence,  root  roots.  disease  Rotten  resulting  i t s toppling  r e s u l t s in areas  in  of  very  windthrows  in a l l d i r e c t i o n s . This multi-directional  with  the  small  1) d i s t i n g u i s h e s r o o t  amount  of  disease  root  induced  mass  exposed,  windthrow  winds.  In  spruce,  windthrow  can  occur  with  little  apparent  from  THE QUALITY OF THIS MICROFICHE IS HEAVILY DEPENDENT UPON THE QUALITY OF THE THESIS SUBMITTED FOR MICROFILMING.  LA QUALITE DE CETTE MICROFICHE DEPEND GRANDEMENT DE LA QUALITE DE LA THESES SOUMISE AU MICROFILMAGE.  UNFORTUNATELY THE COLOURED ILLUSTRATIONS OF THIS THESIS CAN ONLY YIELD DIFFERENT TONES OF GREY.  MALHEUREUSEMENT, LES DIFFERENTES ILLUSTRATIONS EN COULEURS DE CETTE THESES NE PEUVENT DONNER QUE DES TEINTES DE GRIS.  23  Figure 1 Exposed root b a l l of an i n t e r i o r infection center.  spruce i n an I_.  tomentosus  24 reduction  i n crown v i g o r .  Trees  apparently  quite  large  part  t o the p a t t e r n of decay  Quite  often,  thin  shell  (Merler  extensively  of l i v i n g  1984).  produced  merchantable estimated  admittedly  killed  time  annual for  (Merler  spruce. such  have a  induced  from  ever  (1984)  windthrow  o f B.C. T h i s e s t i m a t e from  of t h i s  was  one y e a r ' s o b s e r v a t i o n s  of the r a t e  induced m o r t a l i t y  losses  xylem  on t h e  of windthrow  (1978),  reported  and windthrow  o l d b l a c k s p r u c e a n d 1.1% f o r 64 y e a r  Prolonged  stands  still  roots.  1984).  tomentosus  c o u l d n o t be o b t a i n e d . W h i t n e y  75 y e a r  spruce  i s d i f f i c u l t . Merler  a s i t was made  I_j_ t o m e n t o s u s  i s due i n a  r o o t s may be  of windthrow  to  i n the i n t e r i o r  crude  i n large  roots w i l l  t h e r e f o r e an a c c u r a t e p i c t u r e  over  b.  loss  This  adventitious  of t h e impact  1% a n n u a l  fall.  green and  surrouding the decayed  volume o f a s t a n d  s i x stands  and  tissue  to r e p l a c e those  they  decayed  In a d d i t i o n ,  Assessment  in  h e a l t h y when  are often  a t 2.7%  o l d white  magnitude c o u l d prevent  producing a merchantable  yield.  STANDING MORTALITY  Standing  mortality  has  killed  been  the moisture tree.  o c c u r s when t h e r o o t  system  t o an e x t e n t w h i c h p r e v e n t s  and n u t r i e n t  Such m o r t a l i t y  will  of the tree  i t from  absorption requirements o b v i o u s l y be h a s t e n e d  on  meeting  of the droughty  25 s i t e s or abiotic  if  the t r e e  i s otherwise  a g e n t s . Whitney  s t r e s s e d by b i o t i c  (1978) l i s t s  s o i l s w i t h deep humus that are  shallow,  subject  acid,  as being most conducive to s t a n d i n g  However,  these s o i l f a c t o r s are p r o b a b l y promoting  promote  (soil)  i n c r e a s e d root wounding.  are c o n d u c i v e t o  tomentosus  (Whitney  1960). These f a c t o r s l i k e l y  outright  mortality  exert t h e i r  through promotion of  infection  pathogen s p r e a d . Sandy s o i l s may a c t u a l l y s o i l nutrient  content  mortality  on the t r e e .  r a t e of m o r t a l i t y  Attack  to  Acid growth effects and  These f a c t o r s  through s t r e s s  by bark b e e t l e s or other  insect  p e s t s c o u l d a l s o be expected t o hasten t r e e m o r t a l i t y . mortality  may,  i n some i n s t a n c e s be a t t r i b u t e d  more o b v i o u s (above ground) result  c.  in underestimation  pest.  of the  on  i n d i c a t e a low  and e x c e s s i v e d r a i n a g e .  may be i n f l u e n c i n g the effects  mortality.  ways. Shallow s o i l s have been p o s t u l a t e d  i n f e c t i o n due to  conditions  sandy  to h i g h moisture  deficits  in d i f f e r e n t  or  Such  s o l e l y to  the  Such an assessment c o u l d impact of  tomentosus.  BUTT CULL  Inonotus  tomentosus decay o f t e n extends  the b o l e of  severely  one to two meters four meters substantial.  infected trees.  into  the b u t t and  Decay commonly extends  i n t o the b o l e and can extend as h i g h as  (Whitney Whitney  1978). Losses due t o decay can be (1978),  found t h a t about  4.4% and  3.2%  26 of  the g r o s s merchantable volume were l o s t t o  butt c u l l  tomentosus  i n n o r t h e r n O n t a r i o b l a c k and white spruce stands  respectively. white p i n e  Losses of t h i s type  (P_j_ s t r o b u s L . )  i n jack p i n e and e a s t e r n  were found to be much lower  (0.3%). While b u t t and stem decay are accounted f o r  in  predictions,  unclear.  This  is  the p r o p o r t i o n due t o  important  require different well,  since  tomentosus i s  tomentosus as a r o o t  rot  management than decays and h e a r t  w h i l e decay and h e a r t  rot  will r o t s . As  l o s s e s may be e x p e c t e d  d e c r e a s e s h a r p l y w i t h s h o r t second growth r o t a t i o n s , may not be t r u e  d.  for  yield  to  this  tomentosus.  INCREMENT REDUCTION  As p r e v i o u s l y s t a t e d ,  K. tomentosus d i s r u p t s and k i l l s  root  bark and cambium and induces decay of the x y l e m . Defense r e a c t i o n s t o i n f e c t i o n and the p r o d u c t i o n of roots u t i l i z e  energy which might  i n c r e m e n t a l growth. As w e l l , i m p a i r water and n u t r i e n t  o t h e r w i s e be put  uptake of the t r e e .  T h u s , as more and more of the  a g r a d u a l r e d u c t i o n (over (Whitney  into  a l o s s of c o n d u c t i v e t i s s u e can  would l o g i c a l l y be expected to r e s u l t rate.  adventitious  15-20  i n a d e c r e a s e d growth  root  years)  Such s t r e s s  system i s  attacked,  i n t r e e growth o c c u r s  1962b).  Merler  (1984), r e p o r t e d a r e d u c t i o n of 20% i n  the  27 latest  five  year  increment of d i s e a s e d white spruce t r e e s  aged 30 to 65 y e a r s . On a per s t a n d b a s i s , Whitney r e p o r t e d annual increment  r e d u c t i o n s of 0.11%  spruce stands sampled and 0.52%  i n white  i n b l a c k spruce s t a n d s .  While these annual r e d u c t i o n s appear  insignificant,  l o s s e s are c u m u l a t i v e and thus r e p r e s e n t over a  (1978),  such  substantial  losses  rotation.  The type of m o r t a l i t y which o c c u r s w i l l , depend on the  inoculum p o t e n t i a l  growing c o n d i t i o n s f o r the  t o some e x t e n t  of the f u n g u s . Where  fungus a r e optimum, more  rapid  death may be expected and thus more s t a n d i n g m o r t a l i t y , a s h o r t e r p e r i o d of The age of the affect  it's  incurred).  increment  reduction p r i o r  to  mortality.  i n f e c t e d t r e e c o u l d a l s o be expected t o  symptom development  (and thus the type of  loss  Younger, s m a l l e r t r e e s may be e x p e c t e d to show  above ground symptoms sooner due t o y o u n g e r , s m a l l e r systems. T h i s would l i k e l y  result  in swifter  of  increment  e.  INDIRECT LOSSES  root  m o r t a l i t y and  hence more s t a n d i n g m o r t a l i t y and a r e l a t i v e l y  short  period  reduction.  There are a l s o i n d i r e c t  s o u r c e s of damage a s s o c i a t e d w i t h  tomentosus. Dendroctonus r u f i p e n n i s K i r b y beetle),  and  uses w i n d f a l l  trees  (spruce bark  t o b u i l d up i t s  populations  I.  28 b e f o r e l i v i n g t r e e s are a t t a c k e d  (Dyer and T a y l o r  Such t r e e s are s u p p l i e d i n abundance by many a r e a s of B . C . As w e l l , facilitated i n the  tomentosus s t r e s s e d t r e e s  Thus  tomentosus can be seen as  an a c c o m p l i c e i n spruce bark b e e t l e  A less distinct  tomentosus i n  spread to l i v i n g t r e e s may be  by the presence of  immediate v i c i n i t y .  1971).  attack.  r e l a t i o n s h i p e x i s t s between  I.  tomentosus and H y l o b i u s warreni  (Warren's  S t r o n g c o r r e l a t i o n s between the  i n c i d e n c e s of these  p e s t s have been r e p o r t e d However,  the nature  (Krebill  1962,  collar  Whitney  attack  1961).  weevils  i n t o the r o o t c o l l a r c a u s i n g bark m o r t a l i t y  eventually  girdle  two  of the r e l a t i o n s h i p between the two has  not been c l e a r l y d e m o n s t r a t e d . W a r r e n ' s c o l l a r tunnel  weevil).  (and thus k i l l )  the t r e e .  and can  Serious weevil  undoubtedly weakens the t r e e perhaps d e p r i v i n g i t  any r e s i s t a n c e mechanisms to Alternatively,  i n f e c t i o n by  of  tomentosus.  advanced i n f e c t i o n by 1^ tomentosus weakens  the host t r e e and can cause r e s i n o s i s . E x u d a t i o n s from such t r e e s may" a t t r a c t W a r r e n ' s c o l l a r w e e v i l s . Once a t t r a c t e d , the weakened s t a t e  of the t r e e may i n h i b i t  thus c l e a r the way f o r  insect  attack.  p i t c h i n g out and  29  6. VARIABILITY OF INONOTUS TOMENTOSUS  Inonotus highly or  tomentosus  variable  fungus.  1944, Haddow  "cultural  experiments  tomentosus several  cultural  on  these  recognized  Haddow  (1941) each  conspicuously  c h a r a c t e r i s t i c s " . However,  states: form (P. comprises  among  further  i s lacking  a n d no p u b l i c a t i o n s  were c i t e d .  Haddow may  in fact  varieties Whitney  o f P_j_ t o m e n t o s u s  and Bohaychuk  tomentosus conifer  in cultural  seedlings.  chlamydospore sharing  and M a i r e  This  in  pathogenicity  accepted  several recently,  t h e v a r i a t i o n of I .  may  resulted  pathogenicity  v a r i a t i o n among  silvicultural  that  studied  analysis  signify  o r b o t h of the r e c o g n i z e d great  i n which  c h a r a c t e r i s t i c s and p a t h o g e n i c i t y  Cluster  variability  of  (1922),  of these  referring to  were p r o p o s e d . More  (1977),  production,  significant  have been  themselves  elaboration  studies  work done by S a r t o r y  of s t r a i n s  i t sdiversity  var. circinatus)  differ  as a  a number  to explain  shown t h a t  a n d P_j_ t o m e n t o s u s  in  data  1941). have  s t r a i n s which  been  So much s o t h a t  v a r i e t i e s h a v e been p r o p o s e d  (Gosselin  one  has long  e x i s t s between  and growth  color, rate  isolates.  a number  species.  importance  i n mat  on  Such  of s t r a i n s  within  s t r a i n s would  i f significant  differences  them. I t i s g e n e r a l l y  c i r c i n a t u s i s less pathenogenic.  be  I f so,  30  c o n t r o l of l o s s e s i n second growth s t a n d s c o u l d be a c h i e v e d by i d e n t i f y i n g t h e s p e c i e s o r s t r a i n p r e s e n t and p l a n t i n g s p e c i e s o u t s i d e of t h e fungus' hypothesized  that  h o s t range. M e r l e r  tomentosus may comprise  (1984)  a number of  s t r a i n s , each s p e c i f i c t o one o r a v e r y few s p e c i e s w i t h i n the o v e r a l l h o s t range. T h i s p o s t u l a t i o n was prompted by t h e absence of fir  tomentosus i n f e c t i o n on p i n e s and s u b a l p i n e  ( A b i e s l a s i o c a r p a (Hook.) N u t t . ) , on h i s s t u d y  sites  d e s p i t e t h e s e s p e c i e s b e i n g r e p o r t e d as h o s t s e l s e w h e r e i n Canada.  S t r a i n s of H e t e r o b a s i d i o n annosum have been documented. More i m p o r t a n t l y , t h e s e s t r a i n s have s i g n i f i c a n t l y  different  p a t h o g e n i c i t y and h o s t s p e c i e s ranges (Korhonen 1978). As w e l l , A r m i l l a r i a m e l l e a s e n s u l a t o has r e c e n t l y been shown t o be a number of d i f f e r e n t s p e c i e s , each w i t h d i f f e r e n t h o s t r a n g e s and p a t h o g e n i c i t i e s ( M o r r i s o n e t a l 1985). Thus, i t can be seen t h a t t h e p r e c e d e n t  has a l r e a d y been s e t and  t h a t t h e v a r i a b i l i t y of IN tomentosus may be e x p l a i n e d by t h e e x i s t e n c e of s t r a i n s o r v a r i e t i e s . V a r i a t i o n i n tomentosus r o o t r o t b e h a v i o u r with the v a r i e t y c i r c i n a t u s  has been o f f i c i a l l y  recognized  (now a s e p a r a t e s p e c i e s ) . A s i d e  from t h e d i f f e r e n c e i n s e t a e shape, 1^ c i r c i n a t u s has a l s o been shown t o be somewhat l e s s p a t h o g e n i c f a s t e r growing  i n c u l t u r e (Whitney  on s e e d l i n g s and  and Bohaychuk  1977).  