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Tomentosus root rot of white spruce in central British Columbia Merler, Hadrian 1984

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TOMENTOSUS ROOT ROT OF WHITE SPRUCE IN CENTRAL BRITISH COLUMBIA by HADRIAN MERLER B.Sc,  The U n i v e r s i t y Of B r i t i s h Columbia, 1982  A THESIS SUBMITTED IN PARTIAL FULFILMENT OF THE  REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE in  THE  FACULTY OF GRADUATE STUDIES Department Of F o r e s t r y  We accept t h i s t h e s i s as conforming to the r e q u i r e d  THE  standard  UNIVERSITY OF BRITISH COLUMBIA May 1984  ©  Hadrian M e r l e r , 1984  In  presenting  this  thesis  in  partial  fulfilment  of  requirements for an advanced degree at the U n i v e r s i t y of Columbia,  I  available  for  permission  agree  for-  purposes may or  her  that  the  Library  shall  reference  and  study.  I  extensive  It  is  for  allowed without my  permission.  Department of  Date:  financial  Forestry  The U n i v e r s i t y of B r i t i s h 2075 Wesbrook Place Vancouver, Canada V6T 1W5  24 August  1984  it  freely  agree  Columbia  Department or  understood  p u b l i c a t i o n of t h i s t h e s i s written  further  British  that  copying of t h i s t h e s i s f o r s c h o l a r l y  be granted by the Head of my  representatives.  make  the  gain  that  by  his  copying or  shall  not  be  i i  Abstract Root stands  disease  in 50-70 y e a r - o l d f u l l y  in c e n t r a l  fashions.  The  e s t a b l i s h the  British first  Columbia  method  stocked white  was  was  investigated  spruce in  two  done by root excavation;  i d e n t i t y of the c a u s a l  organism,  to  investigate  the e t i o l o g y , and assess the damage caused by the d i s e a s e . second method i n v o l v e d , a survey estimate was  the i n c i d e n c e of the d i s e a s e .  the  only  pathogen  r e s u l t e d when r o o t s Although  tree  involved.  contacted  species  other  i n f e c t e d r o o t s , only spruce consistent  crown  symptoms  root weevil H y l o b i u s warreni survey occupied  estimated by  20%  season  that  diseased  for whole stands, one  of s i x 1.2  as  was  Infection  than  were in contact  apparent.  Wood was  trees.  of spruce  i n f e c t e d by the  of  the  Of these  r e d u c t i o n of b a s a l area increment.  measure the m o r t a l i t y caused by the  rot.  A complex with  with No the  R e s u l t s from the  area  investigated  22.6%  had butt r o t , and  1% of the standing t r e e s a r e s u l t of d i s e a s e .  trees  inoculum.  root  evident.  to  tomentosus F r .  existing  spruce  The  in order  previously  were  28.4%  ha stands  Polyporus  to  were  Diseased I t was  fungus.  windthrown  was  in  trees suffered a not  possible  to  i i i  Table of Contents Abstract L i s t of Tables L i s t of F i g u r e s Acknowledgement  i i iv v vi  Chapter I INTRODUCTION  1  Chapter II A REVIEW OF THE LITERATURE  3  Chapter I I I METHODS  11  1. SELECTION OF EXPERIMENTAL AREAS  11  2. PLOT SELECTION  13  3. EXPERIMENTAL PROCEDURES  14  4. INCIDENCE SURVEY  16  Chapter IV RESULTS AND DISCUSSION  .18  1. IDENTIFICATION OF PATHOGENS  18  2. CROWN SYMPTOM DEVELOPMENT  19  3. INFECTION AND SPREAD  22  4. SUSCEPTIBILITY OF CONIFER SPECIES  27  5. IMPACT OF TOMENTOSUS ROOT ROT 5. 1 Survey 5.2 E s t i m a t i o n Of Losses 5.2. 1 Butt Rot 5.2.2 Increment Loss 5.2.3 M o r t a l i t y And Windthrow 6. ECOLOGICAL CLASSIFICATION Chapter V CONCLUSIONS  31 31 32 32 34 40 43  LITERATURE CITED  48  46  APPENDIX A - ECOLOGICAL SITE CLASSIFICATION AND VEGETATION LISTINGS 52 APPENDIX B - STEM MAPS OF THE TEN STUDY PLOTS  57  iv  APPENDIX C - RELATIONSHIP BETWEEN TREE SIZE AND BASAL AREA INCREMENT 68  V  L i s t of Tables  1. Frequency of i s o l a t i o n of P. tomentosus as a percent of a l l fungal i s o l a t i o n s i n Canadian decay s t u d i e s 5 2. Average d i s e a s e r a t i n g ( s c a l e : 1-6) of s e e d l i n g s i n o c u l a t e d with Polyporus tomentosus and the v a r i e t y c i r c i n a t u s (from Whitney and Bohaychuk 1976) 3. Average age, area, s t o c k i n g , and volume of spruce in the p l o t study  6  found 14  4. Incidence of tomentosus root d i s e a s e f o r a l l l i v i n g s p e c i e s examined  tree 29  5. Incidence of Polyporus tomentosus root r o t i n white spruce w i t h i n the P r i n c e George survey area ( F i g u r e 1). 31 6. Number of white spruce by age c l a s s i n a l l p l o t s , showing v a r i o u s disease symptoms  33  7. Equations of the r e g r e s s i o n l i n e s f o r the l o g a r i t h m of basal area versus the l o g a r i t h m of diameter squared times height and r e s u l t i n g t e s t f o r common slope 36 8. A n a l y s i s of c o v a r i a n c e t a b l e based on the n a t u r a l l o g a r i t h m of b a s a l area increment, using the n a t u r a l l o g a r i t h m of D.B.H. squared x Height as c o v a r i a t e (confidence = 0.95)  36  9. N a t u r a l logarithms of 5 year b a s a l area increment f o r healthy and d i s e a s e d t r e e s 37 10. T o t a l number of spruce t r e e s i n v e s t i g a t e d , with the percentage i n f e c t e d and percentage dead  40  11. Percent of the t o t a l number of spruce t r e e s by age c l a s s 42 12. Percent of t o t a l spruce t r e e s by diameter c l a s s  43  vi  L i s t of F i g u r e s  1. L o c a t i o n of the study p l o t s ( • ) (Table 3) and the i n c i d e n c e survey stands ( ° ) (Table 5 )  12  2. A blown down spruce at the center of p l o t B4.  13  3. F r u i t i n g s t r u c t u r e of Polyporus tomentosus emerging from a 4 cm diameter stump of a destroyed white spruce. ...19 4. F r u i t i n g s t r u c t u r e of Polyporus tomentosus at the base, of a white spruce 19 5. Exposed r o o t s of white spruce. Note the t y p i c a l white pocket appearance of a decayed root at the c e n t e r . ...19 6. A windthrown white spruce due to complete s t r u c t u r a l and f u n c t i o n a l f a i l u r e of the r o o t s . Red-stain appears i n the f i r s t notch i n the l o g 22 7. Root g r a f t i n g of spruce r o o t s , and red s t a i n Polyporus tomentosus  from  23  8. Exposed root c o l l a r of white spruce. Stem occurs at l e f t , an a d u l t weevil i s at the center of the r o o t , and a pupal chamber l i e s at bottom l e f t (arrows) 26 9. Mean b a s a l area increment over time f o r healthy white spruce, and t r e e s i n f e c t e d with Polyporus tomentosus.  38  10. S o i l p i t from J e r r y Creek with w e l l d r a i n e d sandy loams. The s o i l p r o f i l e had a t h i n organic l a y e r , with an e l u v i a t e d Ae zone, and an e n r i c h e d Bf l a y e r 43  vi i  Acknowledgement  I  would l i k e to thank A s s o c i a t e P r o f e s s o r Dr. B.J. van der Kamp  for both h i s guidance and c o n t r i b u t i o n s to t h i s his  help  study.  Without  t h i s work would not have been completed. I would  to thank Dr. D.J. M o r r i s o n , whose suggestions drew  my  like  interest  to t h i s study, and to Dr. R. Whitney whose research i n Polyporus tomentosus i n s p i r e s , and whose personal a t t e n t i o n s encouraged. I would  also  like  to  thank  the  staff  M i n i s t r y of F o r e s t s f o r t h e i r c o o p e r a t i o n field  work. In a d d i t i o n  of the B r i t i s h Columbia in aiding  me  in  the  I would l i k e to thank my wife, M a r i l y n ,  whose endurance I always marvel a t . T h i s p r o j e c t was Forestry. Service Research  by  to  (Environment  Universities  material discussed report  funded by a  in  contract Canada)  Forestry  with  under  the  the Program f o r  (PRUF).  Most  of  the  i n t h i s t h e s i s i s a l s o presented i n the f i n a l  the Canadian F o r e s t r y S e r v i c e e n t i t l e d  of spruce in c e n t r a l B r i t i s h Columbia" which the C.F.S. i n March  Canadian  was  "Root d i s e a s e submitted  1984 as r e q u i r e d by the PRUF c o n t r a c t .  to  1  I.  It  i s well  INTRODUCTION  known  that  (Moench) Voss, P. engelmani i c e n t r a l B r i t i s h Columbia root  decays.  tomentosus  Fr.,  are a f f e c t e d by a number  of  butt  and  mellea  Polyporus  (Fr.)  sulphureus  Kumm.,  Polyporus  B u l l , ex  F r . , and  In a d d i t i o n to butt r o t , these fungi may  standing mature spruce, or lead to windthrow of l i v i n g  There  i s l i t t l e doubt that endemic p o p u l a t i o n s of spruce  bark b e e t l e (Dendroctonus trees  (Picea glauca  Parry, and t h e i r h y b r i d s ) stands of  F r . , Armiliar ia  Flammula a l n i c o l a F r . .  trees.  