31 B.  METHODS  1.  S E L E C T I O N OF  S I T E S AND  PLOT LOCATION  T w e n t y p l o t s were s a m p l e d biogeoclimatic  zone  (Figures  3,  from  2 and  of  L a k e s and  Bulkley  Region.  the  status,  Since  stands  root  and  challenged  trees  challenged  tree  meters  of  of was an  above ground  root  symptoms  examinations. a l l trees  50  and  not  to being  In  as  Rupert  100  show  a tree  contact was  or  and  West  the  Morice,  Forest pine were  infection presence  selected  for  sampling.  least  were r e q u i r e d .  found  p l o t had  a  A  within (upon  infected root.  t e n t a t i v e l y estimated  located within  I.  ten  which o c c u r r e d  an  of  t o e n c o m p a s s an  w h i c h was  with  selected  years  f o r the  pine  Columbia  lodgepole  (where e x p r e s s e d ) and  the  and  each p l o t , at  s p r u c e and  center  P l o t s were within  Prince  maps do  infected tree  infection  the  Region  were p o s i t i o n e d  defined  size  George E a s t  were s u r v e y e d  both  t o have  of  the  between  center.  excavation)  s t a n d s were  i n f e c t e d mixed  rot prior  were c i r c u l a r  Prince  of  Spruce  central British  George F o r e s t  stands  infection  three  the  f o r e s t cover  candidate  north  Sub-boreal  Candidate  Districts  tomentosus  that  5).  Prince  spruce  tomentosus  Plots  of  Inonotus tomentosus  interior  desired.  I.  Table  f o r e s t c o v e r maps o f  Districts  and  (SBS),  i n the  cursory  centers  reasonably  to  The using  root ensure  high  chance  F i g u r e 2. L o c a t i o n Forest Region.  of Study  Sites  l n the P r i n c e  George  Scale  = 1:300,000  F i g u r e 3 . L o c a t i o n of s t u d y s i t e s r e s t Region  In the P r i n c e Rupert  Table 5 Stand Age, Region and Biogeoclimatic of Sample Sites Site no.  Stand age  1  105  2  Region  Classification  Subzone or Variant  Association *  PG **  SBSK3  04.1  88  PG  SBSK3  04.1  3  103  PG  SBSK3  01  4  75  PG  SBSK3  04.1  5  50  PG  SBSe2  01  6  54  PG  SBSe2  01  7  110  PG  SBSe2  04  8  104  PG  SBSe2  01  9  45  PG  SBSe2  04  10  52  PG  SBSe2  01  11  50  PG  SBSe2  01  12  86  PR  SBSel  04  13  87  PR  SBSel  01  14  90  PR  SBSel  04  15  80  PR  SBSel  01  16  73  PR  SBSd  08  17  74  PR  SBSel  04  18  54  PR  SBSd  07  19  54  PR  SBSd  07  20  87  PR  SBSel  01  * described  i n Appendix B  ** PG = Prince George  PR = Prince Rupert  35 of c o n t a c t i n g inoculum (and thus becoming i n f e c t e d ) . s i z e was 0.049 ha (12.5 pine),  2.  whichever was  m radius)  or 50 t r e e s  Plot  (spruce and  less.  DATA COLLECTION  Each main canopy t r e e i n each p l o t number. A c e n t r a l  tree,  approximating  was marked w i t h a  the c e n t e r  i n f e c t e d area was d e s i g n a t e d . The c e n t e r be the  the  t r e e was taken  t r e e which showed most advanced symptoms of  tomentosus root point  of  rot.  This central  from which the other  trees  I.  t r e e s e r v e d as a i n the p l o t  to  reference  were mapped.  T h i s was a c c o m p l i s h e d u s i n g d i s t a n c e s and compass r e a d i n g s from the c e n t r a l  tree.  On each t r e e ,  sampled w i t h i n 30 cm of the b o l e . a 3/8  i n c h diameter  attachment  t h r e e major  Sampling i n v o l v e d  h o l e i n each root  f o r a chainsaw.  r o o t s were  Inonotus  drilling  u s i n g an Atom D r i l l  tomentosus i n f e c t i o n was  i n d i c a t e d by red s t a i n e d wood c h i p s or white p i t t e d ,  rotted  wood. Root e x c a v a t i o n s were performed and samples taken c u l t u r i n g where any doubt r e g a r d i n g the  infection  a t r e e e x i s t e d . T r e e s d e s i g n a t e d as h e a l t h y sampling were not e x c a v a t e d .  upon  status drill  A number of samples were  c o l l e c t e d from s t a i n e d and r o t t e d  r o o t s i n each p l o t  i s o l a t i o n and study of c h a r a c t e r i s t i c s i n  culture.  for  for of  36 Each p l o t within. center  Trees were  map  was  examined  suspected  of  being  t h r e e m e t e r s away  trees  w h i c h had  center  than  checked source a  when t h e  plot.  For  t r e e was  found, the  infection  adjacent,  trees  system  were r e c o r d e d .  A  were r e c o r d e d  an  f u r t h e r away  re-entered. the  was  from  the  ecological  and  of  the  depth  and  i n the  field.  were c o l l e c t e d  for determination  o f pH.  Measurements of  height  and  age  of  clinometer  and  increment  were s e l e c t e d a t increment  3.  infected of  each  random a n d  In  10  plots,  Soil  using  a  up  20  to  d i a m e t e r measurements from  samples the  trees  and  them.  SUSCEPTIBILITY  s p e c i e s w i t h i n each p l o t ,  t r e e s was  challenged  c a n o p y were t a k e n  borer.  b o r e s were t a k e n  HOST S P E C I E S  For  main  with  to  features  excavated  the  contact  horizon  the  by  from were  classified  biogeoclimatic  p i t was  were  infection  removed  ( K r a j i n a 1965). S i t e soil  and  Where no  t r e e was  the  which  infected  I ^ tomentosus  vegetation  a s s o c i a t i o n using  classification  texture  with  p l o t s were  each p l o t  ecolgical  area  contacts  center  t r e e s were t h o s e  f r o m any  infected  the  t h e m s e l v e s . S u c h t r e e s were n o t e d  for root  diseased  no  r e d e f i n e the  outside  re-examined. Suspect  more t h a t and  to  trees  totalled  and  ( h e a l t h y and  the  d i v i d e d by infected),  number the of  of  total that  number species,  37 w i t h i n the p l o t . rate for  the  This proportion  species in that p l o t .  performed on s e l e c t e d t r e e s c o n t a c t s or g r a f t s  infection  whether  root  study the growth and  the d i s e a s e . O b s e r v a t i o n s of  and induced symptomology of  and any v a r i a b i l i t y  the  Root e x c a v a t i o n s were  to determine  e x i s t e d and to  spread b e h a v i o u r of etiology  represented  the  t h i s pathogen were made  i n behaviour w i t h t r e e  s p e c i e s or  g e o g r a p h i c a l l o c a t i o n was n o t e d .  A t-test determine  if  and an a n a l y s i s of v a r i a n c e were performed g e o g r a p h i c a l l o c a t i o n or e c o l o g i c a l  c l a s s i f i c a t i o n affected rates  the  differences  of p i n e and spruce on each s i t e .  (within centers) paired  were compared f o r  sample t - t e s t  w i t h i n each p l o t  infection  were not  equal  if  infection  rates  the  a non-parametric  statistical  1976)  it  test  by s p e c i e s and by  was performed  within centers  (Fox and G u i r e  As  two s p e c i e s  to  (sites)  f o r p i n e and s p r u c e . These a n a l y s e s were  u s i n g the MIDAS  rates  two s p e c i e s u s i n g a  rates  test  infection  Infection  of  independent,  A W i l c o x i n matched p a i r s  between  being one p a i r ) .  rates  was performed to compare i n f e c t i o n  determine  the  ( w i t h each p l o t  was b e l i e v e d t h a t the  site.  to  performed  and SPSS.X  packages on the UBC MTS mainframe  were  (Lai  1983)  computer.  38 4. E F F E C T OF  INFECTION ON  INCREMENT  One h u n d r e d a n d e l e v e n taken pine  at breast trees  height  1/4  from  i n the P r i n c e  healthy  Where o b s t r u c t i o n s  close  t o the north  a plastic diameter tree  until  increment five  which  were u s e d  been  height,  the increment  could  The  basal from  variance  area  status  of each  and t h e l a s t  five  was t h e maximum  decay  occurred  s a m p l e was d i s c a r d e d  the r a d i a l  was p e r f o r m e d  t r e e diameter  covariates. variable.  be made. T h e  of the tree c o u l d  Where b o l e  stored in  an Addo-X m e a s u r e r .  as t h i s  status  increment  subzone and i n f e c t i o n and  using  was  since  Only period  be a s s u m e d at breast the tree's  n o t be d e t e r m i n e d .  calculated of  constant.  Forest  s i d e of the  bore  and t h e i n f e c t i o n  was m e a s u r e d  t o have  age  measurements c o u l d  the i n f e c t i o n  and  e x i s t e d , t h e s a m p l e was t a k e n a s  height  years  were  spruce  Rupert  the north  s i d e as p o s s i b l e . Each  at breast  last  over  from  bores  and d i s e a s e d  were n o t e d . The a g e was d e t e r m i n e d  years the  straw  increment  George and P r i n c e  R e g i o n s . A l l b o r e s were t a k e n tree.  inch  Basal  The a n a l y s e s  the l a s t  five  years  increment. A two-factor f o r each  species.  was  analysis  Biogeoclimatic  s t a t u s were u s e d a s f a c t o r s . T r e e age  were t e s t e d area  over  for possible  increment  inclusion  as  was t h e i n d e p e n d a n t  of v a r i a n c e  were p e r f o r m e d  u s i n g the  39 GENLIN s t a t i s t i c a l  package ( G r e i g and B j e r r i n g  1980)  on the  UBC MTS mainframe.  5.  CULTURAL IDENTIFICATION AND DESCRIPTION  R o t t e d a n d / o r s t a i n e d wood was c o l l e c t e d from each host s p e c i e s on each s i t e culture.  for  i s o l a t i o n and i d e n t i f i c a t i o n  Samples from each s i t e were s u b s e q u e n t l y s t o r e d at  1 degree C e l s i u s f o r  3 months. The samples were then removed  from c o l d s t o r a g e and c u l t u r e s of agar  I_j_ tomentosus on 3% malt  i n d i s p o s a b l e 100 X 15 mm p e t r i  from e a c h . I d e n t i f i c a t i o n s Nobles I d e n t i f i c a t i o n  of  p l a t e s were  room temperature  at  of C u l t u r e s of Wood R o t t i n g Fungi  which time  ten days at  d i s h w i t h an  The growth margin was a g a i n d e l i m i t e d at the two r a d i a l  t o c a l c u l a t e the growth r a t e isolate.  for  the growth margin was  d e l i m i t e d on the bottom of the p e t r i  d a y s . The average of  obtained  I_j_ tomentosus were made u s i n g  (1948). Each c u l t u r e was s t o r e d i n the dark  marker.  in  (in  indelible 20 and 30  increments was then used  mm per day)  Mat c o l o r and changes t h e r e i n  f o r each  were r e c o r d e d  c o n c u r r e n t l y with growth margin d e l i n e a t i o n . Mat c o l o r and growth c h a r a c t e r i s t i c s of each i s o l a t e were then compared t o those of other  isolates.  40  C.  RESULTS AND  1.  ETIOLOGY  DISCUSSION  Differences of  were n o t e d  tomentosus between  spruce  studied  detectable roots  as  a  the  presence  light  4).  (Figure  tissue  the  i n the  red  This  of  was  brown c o l o r clearly  at  i n which white  evident  present  (Figure  the  system Such  conductive  t i s s u e was  alive.  structural  support  the  provide maintain  the  tree with  an  root  collar.  and  i n the  root  cambium and occurred  resulted  was  creamy  white  origin  yellowish r o t had  even roots  to a  large  the  provided  appeared,  were o f t e n  mycelium  s i z e s or and  rot  a  with was  little  however,  where  to  to severe  abundant  progressed  phloem  was  outer  c a u s e d d i s r u p t i o n of of  rusty  pits  nutrients  cases  the  of  pitting  though  In  of  xylem  to advanced  water and  in necrosis various  the  inner  progressed  crown.  some c a s e s ,  of  center  near  as  roots  cambium. T h i s  in patches  mycelium  healthy  adventitious In  the  t r e e . They  sufficient  apparently  occurred,  the  to  or  early  root  the  first  then  Often  On  was  Advanced  6).  (Figure  throughout  5).  appearance  disease  stain  and  and  studied.  s t a i n i n g i n the  a p p r o a c h e d . Such  white mycelium  The  the  stain  red  hosts  became p r o g r e s s i v e l y d a r k e r  infection  decay  two  behaviour  at  along the  t i s s u e . Mycelium continuous  flake-like.  In  mat.  places  41  Figure Light  red s t a i n  in infected  4  pine  with  l_.  tomentosus.  42  43  44 mycelium  adhered  t o or  soil  the  clumps  of  to  soil  the  the  later  between the  stages bark  wood. The  xylem.  or In  present had  tended  the  precede  on  grew root  or  s u r f a c e . T h i s h a b i t of  adhering  some c a s e s ,  of mycelium  any  staining  extensive  appeared  t o be  (although  less)  cambial  diameter  no  related  small roots  had  trend. Exceptions cm)  between  on The  and  the  of  the  was  larger  living  bark  roots and  of m y c e l i a l  12  cm  It that  diameter  internal  root  and  experienced  cambium. T h i s was  were p r e s e n t  was  many  recorded)  (approximately  inner  pitting  root  type  concurrent  from  necessarily  to root diameter. data  In  sheets  separated  rotting  staining  quantitative  a t t a c k o n l y and  (12  and  staining.  sometimes e x p e r i e n c e d  observed  formed  d i d not  i n and  fungal progression while  internal  the mycelium.  s u r f a c e mycelium. Conversely,  development  and  of  s h i c h were e a s i l y  xylem  f o l l o w severe  sized  presence  and  root  medium  soil  the mycelium  apparent  appeared  the  of d e c a y ,  no  an  hardened  e x t e n s i v e m y c e l i a l growth  light  and  causing  t o mask t h e  presence  without  into  and  classes i s admittedly  the  only  cutoff  very  approximate.  Merler diameter unable  and  (1984) a l s o  correlations  disease behaviour.  to penetrate  progression  found  from  the  thick  a column  The  bark  of decay  between  f u n g u s was of  root  found  to  be  large roots. Instead  i n the  stump t o a  healthy  45 root of  occurred  i n the  approximately  the  b a r k and  observed  study,  five  trends,  suggest  progresses  cm  grow a l o n g  proximally  behavioural  x y l e m and  that  as  the the  along  with  longer  able  t o c o l o n i z e the  roots.  The  of  after  the  center  differently ground  in pine.  symptomology  x y l e m was  again  infection  outer  or  of  only  with  stopping was  near  the  differed. as  resin  soaking  of  the  commonly  resin  d i d not  progression isolated  of  and  disease  is  no  diameter of  behaviour  a  toward  large stages  t o be  fungus. resin  of  the  origin  the  often  tissues. This  Such copious  effective Inonotus  iself  above  the  before  T h i s was  root  and  staining  occurred  the  from w i t h i n  roots  manifested  quite b r i t t l e .  the  this  i s i n advanced  a light  killed.  appear  These  diameter  surface  progressed  of  in  large  disease  a medium s t a i n i n g cambium were  butt.  rotted).  While  one  was  external  the  the  the  root  p l o t s the  r o o t s were c h a r a c t e r i s t i c a l l y production  of  order  cambium)  collar,  Both below ground  present,  sapwood and  associated  study  smaller  diameter  penetrate  the  discovered  cambium  a  reverse  root diameter  c o l o n i z a t i o n ( d a r k l y s t a i n e d or  the  to  xylem and  root  f u n g u s a p p e a r s on  I n many of  of  the  those through  Beyond a c e r t a i n ceases.  only  able  fungus approached  progression  root  at  s u r f a c e . The  ( i n the  Near  until  f u n g u s was  infection  internally  surficially.  the  cambium  in  tomentosus  s o a k e d wood  and  46 progression  o f t h e f u n g u s was a p p a r e n t . L i t t l e  was o b s e r v e d decay  i n these  occurred  diameter), during  i n small  lateral  w h i c h were l e f t  was n o t e d  what a p p e a r e d and  r o o t s . I t i s p o s s i b l e , however,  the excavation.  mycelium  o r no d e c a y  roots  i n the s o i l  Although  a t the time  little  after  1 cm  than  or otherwise  obscured  o r no s i g n o f  of excavation,  t o be r u d i m e n t a r y  i n many o f t h e r o o t s  (less  that  fruiting  m y c e l i u m and  bodies  a p p e a r e d on at 1  two months s t o r a g e  degree C e l s i u s .  This ground  form of a t t a c k  symptoms  pronounced Trees  presumably roots  in this  utilization  mortality  usually died that  (Figure 7 ) .  standing,  decay of t h e main  A f t e r death  showed  of t h e h o s t ,  t i s s u e s by s a p r o p h y t e s a p p e a r e d  supportive d r y i n g and to  prevent  pitted  r o t and p r o d u c t i o n  of f r u i t i n g  bodies  not occur.  In  and  trees. Infected pines  above  o f t h e t i s s u e b y I_;_ t o m e n t o s u s . T h u s t h e  characteristic  of  manner  due t o t h e f a c t  of tree  r e s u l t e d i n obvious  t h i n n i n g and e v e n t u a l  had n o t o c c u r r e d .  invasion  did  in diseased  crown  killed  i n pine  three  the Prince below  Living  sites Rupert  i n the Prince sites,  ground) s i m i l a r  trees  pine  George  displayed  t o those  showed e x t e n s i v e  region  root  symptoms  described and b u t t  a n d i n many (above  f o r spruce. r o t and green  47  Figure Above  ground  symptoms  7  i n an  infected  pine.  48 windfall stands  o c c u r r e d . Such b e h a v i o u r  i n the Cariboo  symptoms o c c u r r e d , cambial  The c a u s e  the disease  physiological spruce  disease  may  from  fails  likely  inoculum  large  roots,  likely  until  source.  between p i n e and  strains  of the  at a l l .  in infected  p i n e s on  t o the succeeding  F a s t e r growing,  of the  one. I f  the tree i s k i l l e d ,  i t will  saprophytic  decay  c o l o n i z e t h e r o o t s a n d stump. W i t h o u t t h e  reserves provided  by d e c a y i n g  n o t be a b l e  stumps a n d  t o s u r v i v e on t h e  i s o l d enough t o s u p p l y  i s the case,  then  pine  i n pines  would c l e a r l y  was s i m i l a r represent  i n the next  to that  food  may i n  disease  i n spruce,  p o s s i b l e inoculum  rotation.  a new  infections  be n o n - t r a n s m i s s i b l e . However, w h e r e  progression  infection  before  regeneration  some c a s e s  five  from  on t h e t r a n s m i s s i b i l i t y  t o m e n t o s u s may  If this  arise  differences  to develop  h a v e an e f f e c t  not develop  large  trees  of decay  to develop  will  than  i n t h e development  i s u n c l e a r . I t may  the c u r r e n t stand  fungi  less  present.  failure  some s i t e s  spruce-type  t o pronounced  in roots  variation  or morphological  being  The  site  soaking  prone  o r i t c o u l d be due t o d i f f e r e n t  pathogen  decay  still  of t h i s  i n pine  observed i n  G e n e r a l l y , where  p i n e s were  n e c r o s i s and r e s i n  cm d i a m e t e r . of  Region.  was a l s o  these  reserves f o r  49 Observations sample p l o t s )  of  infected  pines  (both i n and o u t s i d e  suggested t h a t the d i s t i n c t i o n  symptomology types may be age r e l a t e d .  symptoms. Whereas younger p i n e s d i d n o t . that  between  Older pine trees  stands seemed to show the more c l a s s i c ,  of  and  spruce-type It  is also possible  symptom e x p r e s s i o n i s r e l a t e d to t r e e v i g o u r ,  fungal  inoculum p o t e n t i a l or even t o r a c e s or v a r i e t i e s  of  disease.  implicated  from t h i s pines  None of study.  in Ontario  these  f a c t o r s c o u l d be c l e a r l y  However, are  Whitney  generally  even when i n c l o s e q u a r t e r s tomentosus. least I.  While  this  study  (1985), r e p o r t e d  infected  with  I.  of  the  at  species,  c o u l d not be made. Thus i t  the s p e c i e s of  c a u s i n g the v a r i a b i l i t y  with  found p i n e s to be i n f e c t e d  circinatus  q u i t e p o s s i b l e that  that  circinatus,  w i t h spruce i n f e c t e d  as o f t e n as s p r u c e , the d e t e r m i n a t i o n  tomentosus or  the  Inonotus p r e s e n t  is  is  i n symptom e x p r e s s i o n .  The absence of advanced decay i n p i n e a l s o has implications  in surveys for  the d i s e a s e i n the  field  the  relies  and the presence of p i t t e d r o t surveys for  This situation reported  heavily and dark  Identification on green  symptoms are  may be p a r t i a l l y  i n c i d e n c e s of  not  reliably  responsible  Thus, inaccurate  produced.  for  the d i s e a s e on p i n e  of  windfalls  red s t a i n .  the d i s e a s e i n pine c o u l d be g r o s s l y  in a r e a s where these  of  disease.  in  the  rarity  British  50 Columbia.  Although be  infected  generally the  u n d e r s t o r y p i n e and spruce  i n some o f t h e s t u d y  fairly  plots,  o l d and s u p p r e s s e d .  symptomology e x p r e s s d  by t h e s e  be e x t r a p o l a t e d t o y o u n g p l a n t a t i o n  t r e e s were these  trees  plantation  trees  can not  trees.  needle  l e n g t h , reduced  rounding,  thinning  Infection appeared warreni  and animal  appressed  noted,  root  no  evidence.  often  found  death.  with  what  Wounds due t o H y l o b i u s  t r a m p l i n g were a l s o  observed.  were a s s o c i a t e d  i n c i d e n c e s of i n f e c t i o n one p r e v i o u s l y  no s p o r e  cases without  t o crown  in association  wounds.  infection  v i a root  tight  source  often  with  The  tightly  o c c u r r e d as a  originating  infected  root  c o n t a c t s appeared  present  roots.  at a  root  present  were  i n t h e s e a r e a s was i n  appression, especially was  spruce  reduced  c o n t a c t between h e a l t h y and i n f e c t e d  Spread  inoculation  b l u e i n g of needles,  contacts. Infection  without thus  of spruce.  o f t h e crown and e v e n t u a l l y  wounds n o t e d  of t h i s  However, contact  was  necessarily  Infected  l e a d e r growth l e a d i n g  t o be c o m p r e s s i o n  compression  result  showed  were  O b s e r v a t i o n s of  p i n e s were n o t o b s e r v e d .  15 y e a r s o l d o r l e s s  to  T h u s o b s e r v a t i o n s on  t h e d i s e a s e w e r e , h o w e v e r , made i n p l a n t a t i o n s Infected  found  (advanced  to occur where a  i n some  large  d e c a y ) . T h i s would  51 seem t o - i n d i c a t e t h a t but  not  necessary  Hylobius study  plots.  tunnels cortex  Feeding  a t and  may  x y l e m . Red weevil  and  by  from  the  was  this  themselves quickly  would or  observed  spores  or m y c e l i a  deterrent  which  of  i n two  by  to  is likely  by  1^  be  washed  i n on  the  t o have  short  or  to  time p e r i o d d u r i n g has  a  be  four  (1969)  filled,  life  weevils  tunnels a  short  effect  spores  year  the  i n . If  itself,  deterred  resin  which  to  samples  blown  wounds. T h u s , a l t h o u g h  found  i n . H_j_ w a r r e n i  such  little  not  least  on  insect  was  inoculated root  the  i s only  tomentosus  were t y p i c a l l y  carried  there  Denyer  tunnels a  carried  and  of  the  contaminated.  Whitney  soaking  Weevil  I.  instances,  w a t e r . As  resin,  carried  infection.  infection  these  could  the  into  to y i e l d  subsequently  with  of  t r e e s . Attempts  failed  of  the  i n c l o s e a s s o c i a t i o n with  wind and  spores  are  resin  to,  resinous  Necrosis  infection  in three  s t a i n e d wood  carried  during  be  avenues of xylem  i n many o f  i n some c a s e s .  involve basidiospores  period  conducive  resulted in  collar.  extensive  i n the  become p l u g g e d  production  weevil  root  s e v e r e l y d i s t u r b e d and  Infection  that  this  have p r o v i d e d  t o m e n t o s u s . However, were  wounds were common  was  staining  wounds a r e  infection.  