mature spruce  Among the common pathogens i n v o l v e d are Polyporus  schweinitzi i  also k i l l  the  rufipennis Kirby) survive  largely  weakened or windthrown as a r e s u l t of these d i s e a s e s .  mature spruce f o r e s t s these pathogens lead to a  steady  on In  decline  in volume and v a l u e . Little  is  Studies in Whitney stands  Saskatchewan,  1977a, can  information  also on  B r i t i s h Columbia. hold  known about  the b e l i e f  Ontario  Gosselin be  these d i s e a s e s i n immature stands.  There  diseases  There in  1962,  immature spruce  i s only  fragmentary  of  that spruce stands w i l l  forest  remain  This  may  managers  to  f r e e of root and be  due  to the  to recognize root d i s e a s e damage that does not r e s u l t  i n obvious group typically  (Whitney  immature stands i n c e n t r a l  i s a tendency  butt r o t s u n t i l a f t e r r o t a t i o n age. inability  Quebec  1944) have shown that  affected.  these  and  dying.  overlooked.  i n f e c t i o n by P.  Infections There  are  in some  immature reports  tomentosus at an e a r l y age.  stands  are  of e x t e n s i v e  2  The matter i s of some importance. seen  the  stands  The  last  stands.  Natural  stands  different  commonly  occupied  than  In  such  cases  and  outlives  regeneration hardwoods.  least  lag  these  species.  Natural spruce lag  The  site  is  during  in age,  and  spruce few  of  regeneration therefore involves a  which the s i t e  that the spruce  in the smaller r o o t s ,  spruce  resulting  i s l a r g e l y occupied  T h i s i s of some importance to root means  the  Eventually  t y p i c a l l y show c o n s i d e r a b l e v a r i a t i o n  them are pure.  natural  by mixed stands with a s u b s t a n t i a l component  of aspen, willow and v a r i o u s shrub s p e c i e s . outgrows  most  The  have u s u a l l y a r i s e n f o l l o w i n g a major  d i s t u r b a n c e by f i r e or i n s e c t s .  the  have  i n i t i a t i o n of a l a r g e p l a n t i n g program in spruce.  so c r e a t e d w i l l be rather  stands  decades  disease  by  because  root pathogens w i l l d i e out, at  before  spruce  roots  of  the  new  generation can come in c o n t a c t with them. Spruce  stands  created  by  planting shortly after  w i l l occupy a s i t e much more q u i c k l y . more root rot problems, and  One  may  expect  logging  therefore  a l s o that such problems w i l l  show up  earlier. The  objectives  i d e n t i t y and  frequency  stands,  elucidate  to  of  the present  of the  root  pathogens  mode  of  d e s c r i b e d i s e a s e symptoms, to estimate growth and y i e l d , incidence standing  of  infection  immature and  spruce  spread,  the e f f e c t of d i s e a s e  to design and conduct a survey  of root d i s e a s e , and crop.  study are to determine the  to estimate  to on the  to f o r e c a s t f u t u r e l o s s e s to the  3  II.  A REVIEW OF THE  Most of the information Polyporus Whitney  tomentosus  host-parasite  spruce  i s gained  (1960,1961,1962,1964/ 1972,  northwestern Gosselin of  on  on  Ontario.  1944,  tree  Other  Lachance 1978,  LITERATURE  Solovieff  biomass and wood value both  in p l a n t a t i o n s . same way  The  e t i o l o g y and d i s e a s e impacts  polypore  Polyporus  fungus with  straight  proposed  the  vary  binomial  nomenclature  (Denyer and R i l e y 1927)  boreal  observed the  forest.  only  can  by Whitney  1953, losses and  Both  the  Fr. signifies  the  from place to p l a c e . tomentosus  setae.  This  corresponds  with  (1944) p o i n t e d out, however, that P.  setae  with  by Haddow (1941) in which P.  c i r c i n a t u s has a duplex hooked  deal  and  i n the n a t u r a l f o r e s t ,  v a r . c i r c i n a t u s F r . represents the fungus Gosselin  Saskatchewan  pathogen does not n e c e s s a r i l y behave i n the  i n d i f f e r e n t p a r t s of  The  of  from s t u d i e s done by  1977b) in  studies  interaction  hymenium, and  occur  that  tomentosus  curved  both  Polyporus  fungus found during t h i s study.  setae.  tomentosus v a r . straight  on the same f r u c t i f i c a t i o n .  (1960) as w e l l .  with  This  tomentosus  None of the  and was was  fruiting  bodies examined contained hooked s e t a e . The primarily  fungus has been i d e n t i f i e d due  to  northern hemisphere. western  Russia  Dzieciolowski northern forest  windthrow, The  at  1927),  implicated in studies West  Poland  1955), North C a r o l i n a (Hepting  Idaho  (Whitney  (Hubert 1977b).  1929),  death,  v a r i o u s l o c a t i o n s around the  fungus was  (Solovieff  from c e n t e r s of t r e e  and  from  (Domanski Downs  and  1944),  and across the Canadian Boreal  4  Twenty-one reported  as  species  host  and  trees  two  to  this  1978a).  I t causes root rot  in  species  (Whitney  Table  frequency to  1977a).  w i t h which P.  other  decay  least 1  31  organisms,  conifers  from  are  (Whitney  commercial  summarizes  tomentosus has been  tree  the  relative  isolated  relative  various Canadian c o n i f e r s in Significant  l o s s e s were  found  i n white and black spruce and p o s s i b l y i n lodgepole p i n e .  A study done by Whitney of  of  pathogen i n Canada  at  s t u d i e s d e a l i n g with stem decay. only  varieties  (1962) i n Saskatchewan showed that  a l l the spruce t r e e s examined  carried  some  P.  19.7%  tomentosus  infection. Host  specificitiy  tomentosus and  P.  and  pathogenicity  of  both  tomentosus var. c i r c i n a t u s was  eleven  species  of  conifer  1976).  I s o l a t e s of both  seedlings  evaluated  (Whitney  fungal v a r i e t i e s  were  collected  on spruce near Candle Lake, Saskatchewan.  tomentosus  caused  Polyporus rating  tomentosus  was  obtained  higher  var. c i r c i n a t u s . on  disease The  test  was  the  infected. spruce,  Earlier  lodgepole  using  white  infected tests  spruce,  semi-mature black spruce two  conifers.  r a t i n g s than  ponderosa pine, f o l l o w e d by  conducted  least  Polyporus  disease  Germinants were r a t e d on a s i x being  from  highest  p i n e , white spruce, and e i g h t other c o n i f e r The  for  and Bohaychuk  sporophores  significantly  Polyporus  on  freshly  point  and  s p e c i e s (Table  scale,  "six"  three  germinated  of  being these  the  value  2).  seeds. "one"  the most h e a v i l y species,  and tamarack (Whitney 1964)  black  showed that  i s more h e a v i l y a t t a c k e d than the other  5  Table 1 - Frequency of i s o l a t i o n of P. tomentosus as a percent of a l l fungal i s o l a t i o n s i n Canadian decay s t u d i e s .  Spec i e s  Locat ion  Reference  % of T o t a l I n f e c t ions  Abies amabi1is (Dougl.) Forb. Abies balsamea (L.) M i l l .  Picea  British Columbia  1.0%  Buckland  et al^_ 1949  Canada  0.5%  Basham et a l .  Ontar i o  1.7%  Whitney 1978b  Alberta  85.0%  Denyer and R i l e y  Ontar i o  16.2%  Whitney 1978b  Ontario  17.9%  Whitney 1978b  Alberta  14.0%  Denyer and R i l e y  Eastern 1953  glauca  (Moench) Voss  1953  Picea mariana ( M i l l . ) B.S.P. Pinus c o n t o r t a Dougl. v a r . latifolia  1953  Engelm.  Pinus strobus L.  1.3%  White  B.C.  0.3%  Thomas and Thomas  B.C.  1.4%  Buckland  0.8%  F o s t e r and F o s t e r  Ontario  1953  Pseudotsuga m e n z i e s i i (Mirb.) Franco  1954  Tsuga h e t e r o p h y l l a (Raf) Sarg  B.C.  et a l . 1949 1951  6  Table 2 - Average d i s e a s e r a t i n g ( s c a l e : 1-6) of s e e d l i n g s i n o c u l a t e d with Polyporus tomentosus and the v a r i e t y c i r c i n a t u s (from Whitney and Bohaychuk 1976)  Tree Species  P. tomentosus  All :  P. tomentosus var.  circinatus  Ponderosa pine  4.4  3.2  3.8  Lodgepole  4. 1  3.1  3.6  pine  White  spruce  3.8  2.8  3.3  Black  spruce  3.9  2.2  3.0  Tamarack  3.2  2.6  2.9  Norway spruce  3.0  2.6  2.8  Scots pine  3.6  2.0  2.8  Blue  spruce  3. 1  2.0  2.6  Douglas-f i r  2.7  2.4  2.6  Red  2.4  1 .7  2. 1  2.5  2.0  2.1  3.3  2.4  2.9  spruce  E a s t e r n white  pine  AVERAGE  The fungus i n c u l t u r e does  not  observed than  reaches maximum growth at pH 4.5  grow at a l l at a pH g r e a t e r than 7.0.  that the pH of d i s e a s e d t i s s u e was  that  of h e a l t h y t i s s u e .  the pH of i n f e c t e d roots  i s not  tissue.  limited  by  Whitney  (1960)  considerably  lower  S t u d i e s by Whitney  that the fungus was able to a l t e r  and  (1962) showed  i t s environment by  decreasing  T h i s means that fungal growth i n the pH.  Fungal  survival  i n the s o i l ,  7  however,  would  observed  that  the  be  affected  by  the pH of the s o i l  pH.  Van  Groenewoud  i n areas s e v e r e l y a f f e c t e d by  root r o t were as low as 4.5, while wherever the  top  soil  was seven or higher,  (1956)  the disease  pH  of the  d i d not occur i n any  damaging amounts. Using both a r t i f i c i a l showed that P. growth.  It  and  natural  tomentosus p r e f e r e d grew  best  media,  a rather  Whitney  (i960)  low temperature f o r  from 17 to 22 degrees C e l c i u s , but grew  w e l l at lower temperatures, and had some  growth  at  0  degrees  Celcius. Penetration from  of Polyporus tomentosus i n t o healthy  basidiospores  fungus i s p r i m a r i l y infection  by  not tree  been observed. to  tree  Inoculations  spruce were only  of P.  f r e s h stumps was not s u c c e s s f u l Studies  contact.  by  Whitney  p o i n t of entry  determined.  (Whitney  (1962,1965)  i n t o the host  Death of the host  cortical  cells  i n t e r - and i n t r a c e l l u l a r l y .  still  also  seedlings  of  showed that the mycelium tips  initially.  The  by the pathogen has not been at  the  epidermal  H i s t o l o g i c a l examinations of cells  are  invaded  both  The p e r i c y c l e and the primary xylem  i n f e c t e d , but t o a l e s s e r degree.  living  Inoculation  1966).  t i s s u e begins  of the r o o t .  of  where a wound of  (Whitney 1962).  i n f e c t e d roots showed that the cortex  are  Basidiospore  tomentosus i n t o the roots  does not n e c e s s a r i l y invade through root  and  The spread of the  successful in instances  some s o r t was made on the host  exact  roots  has only been demonstrated i n deep wounds (5 cm deep)  of root wood. white  has  tree  Badly diseased but  seldom had hyphae i n s i d e the endodermis.  8  When the c o r t e x t i s s u e was f u l l y the  root  ensued.  The  invaded,  fungus  the u l t i m a t e death  advances  in  a tree p r i m a r i l y  through the heartwood, but can advance i n and k i l l particularly  i n white and black spruce.  " t h i s i s rare f o r h e a r t r o t c o n s i d e r e d to be t r u l y The roots. brown  root  decay  I t spreads stain,  stages.  fungi,  Whitney  which  appearance. girdled  process  i n the  occurs  secondary  first  1962).  intervals,  In advanced  xylem  resulting  stages  of  causing  formation  are not  in  white  flakes  the  of  a  the  spots  white  large  tree  reddish  i n the l a t e r  leaving  attack,  i s g i r d l e d , at which time  Yellowish  (1962) s t a t e d ,  through small branch  f o l l o w i n g i n v a s i o n of sapwood and bark.  root c o l l a r  bark  parasitic."  followed by a white pocket  at  living  ordinarily  L i g n i n i s decomposed by the fungus,  cellulose  of  of  pocket  roots  are  E v e n t u a l l y the dies  (Whitney  mycelium form beneath the  outer bark s c a l e s and the fungus transmits to healthy roots upon contact  (Whitney  1962).  A statement made by Whitney decay develops  (1962)  that,  "Extensive  before the above-ground symptoms become apparent"  goes l a r g e l y unheeded by many f o r e s t e r s who tend Instead, classic  recognition  of  the  root  disease  root r o t centers and stand openings.  spruce  Absence of symptom noted  l a r g e p a r t s of the root system were k i l l e d on a p p a r e n t l y In  stands  with  high  incidence  symptoms would p o s s i b l y become more e v i d e n t . excessive  branch  mortality  from  stands.  i s based on more  expression was a l s o d e s c r i b e d by Whitney (1965) who  seedlings.  root  of  that  healthy  the d i s e a s e ,  Symptoms  include  the lower p a r t of the crown,  9  shortened height and  lateral  trees,  t h i n n i n g of the crown (Whitney  and  severely  general  increments  i n f e c t e d t r e e s , the  shorter,  needles  relative  were  healthy  1977b).  observed  and having a yellow-green c o l o r .  these needles were c u r l e d at t h e i r ends.  to  as  On  being  The branches b e a r i n g Resinosus  may  result  from b a s a l bole cankers or sunken areas on the lower part of the tree  (Whitney The  1962).  estimation  d i s e a s e d p l o t s has Groenewoud  of  disease  only  (1964)  been  and  impact  by  remeasurement  attempted  by  Whitney  and  van  Whitney  and  van  Lachance  (1978).  Groenewoud (1964) returned to two n a t u r a l stands of white a f t e r a ten year p e r i o d . years  old.  offset  the volume l o s s e s a t t r i b u t e d to m o r t a l i t y .  the  case  of  In  The stands were i n i t i a l l y  by  16%.  in  The  in t r e e s examined v a r i e d and  decrease  numbers  average  by  and  However,  and  dead  to 15.2  3.8  cm.  5-15  I t was years  for  in 24%,  trees  The  rate  cm per year.  van Groenwoud (1964) a l s o examined the  until  infected  51  annual rate of growth of P.  shown that  the  radial  increment  did  p r i o r to death of the t r e e s .  Based on f u n g a l growth r a t e s , however, the been  30%,  i n f e c t e d r o o t s was  from 0.2  increment of dead t r e e s . not  38  the o l d e r stand, b a s a l area had decreased by  tomentosus i n a r t i f i c i a l l y  Whitney  spruce  the younger stand, the growth of h e a l t h y t r e e s  healthy t r e e s decreased increased  of  trees  had  20-30 years before the t r e e s death  to  have  (Whitney  1962). In  a study of p l a n t a t i o n spruce i n Quebec, Lachance  r e p o r t e d a 27.3%  (1978)  ten year volume l o s s where Polyporus tomentosus  10  was and  r e s p o n s i b l e f o r two-thirds of the l o s s e s , both  in frequency  volume. Tunneling by weevil l a r v a e i n the roots of examined  has  been  a s s o c i a t e d with the i n c i d e n c e of root s t a i n and decay  (Whitney 1961). warreni  The  Wood.  insect  Whitney  has  been  weevil)  but  is  identified  as  Hylobius  (1961) s t a t e d t h a t , "entry of fungi does  not always appear to be d i r e c t l y the  spruce  often  through  through  the  wound  (caused  by  a more d i s t a l p a r t of the  root. " Warren's root c o l l a r banksiana been  observed  The  1973).  Severe  soils  Abies  flightless  old.  balsamea  adult  damage occurs i n  The  most  however  and c l i m a t e may  requirements (Warren  and  of  1956).  spruce.  trees  The  cambium  can l i v e  certain  damaging  (L) M i l l . .  to  which  local  are  Pinus  I t has  never  larval  stage  of  the  f o r four years  g r e a t e s t weevil a c t i v i t y  conditions,  composition the  on  years feeding on bark  tree.  duff  is  Lamb., lodgepole p i n e , and white  l a s t s f o r two  years  weevil  host  (Cerezke  about  forty  occurs i n moist to wet factors  of  stand  produce moist c o n d i t i o n s s u i t a b l e to  these i n s e c t s , even on sandy w e l l d r a i n e d  11  III.  METHODS  I n v e s t i g a t i o n of the d i s e a s e followed two approaches. first the in  was  to observe  host  the a c t i o n of the fungus and the e f f e c t s on  in some d e t a i l .  order  to  development and y i e l d . selected  Areas of severe disease were s e l e c t e d  efficiently and  The spruce  The  study  pathogen  identity,  disease  spread, and the e f f e c t s of i n f e c t i o n on growth second approach stands.  involved a  This  survey  survey  of  provided  i n c i d e n c e i n stands which were s e l e c t e d on the  randomly  a measure of  basis  of  cover  type. 1.  SELECTION OF EXPERIMENTAL AREAS Several  objectives  spruce stands.  The  were  met  in s e l e c t i n g representative  stands had to be immature (50-70 years o l d ) ,  even-aged, predominatly  spruce, and  fully  o b s e r v a t i o n s of the d i s e a s e i n i t s e a r l y  stocked.  T h i s enabled  stages.  Such  are a l s o r e p r e s e n t a t i v e of c u r r e n t restocked spruce Major district  amounts  of  white  spruce  of the P r i n c e George F o r e s t Region.  were a l l found w i t h i n the Sub-Boreal In  this  study,  all  investigations  c o n t a i n e d i n the SBS J/01, the SBS  J/06,  Three  in  Spruce  forests.  in the e a s t e r n  Candidate  stands  " J " subzone (SBS J ) .  took  the Mesic Oak  place  Fern  w i t h i n areas  association,  and  the Submesic Queen's Cup s u b a s s o c i a t i o n .  stands were s e l e c t e d as matching  one at the Bowron R i v e r , one one  are found  stands  i n the J e r r y  the Hixon F o r e s t D i s t r i c t  J e r r y Creek stand was  the above c r i t e r i a ,  Creek  ( F i g u r e 1).  a i d e d by c o n s u l t i n g  fire  drainage,  and  S e l e c t i o n of the histories  from  12  the  past  fast  r e g e n e r a t i o n of spruce  "Nukto Laie  seventy  years.  A l l three stands  showed evidence of  following f i r e .  "  \Swemp )L. \  Figure  1 - L o c a t i o n of the study p l o t s ( • ) (Table 3) and the i n c i d e n c e survey stands ( ° ) (Table 5 ).  