around  tunnels  isolate  for  warreni  occurred  tunnels  compression  the  as  a  reported by H_j_  resin warreni  there  could cycle  i s at  wash with  or  52  s t a g e s of development  which are not t i e d  to s p e c i f i c  s e a s o n s . Thus at any g i v e n time i n the y e a r ,  all  of  s t a g e s of development are l i k e l y  to be p r e s e n t .  is therefore  d u r i n g the p e r i o d of  likely  to be a c t i v e  (1963,  1966)  The  the weevil spore  release.  Whitney  reported staining  of white spruce i n o c u l a t e d w i t h Such s t a i n i n g d i d not occur but not  tomentosus b a s i d i o s p o r e s .  i n r o o t s which had been wounded  i n o c u l a t e d . C h r i s t e n s e n (1940) r e p o r t e d t h a t  tomentosus hyphae were s p a r s e i n l i g h t l y Since  I.  d i s c o l o r e d wood.  tomentosus c o u l d not be i s o l a t e d from many of  r e d s t a i n e d specimens i t have been p a r t i a l l y  i s p o s s i b l e t h a t the  a r e a c t i o n by the t r e e t o  T r a m p l i n g was e s p e c i a l l y e v i d e n t one and two where c a t t l e the  i n root wounds  root c o l l a r  o b s e r v e d e x t e n d i n g to or  beneath the bark at  8).  s t a i n i n g may infection.  i n P r i n c e George s i t e s  g r a z i n g had o c c u r r e d . Wounding at  was e v i d e n t .  r o o t wound ( F i g u r e  S t a i n i n g on one sample was  from what appeared t o be an o l d  Inonotus  tomentosus was i s o l a t e d  the o l d wound. As the e x t e n t of avenue of  i n f e c t i o n was  apparent,  implicate  this  t h i s would s t r o n g l y infection  in t h i s case.  inoculum was in e v i d e n c e ,  it  from  staining  was v e r y l i m i t e d and no other  the avenue of  the  root wound as  S i n c e no source of  appeared t h a t  i n o c u l a t i o n must  53  Figure Red  stain extending  8.  from an  o l d root  wound.  54 have o c c u r r e d  by a means o t h e r  than  root  contact.  2. SPREAD  Spread occurred  by r o o t  where r o o t s  appressed. required.  Root  contact  spread  the  between  roots  occurred. staining healthy  or r o t t i n g ,  through  3.  HOST S P E C I E S  Figures the  Prince  roots  loosely was  i n t e r - and  Roots of  various  was a l s o o b s e r v e d . stages  of  which  grafts involved  one r o o t  t r a n s f e r of inoculum There  Such  infection.  i n many  cases,  prevented  t i s s u e through which  regularly occurred.  spread  This  involved.  t o an e x t e n t  of the root  of  mycelium  x y l e m was a p p a r e n t . However,  Where r o o t  root  was o b s e r v e d  and s p r u c e .  were  i n evidence.  o r i n some c a s e s ,  i n e a r l y and l a t e  were d e c a y e d  determination  frequently  ectotrophic  spread  between g r a f t e d  was o b s e r v e d  Transfer  case,  i n pine  (2 cm. t o 12 cm.)  Spread  was  were t i g h t l y ,  In the l a t t e r  intra-specifically sizes  contact  spread with  had  dark  to the previously  were no i n d i c a t i o n s  the s o i l . *  SUSCEPTIBILITY  2 a n d 3 show t h e l o c a t i o n s o f s a m p l e p l o t s i n  George and P r i n c e  Rupert  forest  regions  r e s p e c t i v e l y . Table  5 lists  the age,  biogeoclimatic c l a s s i f i c a t i o n r e f e r s not  region  and  of a l l s i t e s sampled. Region  to the B r i t i s h Columbia M i n i s t r y of F o r e s t s  and  Lands a d m i n i s t r a t i v e r e g i o n but t o the g e n e r a l l o c a t i o n of the p l o t i n two  w i t h i n B r i t i s h Columbia. The  widely  s e p a r a t e d c l u s t e r s . One  around P r i n c e George and Houston. The  stands  f o l l o w i n g Sub-Boreal  located  l o c a t e d around  from 45 t o 110  years.  Spruce subzones were r e p r e s e n t e d ; the S u b a l p i n e  The the  f i r Subzone, Western  ( S B S e l ) , the F r a s e r B a s i n M o i s t C o o l C e n t r a l  Sub-Boreal Dry  c l u s t e r was  the o t h e r was  ranged i n age  Spruce Subzone (SBSd), Variant  p l o t s were d i s t r i b u t e d  Spruce Subzone (SBSe2) and  Southern  Sub-Boreal  c o n t a i n s a map  Challenge  the Nechako R i v e r Warm  Spruce Subzone (SBSk3). Appendix A  of the c h a l l e n g e t r e e s i n each sample  t r e e s were those  trees ( l i v i n g ,  plot.  dead,  i n f e c t e d or h e a l t h y ) which o c c u r r e d w i t h i n t h r e e meters of an  infected  t r e e or which were found  have r o o t c o n t a c t w i t h an Table  6,  infected  infected  (PPIN) was  i n a l l cases  interior  spruce  p r o p o r t i o n of l o d g e p o l e p i n e  ranged from 0.125  g r e a t e r than  infected  (PSIN).  pine  the p r o p o r t i o n The  i n f e c t e d ranged from 0.429 to  w i t h a mean of 0.830. The  infected  to  r o o t . As can be seen i n  the p r o p o r t i o n of c h a l l e n g e d l o d g e p o l e  of c h a l l e n g e d  1.00  (upon e x c a v a t i o n )  p r o p o r t i o n of i n t e r i o r  spruce  t o 0.897 with a mean of 0.563.  Table  6  P r o p o r t i o n s of Pine and Spruce I n f e c t e d and the D i f f e r e n c e i n P r o p o r t i o n of Pine and Spruce I n f e c t e d by S i t e  Site No. *1  H. *2  Spruce I. PSIN *3  1 2 3 4 5 6 7 8 9 10 11  10 8 3 4 14 12 13 15 7 9 10  12 10 20 12 2 7 7 7 4 10 6  0.55 0.56 0.26 0. 43 0.13 0.37 0.35 0.32 0.36 0.53 0.44  3 5 5 6 3 1 3 2 4 2 2  12 13 14 15 16 17 18 19 20  11 3 6 6 3 5 4 3 2  14 16 18 13 26 26 14 12 10  0.56 0.84 0.75 0.68 0.90 0.84 0. 78 0.60 0.83  3 2 0 0 1 1 1 0 3  H.  Pine I.  PPIN  DIFF  14 7 6 15 14 12 16 10 3 13 10  0.82 0.58 0.55 0.71 0.62 0.92 0.64 0.83 0.43 0.87 0.63  0.26 0.03 0. 29 0.29 0.70 0.55 0.49 0.52 0.07 0.34 0.39  14 19 19 11 11 9 20 23 23  0.82 0.91 1.00 1.00 0.92 0.90 0.95 1.00 0.69  0.26 0.06 0.25 0.32 0.02 0.06 0.18 0.20 0.05  *1 S i t e s 1-11 l o c a t e d l n the P r i n c e George F o r e s t Region S i t e s 12-20 l o c a t e d i n the P r i n c e Rupert F o r e s t Region *2 The  number of h e a l t h y  t r e e s i n the p l o t .  *3 The  number of i n f e c t e d t r e e s i n the p l o t .  PSIN= p r o p o r t i o n  of  PPIN= p r o p o r t i o n  of l o d g e p o l e  DIFF = PPIN - PSIN  i n t e r i o r spruce i n f e c t e d pine  infected  57 The d i f f e r e n c e  between  the p r o p o r t i o n of  i n f e c t e d and the p r o p o r t i o n of (PPIN -  interior  lodgepole spruce  infected  PSIN) was r e p r e s e n t e d by the d i f f e r e n c e  proportion infected  infected  ( D I F F ) . The d i f f e r e n c e  in  were a t t a c k e d  on a l l  A paired  proportion  i n d i c a t e d that both c o n i f e r sites  studied.  spruce i n f e c t e d ,  representing a p a i r ,  with each  showed t h a t the d i f f e r e n c e at  The W i l c o x o n p a i r e d sample t e s t  performed on the  level  (Table  8).  Correlation  Thus the  of  infection  higher  0.30.  (Table  9,  The s l o p e of Figure 9).  showed s i g n i f i c a n t  at  level  0.01  correlation one i n d i c a t e  (Table  and the that  10).  spruce  proportions  in  at  the  lodgepole  pine  infected  (P=0.012 ) w i t h an was p o s i t i v e  performed on  deviation  deviation in host  this  from a s l o p e of  The p o s i t i v e  the v a r i a t i o n  7).  interior  this correlation  significant  the  (Table  lodgepole  interior  A t-test  correlation the  levels  the p r o p o r t i o n of  yielded a significant correlation  (0.369)  level  than those of  i n f e c t e d w i t h the p r o p o r t i o n of  R-square of  0.01  between  i n f e c t e d was found to be s i g n i f i c a n t  p i n e were s i g n i f i c a n t l y spruce.  the  lodgepole  site  two p r o p o r t i o n s was s i g n i f i c a n t  each s p e c i e s  0.267. species  sample • t - t e s t on the p r o p o r t i o n s of  p i n e and i n t e r i o r  0.05  in  ranged from 0.020 to 0.699 w i t h a mean of  These r e s u l t s c l e a r l y  of  pine  s l o p e of  this  from a s l o p e of  species  one  Table Paired  7  58  T - T e s t on t h e P r o p o r t i o n o f P i n e I n f e c t e d and t h e P r o p o r t i o n o f S p r u c e I n f e c t e d  No. o f  variable  Cases  Proportion of s p r u c e i n f .  Mean  Standard Deviation  0.563  0.228  0.051  0.830  0.154  0.034  20 Proportion of p i n e i n f .  Difference Mean -0.267  2-tailed Prob. 0.012  standard Deviation  Standard Error  0.193  0.043  T-value  -6.18  Standard Error  Degrees o f Freedom 19  2-tail Prob. 0.000  Table wilcoxin Proportion  Mean Rank  cases  20  .00  - Ranks ( P r o p o r t i o n o f p i n e l e s s than P r o p o r t i o n of infected)  infected spruce  + Ranks ( P r o p o r t i o n o f p i n e greater than P r o p o r t i o n infected)  infected of pine  Ties  20 Z=3.920  59  Matched-Pairs signed-Ranked Test on t h e of Pine I n f e c t e d with P r o p o r t i o n of Spruce I n f e c t e d  00  10.50  8  (Proportion of pine i n f e c t e d equal t o the p r o p o r t i o n of spruce infected)  Total 2-tailed  P=0.0001  Table 9 60  C o r r e l a t i o n of the P r o p o r t i o n of Lodgepole pine on t h e P r o p o r t i o n o f S p r u c e I n f e c t e d .  Infected  N = 20 Source  Df  Sum o f Squares  Mean Square  F-Stat  Correlation Error Total  1 18 19  0.135 0.314 0.449  0.135 0.017  7.755  Mult.  R.  = 0.549  Variable  Constant Proportion of Spruce Infected  Partial  0.549  R-Square  0.622 0.369  0.080 0.133  0.012  SE  = 0.301  Coeff. Std. Error  Signif.  T-Stat  7.738 2.785  = 0.132  Signif.  0.000 0.012  61  Figure Correlation  of  the  proportion  of  9  proportion spruce  of  pine  infected  infected  on  the  Table T - T e s t of of P i n e  Ho: Ha:  10  t h e s l o p e of t h e C o r r e l a t i o n of the P r o p o r t i o n I n f e c t e d on t h e P r o p o r t i o n of S p r u c e I n f e c t e d  Regression Regression  has has  a slope equal to 1. a s l o p e not e q u a l t o  Slope of Regression  R e s i d u a l SS for Regression  0.369  0  .133  Residual Df  18  1.  T-Value  Prob.  -4.74  <0.001  63 susceptibility That  was p o s i t i v e ,  i s the i n f e c t i o n  same d i r e c t i o n  rates  of both  (hence p o s i t i v e ) ,  (hence d i f f e r e n t i a l ) . S i n c e a  continuum  from h i g h  quantitative. species  susceptibility was a l s o  would  increase  Qualitative  discrete  representing  than  variation of  varying  exists  one s p e c i e s  (and  vice  versa).  Positive  species  a r e t h e same  Positive  qualitative  centers centers.  infection  with  host  species  along  was i n host  would  have  species  differential i n host  infection  variation  species forms  levels, qualitative  s i t e the s u s c e p t i b i l i t y  of the other  qualitative  species  variation  i s low  exists  t h e s u s c e p t i b i l i t i e s o f t h e two r e l a t i v e l y high  variation rates  t o be l o w e r  l_y_ t o m e n t o s u s  within  i f one were  infection  I_j_ c i r c i n a t u s  i s considered  one w o u l d e x p e c t  or l o w ) .  w o u l d be e x p e c t e d  within  rates  As I_;_ c i r c i n a t u s  pathogenic,  that  (either  infection  varied  a continuum. Negative  when  site,  extents  susceptibility  relative  w h e r e , on a g i v e n  i s high  w h e r e , on a g i v e n  comparing  along  i n the  the s u s c e p t i b i l i t y  variation  o c c u r s where s p e c i e s  rather  case  as the other  susceptibility cases  rates  the c o r r e l a t i o n  decreased. Qualitative  possible.  varied  differential variation  1 0 ) . In t h i s  (Figure  quantitative.  but t o d i f f e r e n t  the i n f e c t i o n  susceptibility existed  one s p e c i e s  species  t o low, t h e v a r i a t i o n  If negative  been n e g a t i v e of  d i f f e r e n t i a l and  t o be  the i n f e c t i o n  in I^ circinatus  infection  less  rates centers  of both  1.0  64  Proportion Figure  10  of Spruce  1.0  Infected  Quantitative Differential Variation in Host S p e c i e s S u s c e p t i b i l i t y  1.0 »  x  x * * x X X X X  Negat i  P r o p o r t i on of P i n e Infected  ve  Posi t i ve  * *  %  X • #  Proportion Figure  11  of  Spruce  Infected  Qualitative Differential Variation in Host S p e c i e s S u s c e p t i b i l i t y  1.0  65 than  in  tomentosus c e n t e r s .  