13  2.  PLOT  SELECTION  The  o r i g i n a l  encompassing s u r r o u n d i n g because  a  s i n g l e  h e a l t h y  t h e  c e n t e r s .  Instead tree  tree  severe  0.01 as  w i t h  ha  (about  d e s c r i b e d  p l o t s  7-9  F i g u r e  30  2  -  A  p l o t recent  symptoms  Each  p l o t  trees)  d i s e a s e  was  was then  l o c a t e d  a t  blown  down  1-6 J e r r y  were  a t  to  a t  and p l o t  t h e  c e n t e r  o b v i o u s l y  ( f i g u r e  2 ) ,  caused  encompass  p l o t the t e n  of  t o  r e c o g n i z a b l e  opening  t h e  each  p o s s i b l e  an  windthrow  l o c a t e d  Creek,  spruce  on  extended on  be  d i s t i n c t  stand  and a l l t r e e s  P l o t s  t o  c e n t e r e d  a  p l o t s ,  and extending  not  i n  i n f e c t e d or  t e n  c e n t e r  proved  o r g a n i z e d  crown  below.  were  a  e s t a b l i s h  This  was not  as  t o  d i s t i n c t  each  such  d i s e a s e .  was  t r e e s .  d i s e a s e  d i s e a s e d  root  i n t e n t  were  p l o t  by  about  examined  Bowron a t  a  R i v e r ,  Hixon.  B4.  14  Table  3 shows the area, s t o c k i n g , and volume of each of the  ten p l o t s . equations  The volume was based on M i n i s t r y for interior  Table  spruce  of  (B.C. M.O.F.  Forest  1976).  3 - Average age, area, s t o c k i n g , and volume of spruce found i n the p l o t study  PLOT CODE  Age  Area  (sq. m)  Stocking  (stems  per ha.)  3.  Volume (cub m per ha.)  B1  52  11 0  1610  240  B2  57  1 60  1820  238  B3  56  180  1 500  260  B4  53  290  1315  260  DC 1  62  80  2500  368  DC 2  58  1 20  1605  260  DC 3  54  240  1595  200  HIXON  56  250  1 1 60  1 70  J1  55  1 30  760  1 45  J2  64  280  640  1 06  EXPERIMENTAL PROCEDURES The  l o c a t i o n of a l l standing t r e e s , l i v i n g or dead, w i t h i n  the p l o t was mapped, using a were  volume  also  mapped.  place  classification  For each t r e e the  table.  Wind-thrown  trees  A v e g e t a t i v e d e s c r i p t i o n was prepared  s o i l p i t dug i n order to ecological  plane  following  the  three  scheme developed were  recorded:  stands  within  and a the  by K r a j i n a (1969). species;  height;  15  diameter  at  breast  height  (1.3 m); age as determined from an  increment core at 0.3m; and crown c l a s s . The  roots of each t r e e on the p l o t were exposed by hand  examine how the fungus i n f e c t e d and spread through host Root  exposure a l s o allowed  g r a f t s with other noted a  trees.  or  white  Fungal  presence  in  tree  xylem  was  followed i n l a t e r stages of decay by  pocket  rot.  A  probe  was made using an  increment borer and an axe to look f o r these Three  tissue.  i n v e s t i g a t i o n of p o s s i b l e contact or  by a red-brown s t a i n ,  yellow  to  characteristics.  l o c a t i o n s on each t r e e were examined; the exposed roots a  meter from the root c o l l a r , the root c o l l a r , and breast  height.  A three number s c a l e r e l a t i n g the presence of decay symptoms was made f o r each p o i n t examined on the t r e e ; one two  for stain,  and  three  for rot.  f o r no  disease,  T h i s r e s u l t e d i n a three  number code f o r each tree g i v i n g a r e l a t i v e measure of d i s e a s e . T h i s code  accompanied  observations  made  on  any  other  tree  symptoms. Each  standing  d i s s e c t e d to look ten  plots  at  Isolations and  and  f o r Polyporus  gathered  in  were  tree  tomentosus.  order  C u l t u r e s were a l s o taken  was thoroughly  From  to  each  culture  from unstained  made using a s t e r i l e technique  tetracycline petri plates.  fruiting  bodies  pathogens  present.  An  windthrown  of the  l e a s t ten samples of s t a i n e d and decayed root or  bole m a t e r i a l was organism.  dead  Later  i n the  the  causal  wood samples.  on 2% malt agar summer,  fungal  were c o l l e c t e d t o a i d i n i d e n t i f i c a t i o n of the  increment core  was  taken  from  each  tree  at  breast  16  height.  The  core  was accepted only  Each core was s t o r e d measurements. years,  in a plastic  Radial  straw and r e t a i n e d  counter.  for  later  increment was measured i n groups of f i v e  s t a r t i n g from the. 1982 latewood  yet complete) back  i f i t i n c l u d e d the p i t h .  to  the  pith  (1983 r i n g growth was not  using  an  Wood moisture was standardized  Addo-x  tree  ring  by soaking the core i n  water f o r an hour p r i o r to measurement. Covariate differences classes. 4.  analysis  in  radial  was  used  to  test  significant  increment between the d i s e a s e  Diameter squared times height  intensity  was the c o v a r i a t e .  INCIDENCE SURVEY A survey of the incidence  six  immature  spruce  of root disease  stands.  forest  cover  which  survey  was  to  do  the  maps.  Accepting  two),  intermixed rejected tissue.  fully  with since  stocked,  distinct  were  met.  stand. then  A  clumps  of  hardwood  The  was v i s i t e d to see  basic  (1980)  baseline  Two g r i d s of three  of  survey  design  for Phellinus 200m  The  Stands  species  were  i f the  by  taking  above  followed weiri i  r e g u l a r l y spaced t r a n s e c t  increment  lines  each were  falling  The a c t u a l i n f e c t e d core borings  that  (Murr.)  was e s t a b l i s h e d w i t h i n  i n f e c t e d areas was determined.  identified  in  o l d or age  e s t a b l i s h e d , and the t o t a l length of t r a n s e c t l i n e  within  in  t h i s would i n d i c a t e a d i s r u p t i o n of s u s c e p t i b l e  developed by Blooomberg Gilbertson.  year  and predominantly spruce.  Each candidate stand  criteria  stands  based on s e v e r a l c r i t e r i a .  stands had to be r e l a t i v e l y even-aged (50-70 class  was conducted  Candidate stands were l o c a t e d on  the F o r e s t D i s t r i c t  was  for  area  of roots of  17  t r e e s s i t u a t e d w i t h i n a given d i s t a n c e of distance transect in  varied line.  from  0.5  that stands with g r e a t e r than  hectare  1.5  meters  transect.  density  1500 stems per hectare had one  stands with  1000 to  1500  stems  per  had two meter wide t r a n s e c t l i n e s , and stands with  less  1000 stems per hectare had three meter wide t r a n s e c t l i n e s .  Presence  of red s t a i n or decay i n the increment  to i n d i c a t e the presence verified  by  the  of Polyporus  culturing  core  tomentosus.  was  was  taken  T h i s was a l s o  of ten t r e e s per survey area.  boundary between healthy and d i s e a s e d areas along line  This  e i t h e r side of the  T h i s d i s t a n c e changed r e l a t i v e to stand  meter wide t r a n s e c t l i n e s ,  than  to  the  the  The  transect  d e f i n e d as the p o i n t midway between the p r o j e c t i o n of  adjacent h e a l t h y and d i s e a s e d t r e e s .  18  IV. 1.  RESULTS AND  DISCUSSION  IDENTIFICATION OF PATHOGENS A t o t a l of 190 attempts to i s o l a t e root  were  made,  plots.  including  One  tomentosus but ten  hundred  pathogens  at  least  ten from each of the research  and  forty  of  cultures.  yielded  disease  these  y i e l d e d . Polyporus  The remainder were l a r g e l y contaminated,  Pleurocolla  compressa  (E&E)  Diehl,  a  pathogenic fungus known to i n h a b i t wood a f t e r other decay Isolation  attempts  were  decay and advanced decay. root pathogen (Nobles  isolated.  1948),  Dr. Whitney.  of In  1  made  from both red s t a i n e d  Polyporus  tomentosus  was  produced  on  cultural addition,  characteristics t y p i c a l P.  The  disturbed  identity  confirmed as being P. tomentosus was t h i s study.  1  2  incipient the  only  and confirmed by  tomentosus sporophores of  them  and exposed i n f e c t e d r o o t s .  Such  sporophores were commonly s t i p i t a t e 3 and 4 ) .  fungi.  I d e n t i f i c a t i o n was based on Nobles' key  were observed on s e v e r a l of the p l o t s i n August. were  non-  of  these  Many  rather than shelved fruiting  tomentosus by Dr. J .  the only root p a r a s i t i z i n g  bodies Hopkins.  fungus  (Figures was  2  also  Polyporus  found  during  A l l the f r u i t i n g bodies had s t r a i g h t setae only.  Dr. R.D. Whitney of the Canadian F o r e s t r y S e r v i c e , Sault Ste. Marie, O n t a r i o , v i s i t e d the study s i t e s during the summer of 1983. Dr. Hopkins of the P a c i f i c F o r e s t Research Center, V i c t o r i a , i d e n t i f i e d a f r u i t i n g body c o l l e c t e d on s i t e .  19  Polyporus tomentosus was wood.  Initially,  red s t a i n .  