variation similar  would have r e s u l t e d  Differential in a c l u s t e r e d  to those shown i n F i g u r e  Table  11 summarizes PSIN,  - t - t e s t on the  difference  spruce i n f e c t e d  distribution  11.  PPIN and DIFF by r e g i o n . A  in- the p r o p o r t i o n s of p i n e  by r e g i o n  i n DIFF due t o r e g i o n  qualitative  indicated  (Table  significant  difference  12).  B i o g e o c l i m a t i c a s s o c i a t i o n was not  t e s t e d as a  as such a t e s t would be very unbalanced and would replicates  for  three a s s o c i a t i o n s . Instead,  were grouped t o g e t h e r the  b a s i s of  was f e l t factors  that  for  similarity  factor  lack  associations  the a n a l y s i s . They were grouped on  in moisture  and n u t r i e n t  regime.  such groups were more homogenous f o r  than were a s s o c i a t i o n s w i t h i n i n d i v i d u a l  Thus these  and  g r o u p i n g s were used i n s t e a d of  It  these  subzones.  subzones i n  the  analysis.  The t h r e e a s s o c i a t i o n groups d e r i v e d are Table  13.  in  Group 1 i n c l u d e d the mesic a s s o c i a t i o n s ; SBSe  Mesic Bunchberry - M o s s , SBSe2 01: SBSk3 01:  listed  Bunchberry - Moss and  P r i c k l y Rose - Queen's Cup. Group 2 i n c l u d e d  submesic a s s o c i a t i o n s ; SBSel 01: SBSe2 01: D o u g l a s - f i r  -  Subalpine  01:  the  Submesic Bunchberry - Moss, fir  and SBSk3 0 4 . 1 :  Wild  T a b l e 11 66 Summary o f t h e P r o p o r t i o n o f P i n e and S p r u c e I n f e c t e d and D i f f e r e n c e b e t w e e n P r o p o r t i o n s o f P i n e and S p r u c e I n f e c t e d  Variable *1  Region *2  N  PSIN PSIN  1 2  PPIN PPIN  DIFF DIFF  Minimum  Maximum  Mean  11 0.125 9 0.560  0.560 0.897  0.389 0.775  0.130 0.102  1 2  11 0.429 9 0.824  0.923 1.000  0.747 0.931  0.158 0.061  1 2  11 0.028 9 0.020  0.699 0.316  0.358 0.156  0.203 0.109  *1 P P I N = P r o p o r t i o n o f P i n e I n f e c t e d PSIN = P r o p o r t i o n o f S p r u c e I n f e c t e d DIFF = D i f f e r e n c e i n P r o p o r t i o n s o f P i n e Infected *2 1= P r i n c e 2- P r i n c e  George Rupert  Forest Forest  Region Region  standard Deviation  and Spruce  Table  12  67  Two Sample T - t e s t on the d i f f e r e n c e i n i n f e c t e d by geographic r e g i o n .  Region  N  Prince George Pr i n c e Rupert  1 1  0.358  0.041  9  0.1 56  0.012  Test  Df  Significance  T=2.680  18  0.015  Mean D I F F  proportions  Var i a n c e  Table  13 68  Grouping of E c o l o g i c a l A s s o c i a t i o n s M o i s t u r e and N u t r i e n t Status  Group  Ecological  Associations  by  N  Included PR  Group  Group  Group  1:  2:  3:  SBSk3 01 : P r i c k l y R o s e - Q u e e n ' s Cup S B S e l 01 : M e s i c B u n c h b e r r y - Moss SBSe2 01 : B u n c h b e r r y - Moss  3  3 3  SBSd 07 : S u b m e s i c B u n c h b e r r y SBSd 08 : M o i s t S h r u b - F o r b  2 1  2  9 * PG = PR =  Prince Prince  George Rupert  Forest Forest  Region Region  1 5  SBSk3 04.1 : W i l d Sarsaparilla - P r i n c e ' s Pine S B S e l 04 : S u b m e s i c B u n c h b e r r y -Moss SBSe2 04 : D o u g l a s f i r - S u b a l p i n e f i r - Moss  * PG  1 1  69  Sarsaparilla  - Prince's Pine. The  t h i r d group included the  Spruce Subzone (SBSd) a s s o c i a t i o n s which d i d not Group 1 or 2 due  f i t in w i t h  to d i f f e r e n c e s in moisture or nutrient  regimes. These were: SBSd 07: Submesic Bunchberry - Moss and SBSd 08: Moist  Shrub - Forb. The  the d i f f e r e n c e in pine and s i g n i f i c a n t at the 0.05  The  infection  e f f e c t of these groups on  spruce infected  l e v e l (Table  DIFF was  not  14).  rates for both species were higher  in the  Prince Rupert Forest Region than in the Prince George Forest Region (Table 11). This may  be due  to the fact that  Prince Rupert stands were older. Older for greater  the  stands would allow  i n t e n s i f i c a t i o n of i n f e c t i o n centers. Such  i n t e n s i f i c a t i o n would l i k e l y r e s u l t rates within i n f e c t i o n centers and  in higher  infection  the expression  of more  advanced symptoms (rot e t c . ) . P l o t s in the Prince Rupert region had more developed symptoms (as well as having infection  higher  rates).  While PSIN and PPIN were both higher the d i f f e r e n c e between them, DIFF was  in Prince Rupert,  l e s s . This was  due  to  a greater average d i f f e r e n c e in PSIN than PPIN between regions  (0.386 as compared to 0.184). A regression of DIFF  on s o i l pH resulted in no s i g n i f i c a n t regression found.  being  Table  14 70  A n a l y s i s of v a r i a n c e of the D i f f e r e n c e l n the P r o p o r t i o n o f P i n e and S p r u c e I n f e c t e d by G r o u p i n g s o f E c o l o g i c a l A s s o c i a t i o n s .  M o d e l : D i f f e r e n c e i n t h e P r o p o r t i o n o f P i n e and S p r u c e i n f e c t e d = Groupings of E c o l o g i c a l A s s o c i a t i o n s Source  Df  Sum o f Squares  Mean Square  F-stat  Signif.  2  0.149  0.075  2.270  0.134  Within  17  0.559  0.033  Total  19  0.708  Between  71 The  d r i v i n g factor behind  could  n o t be d e t e r m i n e d  stand  age  o r t h e age  responsible. the  rooting  rooting grafts that  could between  biotic  species  could  affecting fungal  there  could  of root  and  (inoculum and  affect  levels  on  o f t h e two  p l o t s were  species  found  All  were c o n s i d e r e d  low  susceptibility  the  p r o b a b i l i t y (P) o f  subalpine centers.  spruce  and  in  root  It i s possible of the  two  (thus e f f e c t s on  have  different  infection, affect  the  thresholds  then  site  infection  differentially.  canopy  t o be  to this  subalpine  f i rlocated  i n f e c t e d with  disease.  infection  This  would  indicate a  A t a 5% c o n f i d e n c e  would have  t o be  would c l e a r l y  0.11  t o be  or l e s s w i t h i n  represent  within  I ^ tomentosus.  such  the p r o b a b i l i t y of i n f e c t i o n  f i r would have This  contacts  and d i f f e r e n t i a l  t o be c h a l l e n g e d .  ( ( 1 - P ) * * 2 5 = 0 . 0 5 ) . Thus  Differences  on t r e e v i g o u r  potential could  None o f t h e 25 m a i n study  be  p o t e n t i a l ) are a l s o p o s s i b l e . If  interior  inoculum  stated,  susceptibility.  inoculum p o t e n t i a l f o r s u c c e s s f u l  effects  region  e f f e c t s on  factors affecting rooting  e f f e c t s of the s i t e  pine  species.  infected roots.  r e s i s t a n c e mechanisms)  lodgepole  could  be d i f f e r e n t i a l  t h e number  differentially  vigour  center  tree  healthy  with  s t u d y . As p r e v i o u s l y  o f t h e two  and a b i o t i c  Differential  of  affect  in this  of the i n f e c t i o n  As w e l l , patterns  the v a r i a t i o n i n DIFF  a much l o w e r  level that  in infection infection  72  rate  than  on s p r u c e  However, not  or pine  i n these  t h e symptomology  of i n f e c t e d  w e l l documented. T h e r e f o r e ,  infections involved  this  made on i n f e c t e d plots  study.  In f a c t ,  subalpine  i n d i c a t e d that  this  staining present  observed  staining area  (from  which  attitude.  distal  decadent  meters).  root  in close Brown  r o t were u s u a l l y roots.  Cambial  i s o l a t e d ) was  portions. Generally,  only  a short  f i r i s currently considered  reputed  study  distance  f r o m an  decay.  propensity  f o r decay  crop  by many i n t h e s p e c i e s . The  i s one r e a s o n  for this  H o w e v e r , many o f t h e p e r c e i v e d p a t h o l o g i c a l  problems a s s o c i a t e d with  study  pitted  i n d u s t r y t o be an u n d e s i r a b l e  species'  observations  or absent  I_j_ t o m e n t o s u s was l a t e r  of t h e xylem extended  Subalpine forest  developed  p o r t i o n s of i n f e c t e d  i n these  of advanced  subsequent  ( w i t h i n 0.6  collar  i n more d i s t a l  t h e methods  may w e l l h a v e b e e n t h e c a s e .  and the c h a r a c t e r i s t i c  necrosis also  t o the root  using  f i ri s  whether  f i r not l o c a t e d within  Symptoms were g e n e r a l l y p o o r l y proximity  subalpine  i t i s uncertain  w o u l d have been d e t e c t e d  with  stands.  stands  of t h i s  recommended  o f 200  species  this  years  t r e e s p e c i e s a r e b a s e d on or o l d e r . Therefore,  i s required before  or dismissed  a s an a l t e r n a t e  further  i t c a n be regeneration  73 species  f o r tomentosus  Due  t o the dark  root  of  methods  a b o v e was  level I.  of i n f e c t i o n  t o m e n t o s u s was  Douglas-fir  tomentosus caused  was  reports  (Thomas a n d Thomas sample p l o t s , these  sites  similar low  therefore  not obtained  E F F E C T OF  a.  LODGEPOLE PINE  years  1954).  was  tomentosus  impossible.  in culture  the of the  However,  from t e n  o r d e c l i n i n g and t h e r e  Where D o u g l a s - f i r  to  t o be g r o w i n g  along  with  tomentosus species  were p r e s e n t  INFECTION ON RADIAL  increment  at breast  measured f o r 5 7 pine infection,  a n d was  would  no  seem t o  species  occurred  in  w e l l . On many o f and s p r u c e of  the species' suggest  on s u c h  within  were  of t h i s  f a r o u t grew p i n e  vigour  of the pathogen  radial  also  o f low s u s c e p t i b i l i t y  regeneration  4.  The  s t a i n i n g using  n e c r o s i s . The e v i d e n c e  Douglas-fir  susceptibility  levels  root  i t appeared  age. T h i s  suitable  heartwood,  s a m p l e s . A s w e l l , none o f t h e D o u g l a s - f i r  of abnormal  agree with  root  not p o s s i b l e . Determination  s a m p l e p l o t s were d e c a y e d signs  stands.  s t a i n i n g of D o u g l a s - f i r  identification listed  rot infested  sites  apparently  that  i t may  where  be a  high  i n the previous  stand.  INCREMENT  height trees  f o r the l a s t in various  converted  to basal  five  stages area  of I .  74 increment  over the l a s t  b a s a l area Table  f i v e years  increment a r e d e s c r i b e d by i n f e c t i o n c l a s s i n  15. As can be s e e n , h e a l t h y  larger  ( B A I ) . The diameter and  than d i s e a s e d t r e e s  t r e e s were on average  (370.01 s q . cm. b a s a l area as  compared t o 300.66 s q . c m . ) . T h i s d i f f e r e n c e t o be s i g n i f i c a n t u s i n g a T - t e s t  («*=0.05)  was found not  (Appendix C ) .  D i s e a s e d t r e e s showed a 24% r e d u c t i o n i n b a s a l a r e a increment to  i n the 1979-1984 p e r i o d (16.51 s q . cm. as compared  12.55 s q . c m . ) .  The b a s a l area two-factor factors.  increment was used as the v a r i a b l e  in a  ANOVA where subzone and i n f e c t i o n c l a s s were the  B a s a l a r e a increment was found t o c o r r e l a t e  significantly  (*=0.05)  w i t h the b a s a l area at the b e g i n n i n g  of the f i v e year p e r i o d b e g i n n i n g of the f i v e  (Appendix C ) . Thus b a s a l area at the  year p e r i o d was i n c l u d e d i n the  a n a l y s i s of v a r i a n c e as a c o v a r i a t e . Age was not found t o significantly affect not  basal area  increment and was t h e r e f o r e  i n c l u d e d i n the a n a l y s i s . T r a n s f o r m a t i o n s of the  b a s a l a r e a d i d not improve the l i n e a r i t y w i t h b a s a l area therefore  i n c r e m e n t . The i n i t i a l  initial  of the r e g r e s s i o n  b a s a l a r e a was  i n c l u d e d i n the a n a l y s i s of v a r i a n c e as a  c o v a r i a t e without t r a n s f o r m a t i o n . r e l a t i o n s h i p between the v a r i a t e  A strong  linear  and the c o v a r i a t e was  r e q u i r e d f o r the GENLIN a n a l y s i s . The e f f e c t  of  infection  Table Mean  Species  15  75  I n i t i a l B a s a l A r e a and Change i n B a s a l A r e a o f H e a l t h y and D i s e a s e d P i n e a n d S p r u c e  N Infection Status  Mean B a s a l A r e a cm-2  Stand. Dev.  Mean Stand Change Dev. in Basal A r e a cm-2  PINE  24 h e a l t h y 33 d i s e a s e d  370.01 300.66  296.98 237.94  13.12 9.03  10.53 6.04  SPRUCE  21 h e a l t h y 17 d i s e a s e d  245.93 248.69  152.79 136.12  13 .05 12.54  8 . 54 12.23  * at beginning  of f i v e  year  period.  status 16).  was  Thus  not  infection  significant  The  found  not  significant  was  no  did  of  summary exist.  necessary  to detect  b.  INTERIOR  trees  would  in  a  pine.  increment  was  indicate that  i n p r o d u c t i v i t y between time of  the  summarized  that of  a l a r g e r sample  A  spruce summary  demonstrated  also there the  s t u d y ) and  in Table  17.  the  From  differences in productivity  variation size  shown  would  in Table  likely  differences in basal  area  t r e e s were b o r e d and  their  of  the  i s presented the  initial  increment in Table  basal  data 15.  area  of  significantly  lower  than  for diseased  Regression  of  basal  area  increment  basal  showed  area  analysis  cause  increment  area  (Table  15  be  increment  due  SPRUCE  sampled  Initial  0.32)  factors.  Thirty-seven recorded.  This  (at the  l a r g e amount  that  two  area  area  basal  subzone a r e  indicated  the  16).  i t i s apparent  The  to  in basal  variation  basal  (P =  tomentosus d i d not  (Table  f o r each  significant  s u b z o n e on  significant  s u b z o n e s . The  this  by  reduction  effect  mean age  t o be  basal of  area  initial was  variance  as  basal  therefore  area  f o r the  A  trees age  t o be  included  a c o v a r i a t e . Age  spruce  T-Test  healthy  with  increments  trees  to  (Appendix and  be  C).  initial  significant. i n the  was  not  two-factor found  to  .Table Analysis M o d e l : (1 ) TDBA «= INF  of  variance  1. INF 2. SUB  9.02 9.27 3.INF*SUB 27.07 180.05 4 .BA1 5 . R e s i d u a l 351.03  Total  692.14  Mean Square  1 . 2. 2. 1 . 50.  9.02 4.63 .13.53 180.05 7.02  of  Pine  Probability  F-ratlo  0.26 0.52 0.15 0.00  1 .28 0.66 1 .92 25.64  56  Overall  TDBA BA1  Increment  BAl  Df  Sum of Squares  Sour ce (1)  in Radial  + SUB + INF*SUB +  77  16  mean  Overal1 standard Deviation  12.52  3.51  32986 .0  26506. -  (1)  INF SUB BAl TDBA  = = = =  I n f e c t i o n S t a t u s ( h e a l t h y or d i s e a s e d ) B i o g e o c l i m a t i c Subzone I n i t i a l B a s a l Area ( c o v a r i a t e ) T r a n s f o r m e d change i n B a s a l Area (DBA * *  .37105)  Table sumrnar y of  Mean  Stand  17  Ages  b y s u b z one  or  variant  Subzone N or V a r i a n t  Min i mum  Max imum  Mean  Standard D e v i a t i on  SBSk3 SBSel SBSe2 SBSd  8u . 0 80. 0 73 .0 75. 0  104 . 0 104 . 0 151. 0 113 . 0  93. 1 87. 4 101 . 3 101. 0  7. 0 9.6 21. 3 10.3  23 5 9 20  Summary o f Mean  Subzone N or V a r i a n t  SBSk3 SBSel SBSe2 SBSd  Tree  M i n imum  23 95 .03 5 206 .12 96 . 77 9 20 93 . 31  Basal  Areas cm  Maximum  564 1281 844 1158  .10 .97 .96 .10  2  by  iS u b z o n e  Mean  225 .07 546 .66 212 . 58 4 79 . 59  or V a r i a n t  Standard Dev i a t i on  1 1 5 . 84 437.30 238.91 284.51  79 have a s i g n i f i c a n t the e f f e c t not  of  effect  infection  and was t h e r e f o r e  s t a t u s on b a s a l area  found to be s i g n i f i c a n t  subzone and of  factor  the 0.05  (Table  level  (Table  interactions  18).  spruce were of  spruce. Merler  The d i s c r e p a n c y between M e r l e r ' s  i n increment  a l s o found t h a t  s t u d y . They may a l s o be due to d i f f e r e n c e s  involved in t h i s  well,  Merler's  5.  of  insufficient  study were 70 t o  ones.  sample s i z e s in  the  120 y e a r s o l d . As  study i n v o l v e d stands w i t h h i g h e r stems per h e c t a r e  of  stocking  as compared to 800 stems  The h i g h s t o c k i n g l e v e l  have a m p l i f i e d the e f f e c t stress  infected  sampled t r e e s aged 30 t o 65 y e a r s . The  trees  related  in M e r l e r ' s  i n f e c t i o n due to  study may  density  effects.  CULTURAL CHARACTERISTICS  Attempts  I.  r e s u l t s and those d e s c r i b e d  in this  (1450  at  Merler of  than h e a l t h y  study may be the  per h e c t a r e ) .  result  f i n d i n g s of  in t h i s  densities  of  18).  s m a l l e r average diameter  stand a g e s . M e r l e r  was  were not s i g n i f i c a n t  r e p o r t e d a 20% r e d u c t i o n  tomentosus i n f e c t e d  increment  The e f f e c t s  These r e s u l t s do not concur w i t h the (1984). M e r l e r  e x c l u d e d . Again  to c u l t u r e  I^  tomentosus from c o l l e c t e d  samples were not always s u c c e s s f u l . V i a b l e c u l t u r e s  from  T a b l e 18  Analysis Mode 1: (1) TDBA = INF Source (1) l.INF 2. SUB 3.INF*SUB 4 .BAl 5.Residual Total  of V a r i a n c e  4  in Radial  SUB • lNF*SUB  (1)  INF SUB BAl TDBA  Df  Mean Square  F- ratio  0.0015 0.0028 0.0045 0.0087 0 . 0222  1 . 2. 2. 1 . 31 .  0.0015 0.0014 0.0023 0.0087 0.0007  2 . 15 1 .92 3 . 15 12 . 12  0.0415  37 .  0.654 24716.  = * = =  + BAl  Sum of Squares  O v e r a l l mean  TDBA BAl  Increment of spruce  Probability  0.152 0. 1G3 0.056 0.001  Overall standard Deviation 0.033 14365.  I n f e c t i o n S t a t u s ( h e a l t h y or d i s e a s e d ) B i o g e o c l i m a t l c Subzone I n i t i a l B a s a l Area (covariate) T r a n s f o r m e d change i n B a s a l A r e a (DBA * *  .37105)  lightly In  t o medium  stained  isolation  more a d v a n c e d (1963,  f r o m p o r t i o n s ,of t h e r o o t  symptoms.  rare  or perhaps  staining  in early  reaction  by t h e h o s t t o  Unsanitary contamination a slow  it's  which  (1940) a n d  absent  s t a g e s may  represent  a  l e d to a large  isolation  amount o f  a t t e m p t s . As I ^  p r e s e n c e was  easily  masked a s i t was  successful  least  collected  was  characteristics cultures  samples  each  from  an e i g h t  often  overgrown was  case  and i d e n t i f i e d  rotting  i n Nobles fungi  month p e r i o d ,  I_j_ t o m e n t o s u s  by  more  bench.  s p r u c e a n d p i n e were (105 s a m p l e s ) , I . using  cultural  key t o i d e n t i f i c a t i o n  (1948).  isolates  at a p p r o x i m a t e l y 1 degree  in isolating  per day),  isolation  infected  In each  as o u t l i n e d  o f wood  stored  site.  isolated  tomentosus  o u t on a l a m i n a r a i r f l o w  two s a m p l e s  from  tomentosus  Over  contaminants. Culture  when c a r r i e d  hyphae a r e  physiological  ( a p p r o x i m a t e l y 1.5 mm  At  Whitney  wood. The  fungus  growing)  showed  infection.  conditions  in field  in stained  growing  (faster  rate  Christensen  difficult.  c o n f i r m e d by  1966) p r e s e n t e d e v i d e n c e t o s u g g e s t t h a t  sometimes  of  were p a r t i c u l a r l y  many c a s e s t h e p r e s e n c e o f t h e f u n g u s was  cultural  is  xylem  was  were c u l t u r e d C. A h i g h  from  success  achieved throughout the  82 storage  period  f o r most  other  fungi  sites  impossible after  Fungal color, and  made i s o l a t i o n  growth r a t e  variation  over  both  advancing  the P r i n c e Rupert isolates.  t o golden  Established  advancing but  dark  zone. brown  characteristics case, with  site  produce  A great d e a l of  isolates.  However,  t o dark  were  studied fan-like  found  found  extensions  with  regularity  much d a r k e r  had h y a l i n e  observed  from  was  darken  being with  than the  advancing  zones  mycelium. V a r i a t i o n i n within  the advancing  brown e s t a b l i s h e d with  were  brown w i t h t h e l a t t e r  was o f t e n  isolates  was a l s o  isolates  o f t h e submerged m y c e l i u m  established  an i s o l a t e  o f 50  two s p r u c e a n d on o c c a s i o n i n  mycelium  h y a l i n e mycelium golden  characteristics  t h e submerged mycelium would  Some  hyphal  i n a l l of t h e o b s e r v e d  mycelium  brown  zone  of the c u l t u r e s  The c o l o r  most common. O f t e n age.  including:  f o r a subset  f o r t h e most p a r t .  in  two o f t h e  five'months.  among a n d w i t h i n  growing  c o n t a m i n a t i o n by  from  in culture,  observed  zones  more r a p i d l y  hyaline  approximately  t o occur  of  other  samples  a two month p e r i o d .  was f o u n d  t o be e v e n ,  However,  and t e x t u r e , advancing were  characteristics  The  from  characteristics  mat c o l o r  maintained  samples.  mycelium  hyaline  isolates.  I n one  z o n e o f an  isolate  was s u b c u l t u r e d t o  subsurface mycelium  which  83 remained  such  The authors  with  complete (Whitney  age.  range 1977,  by  variation  d i d o c c u r . The  one  chestnut mat  color  changes  isolates  i n mat  d e s c r i b e d by  Bohaychuk  colors  dark  expressed  color  and  as  1977)  other  was  not  s t u d i e d . However, c o n s i d e r a b l e mat  brown a n d was  colors  Whitney  expressed  tan,  the  of mat  expressed  brown. In by  a  to  cream,  some c a s e s , more  single  corresponded  were  isolate.  increasing  than  Typically, age  of  the  culture.  The  m y c e l i a l mats  downy t o c o t t o n y present  the  i n h a b i t e d agar. the  isolate, less  i n one  aerial sparse  variation  continuous In  clumps i n mat  and  older  cultures,  s u b c u l t u r e d from  t o p r o d u c e more a e r i a l directly  from  and  subsurface  those  with  downy  sparse.  In  one  occurred. There  age.  However,  p r e v i o u s agar  isolates  mycelium  than  combinations  mycelial color  o f mat  was  isolates  young c u l t u r e s  samples.  Some c h a r a c t e r i s t i c texture  similar  mycelium of  texture within of  to  most  specifically  o f downy m y c e l i u m color  aerial  which c o v e r e d  h y p h a e were s c a t t e r e d and  remained q u i t e constant)  isolated  some c a s e s ,  sheet  others,  (they  tended  m y c e l i u m ) were a p p r e s s e d  i n t e x t u r e . In  was  mats,  (aerial  color  existed.  and  Three  such  84 combinations of  this  study  chestnut which  were d e t e c t e d as  m y c e l i u m had  and  the  mat  mat had  No isolates high would  dark was  to  host  be  concluded  existed  the  of  likely  in texture  (Figure  was  to produce  these  than  mycelium.  These  The  three  19.  made b e t w e e n p i n e  and  spruce  characteristics studied. b o t h among and  within  The  isolates  ( i f such v a r i a t i o n d i d e x i s t ) .  It could  not  the  r e s u l t s of  this  not  appear  to  from which  isolates 13).  study  No  be  there  plots  than  cultural related to v a r i a t i o n in  the  isolates  were  rudimentary  fruiting  bodies  special conditions  s t r u c t u r e s . However, o n l y  four  that  less variation within  well, variation in  hosts  (Figure  i n Table  three  due  etiology  several  12c).  Type  mycelium  Type  subsurface  mat  isolates  from  the  and  12b).  a  12a).  subsurface  (Figure  had  v a r i a t i o n between  e i t h e r more o r  on  be  h a v e masked any  species  and  (Figure  cultural  did  (greater  down  could  characteristics  produced  velvety  variability  b e t w e e n p l o t s . As  In  brown a e r i a l  purposes  mycelium  cottony  summarized  distinction  Type one  m y c e l i u m and  brown a e r i a l  types are  degree  types'.  f o r the  subsurface  sparse  using  were d e n o t e d  and  dark  texture  texture  mycelium  aerial  typically  two  mycelium  'mycelium  colored  was  and  months s i n c e  isolation),  were older  derived.  were  necessary cultures  produced  these.  Figure 12b I. tomentosus i n c u l t u r e  - Type  2.  86  Figure I.  tomentosus  12c  i n c u l t u r e - Type  3.  