invasion  Shortly thereafter, typical  turns  other  is  the  butt  common u n t i l a l a t e r age. restricted  to  conditions  immature spruce.  other  not  remains  Polyporus  rot fungi of spruce may  Alternatively,  ecological  the  It i s concluded that  dominant root pathogen of  known root and  light  small pockets appear,  with white m a t e r i a l , a mixture of hyphae and  tomentosus  normal  tomentosus produces a l i g h t  the a f f e c t e d wood  of the decayed wood (Figure 5).  The  i s o l a t e d from unstained  by P.  As decay progresses,  yellow-brown. filled  not  not become  fungi  may  be  encountered in t h i s  study. 2.  CROWN SYMPTOM DEVELOPMENT Diseased t r e e s d i d not  show c o n s i s t e n t crown symptoms  c o u l d be r e l a t e d to the extent Some t r e e s were apparently these  were  understory.  smaller The  k i l l e d by P.  than  actual  of i n f e c t i o n i n r o o t s and  the cause  tree  death  trees  with  crowns  i n d i v i d u a l s showed t y p i c a l height  growth,  However others  chlorosis,  in  the  such and  than  cases  root  symptoms  decrease in needle  On  was  disease.  of  diseased reduced  longevity.  with e q u a l l y severe or worse d e t e r i o r a t i o n of  root system e x h i b i t e d no obvious crown symptoms. diseased  found in the  main canopy, a few  root rot crown and  and  in  probably a combination of n a t u r a l suppression Among  boles.  tomentosus, but most of  average of  that  the  the  whole,  t r e e s with obvious crown symptoms were much l e s s common  diseased  t r e e s with normal crowns.  to q u a n t i f y these o b s e r v a t i o n s ,  No  attempts were made  p a r t l y because of the  of r a t i n g crown,symptoms o b j e c t i v e l y .  difficulty  20  F i g u r e 3 - F r u i t i n g s t r u c t u r e emerging f r o m a 4 cm d i a m e t e r stump  F i g u r e  4  -  of of  F r u i t i n g s t r u c t u r e of the base of a white  Polyporus tomentosus a d e s t r o y e d white s p r u c e .  Polyporus s p r u c e .  tomentosus  at  21  Figure t y p i c a l white  It p e r i o d while  5 - Exposed roots pocket appearance  seems  that  d u r i n g the  the  rate  d i e ,  i s  which  crown of  low  most  i . e .  windthrow,  often  in  the  root  Trees system  to  the  were  are  any  a l l  a c t i v e  Note the at the c e n t e r .  through in  the It  t r e e s  number  of  i n f e c t e d  w i t h major  symptoms e x t e n s i v e ,  system  standing  which  from  root  long  that  at  removed  a  f o l l o w s  rate  crown  observed  a f f e c t i n g  i s  pass  u n a f f e c t e d .  the  t o t a l  t r e e s  before  t r e e s  pathogen  m o r t a l i t y ,  i n f e c t e d  6).  the  r e l a t i v e l y  compared  white s p r u c e . a decayed root  i n f e c t e d  remains  a d d i t i o n ,  (Figure  of of  the have  t r e e s . stand  by  developed  advanced  s t r u c t u r a l  In  r o o t s ,  decay but  22  without  apparent  candidates It  crown  symptoms.  of  the extent  symptoms  of d i s e a s e  of the  incidence of P.  major  roots f o r the s t a i n and  On  each  completely infection  are  prime  the  excavated  ten in  spread.  Initial  infection  healthy  root  Hence  examination  systems  of  by P.  in  a  the the  conservative  tomentosus  order The  (<2  via  several  to  determine  systems were  the  manner  of  following  pathway  i s proposed.  contact  between  infected  ( F i g u r e 7). cm  root  In most cases  and  such c o n t a c t s  diameter).  the pathogen grows e c t o t r o p h i c a l l y on the  of small r o o t s .  surveys  At best, d e t e c t i o n of result  plots,  occurs  i n v o l v e small r o o t s Initially  unreliable  SPREAD  of  and  an  i n a stand.  decay.  measure of the a c t u a l area occupied INFECTION AND  give  tomentosus r e q u i r e  d i s e a s e by crown symptomology would  3.  trees  f o r windthrow.  i s a l s o c l e a r that crown  indication  Such  Penetration  of bark  and  cambium  surface  occurs  soon  a f t e r , and as the mycelium on the root s u r f a c e grows p r o x i m a l l y , it  continually  wood.  The  initiates  new  points  s i t e s of p e n e t r a t i o n  show as  of i n v a s i o n of bark small  patches  diameter) of dead brown bark.  These patches grow and  until  killed.  the root i s g i r d l e d and  p r o x i m a l l y the root i n c r e a s e s  in  mycelium  longer  can  apparently  Instead  i t advances in the  center  of  no  xylem,  f r e e of d i s e a s e .  cm  amalgamate  As the mycelium advances  size.  At  some  penetrate  typically  the r o o t , while the outer  (<1  and  point,  healthy  at  or  the  bark.  near  the  s e v e r a l xylem r i n g s remain  At the p o i n t at which t h i s switch  occurs  the  23  F i g u r e s t r u c t u r a l  6 - A windthrown white s p r u c e due to complete and f u n c t i o n a l f a i l u r e of the r o o t s . R e d - s t a i n appears in the f i r s t notch in the l o g .  root  i s  about  which  the  primary  It  i s  not  e c t o t r o p h i c tree  in  small the  c l e a r  absence l i v i n g  cm  diameter)  root but  c o l l a r . such  in  diameter.  periderm  from (1  a r e a s ,  cm  growth  the  energy  5  i s  r e p l a c e d  whether on of host  the  l a r g e r the  r o o t s  t i s s u e s .  Polyporus do  not  may by  a l s o  a  and to  There  patches  appear  can  to  the  the  penetrate are  was  point  at  p e r i d e r m .  a l s o  along  into  tomentosus  be  secondary  pathogen  a b i l i t y  n e c r o t i c  patches  T h i s  maintain  base  of  and  the  d e r i v e  rare  o c c u r r e n c e s  the  cambium,  i s o l a t e d  spread  or  from  e n l a r g e .  of  near such  24  F i g u r e  The  7  -  host  a d v e n t i t i o u s diseased  by  continues  to  increment  energy It through P.  commonly roots  r o o t s .  invasion  because  Root g r a f t i n g of from Polyporus  produce i s  not  s o i l ,  tomentosus  t r e e new  them.  i s  and It  t r e e s , forced  roots  as to  rather  whether  i f  so,  how  u s i n g  the  Warcup  f a r .  be  than the  a in  the  nor  l a r g e bole  but  the  below, part  to host  r e d u c t i o n  can  (1962)  screened  l a r g e  i n  a r i s e s of  i t s  wood.  pathogen  Whitney  of  s u s c e p t i b l e  demonstrated spend  p r o d u c i n g  end  k i l l e d ,  that  s t a i n  by  are  r a p i d l y  may  red  s u r v i v i n g  roots  are  and  a t t a c k  d i s t a l  known  and  to  a d v e n t i t i o u s  d i s e a s e d  host  the  p a r a s i t e  r e p l a c e  of  the to  the  responds  from  Such  spruce r o o t s , tomentosus .  a l s o  c o u l d  not  immersion  spread b a i t p l a t e  25  techniques.  However,  the absence of decay f l o r a may  the l i m i t a t i o n s of the the  plots,  observed.  a  r i n g of P.  T h i s c o u l d only r e s u l t  l i v i n g hyphae. this  fairy  i s o l a t i n g technique i t s e l f .  case,  The  one  of  tomentosus f r u i t i n g bodies  was  from  still  horizon)  which i s r e s t r i c t e d s o i l and  spread.  I t may  acquire  to  ectotrophic  totally  organic  h o r i z i o n rather than in quickly  energy expenditure  on  a  dense mat  produced by the  above,  it  growth  on  roots  the  and  the b o l e .  the  xylem  of  the p o i n t  uninfected  soil.  These  a major root  emergence  pustules erupting Every  then a  fair  in  the  fashion  developing  i n t o the  fungus was  base  observed  level.  diameter  the  by P.  has  t r e e s , and  decreased  to  into  when i t a  few  i n i t i a t e s e c t o t r o p h i c growth. fungus  from under the root  t r e e attacked  roots  to grow, e v e n t u a l l y  r o o t s of d i s e a s e d  i t reemerges and  of  the  fungus i s able to grow i n t e r n a l l y  reaches a p o i n t where root centimeters,  for  of f i n e spruce roots in the  s t a i n caused by the  up to f i v e meters above ground It appears that the  ring,  consumed, r e p r e s e n t i n g  entered  continues  Decay and  the  of the host  i n t o a column of decay that extends from the root of  in  host.  Once the p a r a s i t e has described  such as P h e l l i n u s w e i r i i  in the case of the f a i r y  is  be  of  roots at random  r e l i e s on root contact  fungus  would  ring  energy to form f r u i t i n g s t r u c t u r e s .  a l s o be that  living  continuous  able to a t t a c k  T h i s i s in c o n t r a s t with a root d i s e a s e  mineral  a  On  to  fungus c o u l d be growing through the s o i l ( i n  the organic  l o c a t i o n s , and  be due  appears  as  small  At  white  bark.  tomentosus was  a l s o a f f e c t e d by  26  the root c o l l a r w e e v i l , (Hylobius warreni)  (Figure 8 ) .  Attacks  on the r o o t s and the c o l l a r of the spruce host were a l l c u r r e n t , with  signs  of previous i n f e s t a t i o n s .  on healthy t r e e s , but at a lower  The weevil a l s o occurred  frequency.  healthy spruce t r e e s had weevils p r e s e n t . taking  the  activity,  opportunity  Twenty of  the  125  E i t h e r the fungus was  to a t t a c k s t r e s s e d roots due to weevil  or the weevil sensed  s t r e s s e d roots due t o the  fungal  disease. Weevil In  almost  g a l l e r i e s were l a r g e l y a l l cases,  the  g a l l e r i e s remained a l i v e . the  case,  there  associated  cambium  directly  underneath  such  Even in i n s t a n c e s where t h i s was  not  with  tomentosus decay and any  the weevil wound.  the root c o l l a r weevil i s a secondary weaken  the  host,  s i t e of i n f e c t i o n . attack  I t i s concluded  agent  that  may  The spruce host normally  responds to  by the production of l a r g e amounts of r e s i n , p r o t e c t e d by  and  buried  masses of r e s i n soaked root bark and s o i l . (1941),  stain  resin.  i n severe  further  s i t e to the requirements  weevil  within  large  However, as noted by  i n f e c t i o n s , a t t a c k through  T h i s may be s u f f i c i e n t  as an a t t r a c t a n t .  that  so that the  the cambial  c e l l s of the roots by the pathogen a l s o causes a c o p i u s of  in  but that weevil g a l l e r i e s are not the normal  g a l l e r i e s were normally  Haddow  bark.  were s e v e r a l r i n g s of u n i n f e c t e d xylem t i s s u e  s e p a r a t i n g the column of P. xylem  r e s t r i c t e d to the outer  release  i n a l t e r i n g an otherwise dry  of the w e e v i l .  The r e s i n may a l s o  act  27  4.  SUSCEPTIBILITY  Table in  the  4  ten  i n s t a n c e s c o n t a c t  OF  shows p l o t s .  stems  with  CONIFER  the  amount  Only  and  i n f e c t e d  SPECIES  roots spruce  of  i n f e c t i o n  spruce of  was  the  but  s p e c i e s  i n f e c t e d .  other  r o o t s ,  by  s p e c i e s  no  observed  In were  s i g n s  s e v e r a l i n  of  d i r e c t  i n f e c t i o n  Figure 8 - Exposed root c o l l a r of white s p r u c e . Stem occurs at l e f t , an adult w e e v i l i s at the center of the r o o t , and a pupal chamber l i e s at bottom l e f t (arrows).  were  observed.  p l o t s  have  Columbia of  a  been  (Lowe  t e s t  s u s c e p t i b l e  of  A l l  the  reported 1969).  as  hosts  Whitney  s e e d l i n g s ,  s p e c i e s .  c o n i f e r  s p e c i e s  of  P.  encountered  tomentosus  and  Bohaychuk  (1976)  l i s t e d  lodgepole  pine  i n on  among  i n  these  B r i t i s h the the  b a s i s most  28  The 170  proportion  of i n f e c t e d c o n i f e r s on the  d i v i d e d by 346,  or 0.491.  The  ten  plots  p r o b a b i l i t y that the  was  51  non-  spruce l i v i n g c o n i f e r s escaped i n f e c t i o n by chance would be 51 -15 P = (1 - 0.491) = 1.103 escape The  h y p o t h e s i s that a l l c o n i f e r s were e q u a l l y  be  rejected.  and  as a r e s u l t i t may  infected  D i f f e r e n t s p e c i e s have d i f f e r e n t r o o t i n g be that root c o n t a c t s  l e s s common than w i t h i n  species.  initially  on  established  spruce i n f e c t i o n would be g r e a t e r  the  habits,  between s p e c i e s  Hence.if P.  spruce,  can  tomentosus  p r o b a b i l i t y of  than i n f e c t i o n  of  are were  further  the  other  species. If  the  likelihood  were very much smaller the is  of  t r a n s f e r from spruce to non-spruce  than that  from spruce to spruce,  non-spruce c o n i f e r s were w e l l separated conceivable  infection. spread  It  from  proportion  that can  spruce of  the  51  non-spruce  be c a l c u l a t e d how to  non-spruce  would  i n f e c t e d non-spruce t r e e s  conifers  would be no have to be  at  smaller  random, than 0.05.  and  if  from each other, i t conifers  escaped  small the p r o b a b i l i t y of be.  The  in the p o p u l a t i o n  would  have to be so small that the p r o b a b i l i t y of spruce  and  have  to  selecting  51  non-  f i n d i n g them a l l u n i n f e c t e d ,  Hence the p r o p o r t i o n  (P)  would  such that 51  (1-P)  <  0.05  P < 0.057 This  compares  with  the  proportion  of spruce i n f e c t e d of  d i v i d e d by 295,  or 0.58.  If the p a u c i t y of root  the  lack  infection  cause  of  of  contacts  170 were  of non-spruce c o n i f e r s , the  29  frequency of such c o n t a c t s would have to be l e s s than average that  number between spruce t r e e s .  lack  of  root  contact  i n f e c t i o n of non-spruce Table  is  10% of the  It i s therefore  responsible  unlikely  f o r the lack of  conifers.  4 - Incidence of tomentosus root disease f o r a l l l i v i n g t r e e s p e c i e s examined.  Number of Trees Healthy  Spec i e s  Abies  lasiocarpa  Infected  28  0  18  0  Pinus c o n t o r t a  5  0  Populus  2  0  Pseudotsuga  menziesii  tremuloides  1 25  P i c e a qlauca  It may be of course that spruce  are  also  much  1 70  in addition,  less  susceptible  species than  other  spruce.  than This  e x p l a n a t i o n runs counter to the r e s u l t s of Whitney and Bohaychuk (1976), and a l s o to o b s e r v a t i o n s of heavy i n f e c t i o n on lodgepole pine i n the Quesnel Wilford.  3  as  reported  by  Morrison  and  3  Another of  Lakes areas  several  e x p l a n a t i o n might be that P. r a c e s , each adapted  tomentosus  to a p a r t i c u l a r  consists  t r e e s p e c i e s or  Dr. D.J. Morrison, P a c i f i c F o r e s t Research Center, and E. W i l f o r d , B.C. M.O.F., i n a communication with Dr. B.J. van der Kamp, U.B.C..  30  group of s p e c i e s .  T h i s phenomenon has  been observed  root d i s e a s e s such as A r m i l i a r i a mellea, (Korhonen,  1978).  "  and  for  Fomes  other annosus  If t h i s were the case, damage caused by  tomentosus on a s i t e may  be  avoided  by  a  switch  of  P.  conifer  spec i e s . Appendix  B  shows  stem  maps  indicating  the p o s i t i o n of  h e a l t h y , d i s e a s e d and dead t r e e s in the ten p l o t s by s p e c i e s . None of the ten p l o t s showed d i s t i n c t disease c e n t e r s evidence  of  progression  p e r i p h e r y to the various  stages  of  center. of  symptom  Instead  disease  A  set  of  (1962) i n d i c a t e s that P.  tree,  it  seems  faster.  The  contact  other  that  Given  experiments  the  exhibiting  it  was  of disease c e n t e r s . conducted  tomentosus grows at 3.8  by Whitney  cm per  year.  be t h i s slow, but once e s t a b l i s h e d in the growth  through  secondary xylem i s much  fungus could t r a v e l through trees  trees  this situation,  the rate of spread  inoculation  E c t o t r o p h i c growth may  diseased  from  development and healthy t r e e s were  randomly mixed w i t h i n the p l o t s . not p o s s i b l e to estimate  development  with  at  a  greater  the  root  rate than  system  and  ectotrophically  growing f u n g i . It should not be tomentosus stands. free  is  truly  concluded random  that  among  infection trees  in  by  immature  Rather, s u b s t a n t i a l s e c t i o n s of a stand may  of  disease.  However,  within  Polyporus  be  spruce totally  the d i s e a s e d p a r t s of  the  stand, healthy and diseased t r e e s appear to be randomly mixed.  " Dr. 1983.  D.J.  Morrison  in a communication with the author,  March,  31  5.  IMPACT OF TOMENTOSUS ROOT ROT  5.1  Survey The  Table  r e s u l t s of the survey f o r i n c i d e n c e  5.  Three  of  are  the s i x stands surveyed were a l s o used to  e s t a b l i s h the ten r e s e a r c h p l o t s d e s c r i b e d above. survey Figure  lines  reported in  d i d not  actually  However,  the  t r a n s e c t any of these p l o t s .  1 shows the l o c a t i o n of the s i x stands.  If  we  assume  that each of the s i x stands surveyed r e p r e s e n t s an equal area of immature  spruce f o r e s t ,  (essentially classifying  the p o p u l a t i o n  i n t o s i x equal s t r a t a ) , the o v e r a l l area i n f e c t e d  i s 28.4 + 13.7  percent. Table 5 - Incidence of Polyporus tomentosus root r o t i n white spruce w i t h i n the P r i n c e George survey area ( F i g u r e 1).  Location  Transect  Number  Width  Length  of Trees  (m)  (m)  examined  Percent of Area Infected  95% conf. . Interval  Highway 16  1  390  1 43  J e r r y Creek  2  360  128  16.8  8.2  Tabor Mtn.  2  336  1 53 .  17.6  8.6  Hixon  2  360  1 45  37.7  16.3  A l e z a Lake  3  300  1 13  42.2  10.4  Beaver Road  1  417  1 26  49.0  24. 