Table  19  Common M y c e l i u m T y p e s Found  Type Type Type  1. 2. 3.  Subsur face Color  Aerial  color  Chestnut Dark brown Brown b l a c k  Chestnut Dark brown Dark brown  in Culture  Mat  texture  Cottony Velvety S p a r s e down  Figure 13 Rudimentary f r u i t i n g body of I_. tomentosus.  89 Growth r a t e s f o r i n d i v i d u a l c u l t u r e s ranged from 0.79 t o 1.72 mm. p e r day. Average growth r a t e s  (for a single  host  s p e c i e s o r s i t e ) ranged from 0.89 t o 1.53 mm. p e r day.  D. CONCLUSIONS  The variation  r e s u l t s c l e a r l y show t h a t s t r o n g d i f f e r e n t i a l i n h o s t s p e c i e s s u s c e p t i b i l i t y t o I ^ tomentosus  does n o t e x i s t between l o d g e p o l e  p i n e and i n t e r i o r s p r u c e i n  the a r e a s s t u d i e d . I t was found t h a t t h e s u s c e p t i b i l i t y of the two s p e c i e s v a r i e d i n a p o s i t i v e , d i f f e r e n t i a l , q u a n t i t a t i v e manner. Such v a r i a t i o n was l i k e l y due t o a d i f f e r e n t i a l e f f e c t of a b i o t i c or a b i o t i c f a c t o r  on t h e  s u s c e p t i b i l i t y of t h e h o s t s p e c i e s , r a t h e r t h a n t o t h e p r e s e n c e o f h o s t s p e c i f i c r a c e s o f t h e p a t h o g e n . The l a t t e r c o n d i t i o n would more l i k e l y have r e s u l t e d differential qualitative  i n negative,  variation.  D i f f e r e n c e s were n o t e d i n t h e b e h a v i o r  o f I_j_ tomentosus  on p i n e and s p r u c e and i n t h e above ground symptoms e x p r e s s e d by t h e s e two s p e c i e s . The pathogen o f t e n c a u s e d p r o n o u n c e d n e c r o s i s o f r o o t phloem i n p i n e s w i t h o u t  causing  decay o f t h e h e a r t w o o d . I n f e c t e d p i n e r o o t s were o f t e n  very  r e s i n o u s b u t t h i s r e a c t i o n d i d n o t appear t o s t o p t h e advance o f t h e f u n g u s . I n s p r u c e t h e r o o t x y l e m was o f t e n  90 extensively still the  decayed  alive.  crown  still  Infected  and  showed no  often  crown  disease  of  found  infection on  centers  t o be  there.  standing.  Infected  often  t i s s u e s were  obvious  thinning  spruce  of  commonly  were w i n d t h r o w n  while  In  rate within  between  the  rate  two  on  i n the  variation  as  was  not  two  i n the  i n the  reduction  of  detected  on  alternate  species  tomentosus many  These  studied to the  species. or  species  for regenerating  infestations.  sites  areas  Douglas-fir  sample p l o t s .  i n the  i n the  Douglas-fir  Prince  George  field.  The  to data  spruce  would not by  with be  and  allow their  tomentosus  f i r within  therefore  region.  formed  extrapolate  disease  would  with  region  Inonotus  sites  spruce  studied,  being  subalpine are  was  higher did  s u s c e p t i b i l i t y of  losses  alternate regeneration  centers  a  infection  Rupert  necessarilly  species  than  regions  Prince  infection  does not the  forest  t o be  pine  showed  infection  tomentosus  use  pines  s i t e s . The  that  effective  lodgepole  centers)  infection  for  fact,  considered  infection  levels  to  the  attacked.  However, t h i s  indicate  i s generally  i n B r i t i s h Columbia,  i n more d i s t i n c t  incidence  for  conductive  (within  varied  resulting  of  died  outer  displayed  I_j_ t o m e n t o s u s  rates  higher  pine  pines  symptoms and  spruce  a l l study  the  the  green.  Although  also  while  any  possible  severe  I.  appropriate  Subalpine  f i r is  91  currently interest used  i n the  not  near  found  to  In  be  variation However,  and  spruce,  to have a fact,  in hyphal  in spruce  on  the  However,  i t may  become  widely  was  the  by  effect  basal  tomentosus on  area  basal  area  increment  i n c u l t u r e showed a g r e a t  c o l o r , mat  of  c o l o r and  occurred  No  on  was  the  and  deal  of  texture. and  among  b a s i s of  distinction  from p i n e s  cultural  mat  both within  into races  impossible.  obtained  basis  infection  in infected trees.  classification  isolates  species.  and  significant  tomentosus  characteristics  spruce  i s growing  since v a r i a t i o n  isolates,  between  regeneration  future.  higher  Inonotus  a  species  both pine  increment. found  used as  i n the  For was  not  could  isolates  characteristics.  cultural be  from  made  92 Literature Bakshi, B.K. 1971. Forest practice in forestry. D e l h i . 400 pp.  Cited  P a t h o l o g y : p r i n c i p l e s and C o n t r o l l e r of P u b l i c a t i o n s ,  B a r n a r d , E.L., B l a k e s l e e , G.M., E n g l i s h , J . T . , Oak, S.W. and R.L. A n d e r s o n . 1985. Pathogenic fungi associated with Sand p i n e r o o t d i s e a s e i n F l o r i d a . P i n t . D i s . 69:196199. B a s h a m , J . T . and Z.J.R. M o r o w s k i . 1 9 6 4 . C u l l s t u d i e s , t h e d e f e c t s and associated basidiomycete fungi in the heartwood of l i v i n g f o r e s t t r e e s i n the f o r e s t s of O n t a r i o . Can. Dep. F o r . , O t t a w a . P u b l . 1072, 69 pp. B o y c e , J . S . 1963. Red r o o t and b u t t r o t i n a G e o r g i a p i n e p l a n t a t i o n . P i n t . D i s . Rep. 47(6):572-573. Christensen, C.M. 194 0 . O b s e r v a t i o n s c i r c i n a t u s . P h y t o p a t h . 30:957-963. C o o k e , M.C. 1886. G r e v . XV, 54,  Praecusores 74.  ad  of  Po1ypo r u s  monographia  polypororum  D a v i d s o n , A.G. and D.R. Redmond. 1957. the Maritime p r o v i n c e s . For. Chron. Dyer,  Decay 33(4)  of  spruce  in  E.D.A. a n d D.W. T a y l o r . 1971. Spruce b e e t l e brood production i n l o g g i n g s l a s h and w i n d t h r o w t r e e s i n B r i t i s h C o l u m b i a . Can. F o r . S e r v . I n f . Rep. BC-X-62, 19 pp.  E l l i s , J.B. and B.M. E v e r h a r t . 1889. hymenomycetous f u n g i . J . M y c o l . V. Fox,  Slash  Some new 24-29.  species  of  D.J. a n d K.E. G u i r e . 1976. Documentation f o r Midas. S t a t i s t i c a l R e s e a r c h L i b . , U n i v . of M i c h . 3rd Ed.  Fries,  E.  F r i e s , E. pp.  1821. 1863.  Systema m y c o l o g i c u m , Monographia  I.  351  hymenomycetum  G i l b e r t s o n , R.L. 1974. Fungi that U n i v . o f A r i z o n a P r e s s . 174 pp.  Decay  pp. sueciae,  II,  Ponderosa  268  Pine.  93  G o s s e l i n , R. 1944 . S t u d i e s on P o l y s t i c t u s c i r c i n a t i u s and i t s r e l a t i o n t o b u t t - r o t of s p r u c e . F a r l o w i a 1 ( 4 ) : 525568. G r e i g , M. and J . B j e r r i n g . 1980. UBC GENLIN: A g e n e r a l l e a s t s q u a r e s a n a l y s i s of v a r i a n c e program. Computing C e n t e r , U n i v e r s i t y of B r i t i s h C o l u m b i a . 55 pp. Haddow, W.R. 1941. On t h e h i s t o r y and d i a g n o s i s o f P o l y p o r u s tomentosus F r i r e s . , Polyporus c i r c i n a t u s F r i e s , and Polyporus d u a l i s Peck. Trans. B r i t . Mycol. Soc. 25(2):179-190. Hobbs, S.D. and A.D. P a r t r i d g e . 1979. Wood d e c a y s , r o o t r o t s and s t a n d c o m p o s i t i o n a l o n g an e l e v a t i o n g r a d i e n t . F o r . S c i . 25(l):31-42. Jahn,  H. 1978. D i e G a t t u n g O n n i a P. K o r s t . W e s t f a l i s c h e P i l z b r i e f e . 11:79-83.  K a r s t e n , P.A. 1889. F i n l . Nat. F o l k  Krit. Ofversigt XLVIII.  Finl.  Filzporlinge  Basidsu.  Bidr.  Kondo, E.S. and R.G. T a y l o r . 1985. F o r e s t I n s e c t and D i s e a s e C o n d i t i o n s , B r i t i s h C o l u m b i a and Yukon. 1984. F.I.D.S. Can. F o r . S e r v . I n f o . Rep. BC-X-259. K o r h o n e n , K. 1978. I n t e r s t e r i l i t y groups of annosum. Commun. I n s t . F o r . F e n n . 9 4 ( 6 ) :  Heterobasidion 1-25.  K o s s u t h , S.V. and E . L . B a r n a r d 1983. M o n o t e r p e n e c o n t e n t o f h e a l t h y Sand p i n e and Sand p i n e w i t h r o o t d i s e a s e . F o r . S c i . 29(4):791-797. K r a j i n a , V . J . 1965. E c o l o g y o f W e s t e r n N o r t h A m e r i c a v o l . 1. Dept. of Botany, U n i v e r s i t y of B r i t i s h Columbia, V a n c o u v e r , B.C. Krebill, R.G. 1962. E t i o l o g y o f a J a c k and Red pine p l a n t a t i o n d e c l i n e c h a r a c t e r i z e d by r o o t w e e v i l i n j u r y a n d f u n g a l d e t e r i o r a t i o n . Ph.D. T h e s i s , U n i v e r s i t y of W i s c o n s i n , Madison, Wisconsin. K u h l m a n , E.G. 1969. s o i l . Phytopath. Lai,  S u r v i v a l of 59:198-201.  Fomes  annosus  spores  in  C. 1983. UBC SPSS s t a t i s t i c a l p a c k a g e f o r t h e s o c i a l s c i e n c e s v e r s i o n 9. U n i v e r s i t y o f B r i t i s h C o l u m b i a . 335 pages.  Lloyd,  C.G.  1908.  M y c o l o g i c a l Notes,  XXIX,  376.  94  Lloyd, CG. 1920. M y c o l . 979, 1014, 1018.  Notes  6:933,  949,  953,  964,  976,  M e r l e r , H. 1984. Tomentosus r o o t r o t of w h i t e s p r u c e i n C e n t r a l B r i t i s h Columbia. Master's T h e s i s , U n i v e r s i t y of B r i t i s h Columbia. Vancouver, B r i t i s h Columbia. M o r r i s o n , D.J., Armiliaria 7:242-246.  Chu D. and A . L . S . J o h n s o n . i n B r i t i s h Columbia. Can.  1985. S p e c i e s o f J . P l a n t . Path.  M u r r i l l , W.A. 1904. The P o l y p o r a c e a e o f N o r t h A m e r i c a . 7. The g e n e r a H e x a g o n a , G r i f o l a , R o m e l l i a , C o l t r i c i a and C o l t i c i e l l a . B u l l . T o r r . B o t . C l u b . XXXI 6, 325-46. Murrill, IX,  W.A. 93.  1908.  Polyporaceae,  i n North  American  Flora,  Murrill, IX,  W.A. 93.  1908.  Polyporaceae,  i n North  American  Flora,  M y r e n , D.T. and R.F. P a t t o n 1971. o f P_. t o m e n t o s u s i n p i n e and J . B o t . 49:1033-1040. Nobles, M.K. 1948. Studies Identification of c u l t u r e s J . Res. 26:281-431. P e c k , C.H.  1878.  3 0 t h Ann.  Rep.  E s t a b l i s h m e n t and s p r e a d spruce p l a n t a t i o n s . Can.  in Forest Pathology of w o o d - r o t t i n g f u n g i . N.Y.  S t a t e Mus.  p.  44.  Redmond, D.R. 1957. I n f e c t i o n c o u r t s of b u t t - r o t t i n g i n Balsam f i r . F o r . S c i . 3(1):15-21. Ross,  VI. Can.  fungi  E.W. 1966. I n c i d e n c e o f P o l y p o r u s t o m e n t o s u s i n S l a s h p i n e p l a n t a t i o n s i n the S o u t h e a s t e r n U n i t e d S t a t e s . P i n t . D i s . Rep. 5 0 ( 7 ) : 5 2 7 .  Saccardo,  D.A.  1888.  S y l l o g e fungorum, V I ,  208.  S a r t o r y , A. and L. M a r i e . 1922. Le P o l y p o r u s t o m e n t o s u s F r . Le t y p e , s e s f o r m e s , s e s v a r i e t i e s . A s s o c . F r . A v a n c . S c i . C o n f . 46e S e s s i o n . M o n t p e l l i e r . p . 773-783. Shaw, T.  1985.  Personal  communication.  S h i r a i , M i t s u t a o and H a r a K a h e s u k e , f u n g i . 3 r d Ed. p. 289.  1927.  List  of  Japanese  95 Teng,  S.C. Contr.  1932. A d d i t i o n a l f u n g i f r o m s o u t h e r n B i o l . L a b . S c i . S o c . C h i n a 8:2,25.  China.  T h o m a s , G.P. a n d R.W. Thomas. 1954. S t u d i e s in Forest P a t h o l o g y XIV. Decay of D o u g l a s - f i r i n t h e c o a s t a l r e g i o n o f B r i t i s h C o l u m b i a . Can. J . B o t . 32:630-653. Linger, L. 1984. C a n a d i a n F o r e s t S e r v i c e disease survey c o l l e c t i o n s l i p s . Van  forest  insect  and  G r o e n e w o u d , H. 1956. A root disease complex S a s k a t c h e w a n White s p r u c e . F o r . C h r o n . 32:11-13.  in  W a l l i s , G.W. and G. R e y n o l d s . 1962. I n o c u l a t i o n o f D o u g l a s f i r r o o t s w i t h P o r i a w e i r i i . Can. J . B o t . 40:637-645. Warren, G.L. The e f f e c t o f some s i t e f a c t o r s on t h e abundance o f Hy p o r n o 1 y x p i c e u s (Coleoptera: C u r c u l i o n i d a e ) . E c o l o g y , 37, 132-139. Whitney, R.D. 1960. Stand-opening disease and the p a t h o g e n i c i t y o f P o l y p o r u s t o m e n t o s u s F r . on W h i t e spruce (Picea glauca ( M o e n c h ) V o s s . ) Ph.D. Thesis, Queen's U n i v e r s i t y , K i n g s t o n , O n t . 