1  6.9  6.1  32  5.2  E s t i m a t i o n Of The  total P.  Losses  survey d e s c r i b e d above estimates  stand area now  of  w i l l be due  a  percent  occupied by d i s e a s e d t r e e s .  tomentosus i n these stands,  consist  the  decrease  to a decrease  from logs with butt r o t .  will  in  the  The  final  of  the  impact  of  analysis,  i n value at the time of h a r v e s t .  This  i n volume, and a lumber recovery  loss  In t h i s study the e s t i m a t i o n of  impact  i s l i m i t e d to a c o n s i d e r a t i o n of r e d u c t i o n of volume. Volume l o s s e s occur  in at l e a s t  four ways, namely  increment  l o s s , butt r o t , m o r t a l i t y of standing timber and windthrow.  The  data  and  c o l l e c t e d allow an e s t i m a t i o n of the c u r r e n t increment  butt rot l o s s e s , but cannot up  to  r o t a t i o n age.  more severe  Whitney  i n o l d e r stands.  contained more decay. trees may  was  expect  certain 5.2.1  project  equal  l o s s e s due  to  the  disease,  (i960) noted that the d i s e a s e Older  infected  trees  themselves  However, the r a t i o of h e a l t h y to d i s e a s e d  i n a l l age c l a s s e s from 40 to 120 y e a r s .  then, that younger stands  may  be  able  to  One  mask  Butt  Rot extent  of P.  for each i n f e c t e d t r e e . Forty-five  percent  tomentosus s t a i n and decay was The  r e s u l t s are presented  measured  i n Table  of the d i s e a s e d t r e e s showed P.  6.  tomentosus  s t a i n and or decay at breast h e i g h t ; i n 34 percent the s t a i n had  progressed  height, while stain  a  l e v e l of i n f e c t i o n b e t t e r than do o l d e r stands.  The  decay  was  and  the  i n the remaining  decay was  of a l l l i v i n g  to  root 21  collar  percent  r e s t r i c t e d to r o o t s .  spruce t r e e s encountered  or  but not to breast of  infected  Expressed  i n the survey  trees  as percent (using  the  33  figure  of  28.4  as the percent  of spruce  trees infected)  values become 12.9, 9.7, and 5.8 r e s p e c t i v e l y . that  i n the stands  described  these  One may conclude  i n t h i s study at l e a s t  22.6 percent  of the t r e e s show butt r o t now.  One would expect t h i s f i g u r e t o  i n c r e a s e , on a stand b a s i s ,  rotation  healthy  to  expressed  diseased  by  trees  in  i n greater p r o p o r t i o n s  in which the pathogen  is  now  the  age.  stand  ratio  i s unchanged  i n butt r o t . restricted  The  to  but i s  Many of the t r e e s the  roots  will  develop butt r o t by r o t a t i o n age. Table  Age C l a s s  6 - Number of white spruce by age c l a s s i n a l l p l o t s , showing v a r i o u s d i s e a s e symptoms.  Healthy  E x h i b i t i n g s t a i n or decay by Polyporus in  roots  only  tomentosus  at the root  at :  collar  30-35  2  -  -  -  36-40  2  -  -  -  41-45  2  3  4  7  46-50  10  4  10  8  51-55  38  12  17  22  56-60  38  13  19  25  61-65  23  3  7  13  >65  10  -  1  2  125  35  58  77  TOTALS  of  34  It on  i s noteworthy that the extent of decay was  tree  age.  not dependent  Table 6 shows a s i m i l a r age d i s t r i b u t i o n  for a l l  three d i s e a s e c l a s s e s . The measurements necessary decay were not c o l l e c t e d .  to estimate the a c t u a l volume of  At present t h i s volume would probably  be l e s s than one percent of the t o t a l volume figure  would  increase  r o t a t i o n age, however,  spruce.  This  with time, but even i f i t quadrupled  i t would s t i l l  that  of  be s m a l l .  the major l o s s here  It  should  by  be  realized  i s a s s o c i a t e d with  decreased  lumber recovery from logs with butt r o t . Current  tables  (B.C. M.O.F. 1976)  for  decay,  estimate  the  waste  class,  percent  decay  of  for diameter  such  be  due  to  the  trees  type  For the 121  is  c l a s s e s 10 to 50  f i g u r e s are somewhat lower  breakage  decay l o s s i n immature  not bearing i n d i c a t o r s of decay as 0%. age  and  to 250  estimated at 1.8  cm  spruce year to  respectively.  than our r e s u l t s .  3.1  These  The d i f f e r e n c e  may  of spruce stand s e l e c t e d f o r the c u r r e n t  study. 5.2.2  Increment An  Loss  increment  core taken at breast height was  each t r e e i n the ten p l o t s . five  year  periods  from  R a d i a l increment the  measurements were transformed the  radius  diameter. years  measured The average  1977-1982  was  by  the  1982  ring  was  to  i n t o b a s a l area increment  f i v e year b a s a l area  collected  from  measured  in  the p i t h .  These  increment,  using  core to estimate t r e e increment  for  the  1708.2 square m i l l i m e t e r s f o r healthy and  1087.3 square m i l l i m e t e r s f o r d i s e a s e d  trees.  The  difference  35  was  not s t a t i s t i c a l l y  s i g n i f i c a n t , being masked by v a r i a t i o n not  a t t r i b u t a b l e to d i s e a s e . size.  Large  trees  One major source  grow f a s t e r than  of v a r i a t i o n was tree  small t r e e s , and both the  healthy and d i s e a s e d t r e e c l a s s e s e x h i b i t e d a wide range of t r e e sizes  (Table  diameter  at  covariate. basal  11). A covariance  increment  squared  and  a  useful  the  logarithmic  between form.  i n Table  these  7.  the diameter squared respectively.  f o r the  The equations  the l o g a r i t h m of b a s a l area  According  to  i n both regression  the  common  and  and  lines  are  logarithm  of  slope,  are shown i n Table 8. fixed.  using  Bjerring  1980),  but  a common y-  The a n a l y s i s of c o v a r i a n c e was then  the s t a t u s was c o n s i d e r e d  natural  the a n a l y s i s of c o v a r i a n c e ,  intercept.  and  between  f o r healthy and d i s e a s e d t r e e s  lines  results  height as the  Appendix C shows the  regression  the  a  using  g i v e the r e l a t i o n s h i p between  the computer package " G e n l i n " (Greig had  the  relationship  increment versus  times height  conducted  c o v a r i a t e , i t was necessary to  variables  The equations  was  times  the data to n a t u r a l l o g a r i t h m s .  relationship  given  height  In order to obtain  area  transform  breast  analysis  not  performed,  P l o t was c o n s i d e r e d  the  and  random,  36  Table 7 - Equations of the r e g r e s s i o n l i n e s f o r the l o g a r i t h m of b a s a l area versus the logarithm of diameter squared times height and r e s u l t i n g t e s t f o r common s l o p e .  Tree s t a t u s  Regression  equations  Healthy y = -1.120 + 0.9988 x Diseased y = -0.2238 + 0.8582 x  Test Hypothesis of common slope  F= 2.21  DF1= 1  DF2= 291  C.L.=0.95  P r o b a b i l i t y = 0.137  Table 8 - A n a l y s i s of covariance t a b l e based on the n a t u r a l l o g a r i t h m of b a s a l area increment, using the n a t u r a l logarithm of D.B.H. squared x Height as c o v a r i a t e (confidence = 0.95).  Source  df  SS  Plot  9-  10..22  Status  1  P l o t x Status Covariate  MS  F-ratio  Probability  1 ,. 1 3  1 .675 .  0.,095  2..47  2.,47  6,.240  0.,034**  9  3..56  0..39  0,.584  0..809  1  252. .18  252. .18  Residual  274  185, .78  0,.68  TOTAL  294  471 . 50  371 .940 ,  0..000**  37  There was a s t a t i s t i c a l l y natural  logarithm  healthy  trees.  significant.  note  is  increment  that be  difference  the  escape.  However,  as  between  the  difference  e i t h e r age or diameter.  the  diseased and was  highly  l a r g e t r e e s grow  trees. tomentosus i s p r i m a r i l y  s t r e s s e d t r e e s while l a r g e vigorous  trees  range of t r e e s i z e s i n the diseased and  c l a s s overlap greatly  significant  of  covariate  expected,  i s tempting to conclude that P. of s m a l l e r ,  using  to  area  f a s t e r than small  a disease  healthy  basal  Also This  significantly It  of  significant  (Tables  between This  11 and 12). There  healthy  i s no  and i n f e c t e d t r e e s f o r  i s based on a t e s t of d i s t r i b u t i o n  the Chi-square technique with both s e t s of t a b l e s . Table 9 - Natural logarithms of 5 year basal area increment f o r healthy and diseased t r e e s .  Healthy  n  1 25  Diseased  1 70  Observed mean  6.749  6.405  P r e d i c t e d mean  6.662  6.470  1 .262  1 .253  0.07 5  0.064  Observed standard  deviation  Standard e r r o r of the mean  The between  calculations basal area  shown  below  partition  the  increment of healthy and diseased  difference trees  into  two p a r t s , namely that due to d i f f e r e n c e s i n t r e e s i z e , and that due to d i s e a s e .  