98 pp. W h i t n e y , R.D. 1961. Root wounds and a s s o c i a t e d W h i t e s p r u c e . F o r . C h r o n . 3 7 ( 4 ) 401-441.  root  rots  of  W h i t n e y , R.D. 1962b. S t u d i e s i n Forest Pathology XXIV P o l y p o r u s t o m e n t o s u s 'Fr. a s a m a j o r f a c t o r i n s t a n d o p e n i n g d i s e a s e o f W h i t e s p r u c e . Can J . B o t . 40:16311658. W h i t n e y , R.D. 1963. A r t i f i c i a l i n f e c t i o n of s m a l l spruce r o o t s with Polyporus tomentosus. Phytopath. 53(4):441443. W h i t n e y , R.D. 1964. H a r d w o o d s a s i n o c u l u m c a r r i e r s f o r P. tomentosus i n o c u l a t i o n s . Phytopath. 54(12):1484-1485. W h i t n e y , R.D. 1964b. I n o c u l a t i o n o f e i g h t S a s k a t c h e w a n t r e e s w i t h P_. t o m e n t o s u s . C a n . Dep. F o r . B i m o n . P r o g . Rep. 20(5):3. W h i t n e y , R.D. 1966. G e r m i n a t i o n and i n o c u l a t i o n t e s t s w i t h b a s i d i o s p o r e s o f P o l y p o r u s t o m e n t o s u s . Can. J . B o t . 44:1333-1343.  96 W h i t n e y , R.D. 1 9 7 7 . P o l y p o r u s t o m e n t o s u s r o o t r o t of c o n i f e r s . C a n . F o r . S e r v . S.S. M a r i e , O n t . F o r . T e c h . Rep. 18. W h i t n e y , R.D. 1978. P o l y p o r u s t o m e n t o s u s r o o t and b u t t r o t o f t r e e s i n Canada. S t a t e m e n t p a p e r f o r 5 t h I n t . C o n f . ( K a s s e l , Germany) Aug. 1978 on P r o b s . o f Root a n d B u t t Rot i n C o n i f e r s . IUFRO Germany 1980. Whitney,  R.D. 1985. P e r s o n a l  communication.  W h i t n e y , R.D. and W.P. Bonaychuk. 1976. P a t h o g e n i c i t y o f P. t o m e n t o s u s a n d P_. t omen t o s u s v a r . c i r c i n a t u s o n s e e d l i n g s o f e l e v e n c o n i f e r s p e c i e s . Can. J . F o r . Res. 6 ( 2 ) :129-133. W h i t n e y , R.D. a n d W.P. Bonaychuk. 1977. V a r i a t i o n of P o l y p o r u s tomentosus i n c u l t u r a l c h a r a c t e r i s t i c s and pathogenicity of conifer seedlings. Can. J . B o t . 55(10):1389-1395. W h i t n e y , R.D. and W.B. Denyer 1969. H e a r t w o o d a s a b a r r i e r t o i n f e c t i o n o f w h i t e s p r u c e by h e a r t r o t t i n g f u n g i . F o r . S c i . 15(3):266-267. W h i t n e y , R.D. a n d H. v a n G r o e n e w o u d . 1 9 6 4 . T h e r a t e o f advance of stand-opening over a t e n year p e r i o d i n White spruce at Candle Lake, Sask. F o r . Chron. 40(3):308-312. Wood, C.S., Van S i c k l e , G.A. and T . L . S n o r e . 1984. F o r e s t i n s e c t and d i s e a s e c o n d i t i o n s , B r i t i s h Columbia and Yukon 1 9 8 3 . F.I.D.S. C a n . F o r . S e r v . I n f o . Rep. BC-X246 32 p p .  APPENDIX A  Maps of Study  Plots  98  PG 1 20  Scale  19 -  0  ie-  4  =  It  200  v  17 -  v  1615-  $  •  +  +  0 +  1413-  iH  A  •  10 H  *  +  9 -  +  7-  +  A  0  v  A  6-  +  A  4 2 1 Q-\—i—|—|—l—I—I—I—I—I  2  I  4  0  Healthy  spruce  0  Healthy p i n e  &  6  I  10  I  1  12  A  Infected  +  Infected pine  r  14  spruce  1t>  tt  99  PG 2 20  Scale -  1:200  I 12  1  ieH 17-  1& •  V  O  +  15-  D  1413-  v  A  12-  11 10  e X  7  A  &  5 4 3 2 1  0  -j  0  1 2  1  1  4  0  Healthy  spruce  O  Healthy  pine  X  Subalpine f i r  1—I  1  1 e  l>  1—I 10  1  I  1  Infected spruce  -f  Infected pine  y  Douglas f i r  " 14  1  15  1  r  ie  100  PG 3 10  16 17 1& 15 14 -  Scale -  19  13 12 11 10 -  X  0.  0  0  A  M.  A  +  X  0  675543-  210  ^  *  +•  +  +  1  2  v  D  0  . +  0  A  1  A  A  X  0  1:200  A  1  4  1  1  1  1  &  6  1  1  10  1  1  1  12  v  Healthy  spruce  A  Infected spruce  Q  Healthy  pine  -f  Infected pine  X  Subalpine f i r  y  Douglas f i r  1  14  1  1  1&  1  1  16  T  2>  101  PG 4 20-  S c a l e » 1x200  19ie17151$ A  14  vi  v  3  +0  +  A  iy 12 11 10 9 6  A  7  V  +  A  &  5  ++ A  4 3 2 1 0  -i—i—i—i—i—i—i—i—i—i—i—i—i—i—i—i—i—i r~ 0 2 4 & 6 10 12 14 1& 16 20  v  Healthy  spruce  A  Infected  spruce  •  Healthy  pine  -j-  Infected  pin*  X  Subalpine f i r  jf  Douglas f i r  102  PG 5 20  Scale «  19 -  1:200  ie171&-  A  154+  1413-  +  4  0  1211 -  0  0  10- +  +  9-  V  e-  +  +  +  0  +  7 a  +0  +  5 4 3H 2  1 H 0 -i—i—i—i—i—i—i—i—i—i—i—i—i—i—i 0 2 4 & e 10 12 14 0  Healthy  spruce  D  Healthy  pine  A  Infected spruce  +  Infected pine  i to  i  i r 16 20  103  PG 6 Scale "  H200  20  19 H  •  1ft-  +  17-  +  n15-  A  •  +  v  14+  +  +  A  v  + • *  « 1  +  .  A  11 -  A  +  0  10-  9-  +  v  ft -  .  7H  *  +  v •  &H 5-  43-  2-  10 - | — i —2 i — l — I4— l — l&— l — l$— I — l10 —I  1  12  1  v  Healthy  spruce  A  Infected  spruce  0  Healthy  pine  +  Infected  pine  1  14  1  1  15  1  1  1*  r  104  PG 7 20  S c a l e « 1x200  19 -  17-  it>-  0  1514-  0  A  + 0  1312-  +  11 -  +  10A  9 -  +  + +  vD  e7-  +  5-  V  5-  + A  4321 0-  |  "1  2  ^ Q  1  4  1  1  1  5  1  S  1  1  10  1  1  1  12  Healthy  spruce  A  Infected spruce  Healthy  pine  •f  Infected pine  1  14  1  1  It  I  I  16  T  2}  105  PG 8 20  Scale =  n -  1:200  1715151 4  "  .  13-  A  *  12-  • A  10 H  0 A  +.  9 -  7i 5  • +  11 -  ft-  +  v  A  +  0  A A  +  +  +  v  *  5 H  +  +  321 OH—i—i—i—i—i—i—i—i—i—i—i—i—i—i—i—i r 0 2 4 5 f t 10 12 14 15 0  Healthy  spruce  A  Infected spruce  D  Healthy  pine  -{-  Infected pine  1ft  20  106  PG 9 20  Scale = Ii200  19-  1617 It 15 14 13 12 11 10 9  X X  A  X X X  + A  e  7 &  5 4 3 2 1 0  X •  n—i—i—i—i—i—i—i—i—r 0 2 4 t 6 10  n—i—i—i—i—i—i—r 12 14 1t 16 2}  0  Healthy  spruce  A  Infected spruce  fl  Healthy  pine  +  Infected pine  X  Douglas f i r  107  PG 10 20 -i 19 -  A  •  S c a l e •> l i 2 0 0  A  14+•  17+  1&-  V  15+  14-  A +  A A  + A  A  + + A  1312-  +  11 -  v  + A  A  109 -  0  475 -  0  +  54 -  V  +  321 Si  1  0  1  1  2  1  4  1  I  1  5  i  4  I  I  10  1  !  1  12  V  Healthy  spruce  A  Infected spruce  •  Healthy  pine  •+•  Infected pine  s  1  14  1  1  15  l  l  14  l  20  108  11  PG 2019 -  Scale •  1:200  16A •  17-  0  + +  0  16-  +  1514-  A  13-  V  V  12-  A  11 -  +  +  •  10-  A  A  Aa  +  +  9 -  +  e7-  +  V  A  65-  +  A  A  43-  V  2-  1A  0  1  0  1  1  2  1  4  1  1  1  &  1  6  1  1  10  1  1  1  12  0  Healthy spruce  •  Infected spruce  O  Healthy pine  +  Infected pine  1  14  1  1  1&  1  1  16  1  2>  109  PR  1  20 S c a l e «=  19 -  ie-  A  17-  A A  i&+  15-  v  +  14 13 H 12  A  A  + ° *  +  1H 10  1:200  A  + +•  +  4  A  +  +  A  A ^  :i  A  A  7  +  + +  & 5  A  4 3 2101—i—i—i—i—i—i—i—i—i—i—i—i—i—i—i—i—i—i—r $ 10 12 14 15 W 20 v*  Healthy spruce  A  Infected spruce  D  Healthy pine  +  Infected pine  110  PR  2  £  Healthy  spruce  A  Infected spruce  rj  Healthy  pine  +  Infected pine  Ill  PR 3 20 S c a l e «=  19 H  1:200  14 1715-  A  15-  A  14+  13-  +  12-1  A  «H 10 H 9  ++  +  +  «  •  . + * A  +  A  *  A  H  ft  7H 5-  1  A  A v  4-  * ^  +  321 Oi—i—i—i—r—i—r~~—i—i—i—i—i—I—T—i—i—i—i—r 2 4 & 4 10 12 14 15 14 0  Healthy  pine  +  Infected pine  ^  Healthy  spruce  A  Infected spruce  T  Subalpine f i r  2>  PR 4  112  20-r S c a l e * 1:200  19161716 -  1514-  A D  A  13-  0 1  A  12-  A  11 -  +1 Jr  + A  +  109 -  e-  A  7-  A  6-  A  54-  A  A +  +  321~i 0  i 2  i—i—i—i—i—i—i—i—i—i—i—i—i—i—i—i—r~ 4 I A 10 12 14 n 16 2£>  D  Healthy  pine  "t"  Infected pine  ^  Healthy  spruce  A  Infected spruce  113  0  Healthy  pine  4  Infected pine  ^  Healthy  spruce  £  Infected spruce  114  PR 6 20-r  Scale »  +  19 -  H200  ie-  + -  17151514131211 -  109676543-  +  2-  +  10-  i—i—i—i—i—i—i—i—i—i—i—i—i—r 2  4  6  e  10  12  0  Healthy  pine  +  Infected pine  v*  Healthy  spruce  A  Infected spruce  X  Subalpine f i r  14  n—i—i—r  H  ia  20  115  PR 7 20 S c a l e = 1:200  ie17-  K'  +  +  1514-  A +  v  A  A  X t  15-1 12 11 10H  4  A  + +  + +  *  +  A  i  +  9 -  en  A  +  o  +  A  +0  7-1 A  6-  +  +  545-j  ++ +  2 1 01—i—i—i—i—i—i—i—i—i—i—i—i—i—i—i—i—i—r  2  4  t  $  10  12  •  Healthy  pine  +•  Infected pine  0  Healthy  spruce  A  Infected spruce  %  Subalpine f i r  14  16  16  20  116  PR 8 ^  +  1  19 H  IH  +  •  S c a l e - 1«200  +  17 „  IH  *  +  1  1 H  +  «  D  +  A  14 H  iH  +  +  +  IH  +  k  +  I H  *  +  » -  »-  »  7-  +  65 -  +  D  +  +  4  A  4J2 1o H — i —2i — i —4i — i —5 I I I —6 i — i — — i — i15 — i — i16— r 10 i — i — 12 i — i 14 D  Healthy p i n *  4  Infected pine  ^  Healthy spruce  A  Infected spruce  2>  PR ^ 20  S c a l e - 1:200  19 1ft+  +  1715 1514-  +  x  •  13-  +  12-  4  11 " A 10-  0  • A  v X  A  + 4  A+  + +  9 -  4  4  ft-  0  7  +  +  5 5  <•  A *  f  0  4  A  3  0  A  2101—i—i—i—i—i—i—i—i—i—i—i—i—i—i—i—i—i—i—r 0 2 4 & ft 10 12 14 15 1ft 2> •  Healthy  pine  4-  Infected pine  4  Healthy  spruce  A  Infected spruce  X  Subalpine f i r  Appendix B The e c o l o g i c a l a s s o c i a t i o n s l i s t e d below:  included  118 i n t h i s study are  symbol  Ecological Association  Moisture Regime  Nutrient Regime  SBSd 07  Submesic Bunchberry - Moss  Mesic t o Submesic  Oligo to Mesotrophic  SBSd 08  M o i s t Shrub - Forb  Mesic t o Subhygric  Submeso t o Subeutrophic  SBSel 01 Mesic Bunchberry - Moss  Mesic  Mesotrophic  SBSel 04 Submesic Bunchberry - Moss  Submesic  Oligo to Mesotrophic  SBSe2 01 Bunchberry - Moss  Mesic  Submeso t o Permesotroph  SBSe2 04 Douglas f i r - S u b a l p i n e f i r  s u b x e r i c t o Submeso t o Mesotrophic Submesic  SBSk3 01 P r i c k l y Rose - Queen's Cup  Mesic  Submeso t o Permesotroph  SBSk3 04.1 Wild S a r s a p a r l l l a P r i n c e ' s Pine  Submesic  Submeso t o Mesotrophic  APPENDIX  Statistical  C  Te  Two  Sample T - T e s t on t h e I n i t i a l B a s a l of H e a l t h y and I n f e c t e d P i n e .  Mean I n i t i a l Basal Area (cm s q )  Area  Standard D e v i a t i o n (cm s q )  Variable  N  Healthy  24  370.01  945.46  Infected  33  300.66  752.42  Test  Df  Significance  T=0.97483 F=1.5789  55 23,  32  0.334 0.115  121 R e g r e s s i o n of the Change i n B a s a l Area w i t h I n i t i a l B a s a l Area  Source  Sum of Squares  Df Mean Square  F-ratlo  Probability  1 0.183E+8 47.363 55 0.386E+6  .000  Mult R = 0.680  R-Square = 0 .463  SE - 612.08  Variable  Coeff  S t d . E r r o r T-Stat  Signif.  364.26 .022  132.04 .003  .008 .000  R e g r e s s i o n 0.183E+8 Error 0.212E+8 Total  Constant I n i t i a l BA  0.395E+8  Partial  .680  2.759 6.882  Two  sample T - T e s t on t h e i n i t i a l B a s a l of H e a l t h y and I n f e c t e d Spruce  Area  Variable  N  Mean b a s a l a r e a cm-sg  Standard Deviation  Healthy  21  245.93  152.79  Infected  17  246.69  136.12  Test  Df  Significance  36 20, 16  .953 .322  Stat.  T= -.058 F= 1.259  The p r o b a b i l i t y o f h e a l t h y i n f e c t e d t r e e s i s 0.52  trees  being  larger  than  

Cite

Citation Scheme:

        

Citations by CSL (citeproc-js)

Usage Statistics

Share

Embed

Customize your widget with the following options, then copy and paste the code below into the HTML of your page to embed this item in your website.
                        
                            <div id="ubcOpenCollectionsWidgetDisplay">
                            <script id="ubcOpenCollectionsWidget"
                            src="{[{embed.src}]}"
                            data-item="{[{embed.item}]}"
                            data-collection="{[{embed.collection}]}"
                            data-metadata="{[{embed.showMetadata}]}"
                            data-width="{[{embed.width}]}"
                            async >
                            </script>
                            </div>
                        
                    
IIIF logo Our image viewer uses the IIIF 2.0 standard. To load this item in other compatible viewers, use this url:
http://iiif.library.ubc.ca/presentation/dsp.831.1-0097051/manifest

Comment

Related Items