38  By t a k i n g the n a t u r a l s of the predicted  means  (Table  logarithmic  values  of  the  9), the d i f f e r e n c e between healthy  and  infected trees i s ; 782 The  - 645  =  137  d i f f e r e n c e of the n a t u r a l s of the observed 853  The  - 605  =  means i s ;  248  p r e d i c t e d mean accounts  for v a r i a t i o n due  to t r e e s i z e which  is; 137  =  55.2%  248 Applying area  these  f i g u r e s to the d i f f e r e n c e s i n untransformed b a s a l  increment c i t e d above, i t i s concluded  l o s s due  to disease  that  the  increment  i s about  1708.2 - 1087.3 x .55  =  20%  1708.2 The  data  were  namely (1) h e a l t h y ,  also  analyzed  using four d i s e a s e c l a s s e s ,  (2) decay r e s t r i c t e d  to r o o t s , (3) decay  s t a i n at root c o l l a r but not at breast h e i g h t , and s t a i n at breast h e i g h t . very area  unbalanced  However t h i s  design,  and  (4) decay  resulted  increment between disease c l a s s e s were s i g n i f i c a n t .  currently  classified  as  increment over  diseased  v a r i a b l e i s represented  by calendar  rings  latewood.  from  the  1982  averages the b a s a l area Table  11  f o r age  and  in  a  none of the d i f f e r e n c e s in b a s a l  9 shows the p a t t e r n of b a s a l area  (see  approach  and  and h e a l t h y . years, It  follows  increment f o r t r e e s distribution).  or  of  time for The the  Figure trees  dependent number  of  that each c l a s s different  ages  Trees c u r r e n t l y healthy  39  ^ J £  Mean Basal Area Increment of Healthy versus Infected Trees 2200-1  3  XT  Number o f Rings from Cambium  Figure 9 - Mean basal area increment over time f o r healthy white spruce, and t r e e s i n f e c t e d with Polyporus tomentosus. show a c o n s i s t e n t l y g r e a t e r basal area increment over time diseased t r e e s . trees  are  difference  T h i s may be a t t r i b u t e d to the f a c t that h e a l t h y  on average somewhat l a r g e r than d i s e a s e d t r e e s . i n basal area increment between h e a l t h y and  trees  f o r the p e r i o d of 1977-1982 i s a consequence  by p.  tomentosus.  just  than  become  In other words, increment l o s s e s  d e t e c t a b l e i n the stands examined.  s t a t e d that the r a d i a l  The  infected  of i n f e c t i o n have  Whitney  only (1962)  increment of dead t r e e s decreased at 5-15  years p r i o r t o death of the t r e e , but i n f e c t i o n , based on fungal  40  growth r a t e s , must have been e s t a b l i s h e d 20 to 30 years p r i o r to a measurable d e c l i n e .  As the  expect  become  the  loss  to  infection intensifies, more  one  might  severe as the t r e e ' s energy  becomes a l l o c a t e d more towards the  production  of  adventitious  roots. The  GENLIN  test  numbered s i x to ten and  was  also  eleven  to f i f t e e n .  group of 5 year annual growth other  words,  only be  shown  the to  basal be  i n f e c t e d t r e e s f o r the to  uncertainty  performed f o r increment  rings  area  showed  different  f i v e years.  about disease  recent  significance.  increment of healthy  significantly last  Only the most  rings  trees  from  T h i s may  In could  that  of  in part be  due  s t a t u s of t r e e s f i v e or more years  ago. 5.2.3  M o r t a l i t y And Mortality  small,  to  i n the ten study p l o t s was  understory  standing P.  Windthrow  trees.  One  hundred  t r e e s were observed, and tomentosus in 82 cases.  largely restricted and  fifty  of these death was It was  seven dead attributed  not p o s s i b l e to estimate  an annual r a t e of volume l o s s due  to m o r t a l i t y s i n c e i t was  determined  standing  Table  how  long  the  10 shows the percent  dead  t r e e s had  of spruce t r e e s i n f e c t e d .  no s i g n i f i c a n t d i f f e r e n c e between p r o p o r t i o n or dead t r e e s . likely  to  die  healthy  small  a t t r i b u t e d to causes such as crown Table  t r e e s , and  not  been dead. There  infected for  T h i s shows that small diseased than  to  is  living  t r e e s are no more m o r t a l i t y may  suppression.  10 - T o t a l number of spruce t r e e s i n v e s t i g a t e d , with the percentage i n f e c t e d and percentage dead.  be  41  Non-infected  Living Dead % Dead  Infected  % Infected  125  170  57%  75  82  52%  37%  33%  Six l a r g e green windthrown t r e e s were observed i n the plots. root was Two  The  cause of windthrow was  system by P. considerably percent  a rate of  tomentosus.  greater  were  rotation, considerable The  remaining  openings  c l e a r l y d e t e r i o r a t i o n of  The  t o t a l volume of these  than that of the dead standing  of the stems examined had  loss  stand  created.  maintained  study  over  been windthrown. the  volume l o s s e s to the  remainder stand  would  the trees  trees. If such of  the  result.  would a l s o be much l e s s wind-firm with  the  42  Table  11 - Percent of the t o t a l number of spruce t r e e s by age c l a s s  Age  Healthy  Diseased  30-35  1.6  0  36-40  1.6  0  41-45  1.6  8.2  46-50  8.0  12.9  51-55  30.4  30.0  56-60  30.4  33.5  61-65  18.4  13.5  >65  8.0  1.9  100 N  125  100 170  43  Table  12 - Percent of t o t a l spruce t r e e s by diameter  Diameter  Healthy  (cm)  class  Diseased  6-14  47.2  50.0  1 5-22  36.8  44.2  23-30  12.8  5.3  >31  3.2  0.6  1 00  1 00  N  If exposure  1 25  to inoculum  1 70  i s the determining  event  of  infection  one would expect l a r g e t r e e s to c o n t a c t inoculum more f r e q u e n t l y than s m a l l e r t r e e s . s i z e of 295 twice  as  However, as seen i n Table 12, with a sample  t r e e s , healthy t r e e s g r e a t e r than 23 cm diameter abundant  as  diseased  trees.  We  cannot a s s e r t  t r e e s of a l l s i z e s are e q u a l l y s u s c e p t i b l e , and appears that l a r g e t r e e s escaped trees.  However  the  are that  in t h i s study i t  i n f e c t i o n more o f t e n than small  difference  was  not  statistically  signi f icant. 6.  ECOLOGICAL CLASSIFICATION A  list  of p l a n t s observed on the study p l o t s i s presented  in appendix  A.  Based on  fitted  an  ecological unit.  to  found d u r i n g t h i s study. not designed to e l u c i d a t e  these  species'  lists,  the  site  was  F i g u r e 10 shows a t y p i c a l  soil  I t should be noted that t h i s study site/disease  relationships,  was  however  44  sandy  the  F i g u r e 10 - S o i l p i t from J e r r y Creek with w e l l d r a i n e d loams. The s o i l p r o f i l e had a t h i n organic l a y e r , with e l u v i a t e d Ae z o n e , and an e n r i c h e d Bf l a y e r .  e c o l o g i c a l  i n t e n s i t i e s such  of  u n i t s P.  to  timber enough,  the  prepare  a  S B S j / 0 1 . 2 ,  and  suggestions  for  are one the  Other  guide  f i r s t  b e n e f i t s  of Of  the  on  Columbia  c l a s s i f i e d as  i n d i c a t o r s  Based  B r i t i s h  i n t e r p r e t a t i o n s . three  p r o v i d e  tomentosus.  c l a s s i f i c a t i o n s .  c l a s s i f i c a t i o n s , able  c o u l d  as  are  M i n i s t r y  f i v e the  SBSj/06.  s u b a s s o c i a t i o n  from  e c o l o g i c a l  of  F o r e s t s  i n t e r p r e t a t i o n s  s i t e s ,  s i t e  ( j / 0 1 . 1 )  was and  i n t e r e s t i n g l y  S B S j / 0 1 . 1 , The  r e l a t i v e  gained  these  s i v i c u l t u r a l the  of  an  are  one  as  the  p r e p a r a t i o n to  mix  the  45  humus l a y e r with the mineral l a y e r a f t e r a broadcast burn and regenerate  the  site  with lodgepole pine and  spruce.  The  site  would then be managed f o r sawlogs on an 80 year r o t a t i o n . practice  would  e s s e n t i a l l y e l i m i n a t e any P.  in small root p i e c e s .  would  This  tomentosus l i v i n g  Larger stumps not t o t a l l y consumed i n the  broadcast burn would pose t h r e a t s i f h e a v i l y i n f e c t e d , l o n g e v i t y of such  to  inoculum  has yet to be determined.  be b e t t e r spruce growth, and as observed  but The  the  result  a good pine crop  as w e l l . For the second for  s u b a s s o c i a t i o n (j/01.2) the  s i l v i c u l t u r e are to prepare  burning  and  technique  recommendations  the s i t e f o r spruce by  broadcast  to manage the stand f o r 80 year o l d sawlogs.  is clearly  a mistake  A broadcast burn  would  continuing  disease,  root  do  i n areas with any  little and  to  the  remove  P.  This  tomentosus.  any  threat  of  r e s u l t i n g stand would soon  become i n f e c t e d . The growing  final on  s u b a s s o c i a t i o n (J/06) should not have had  it  at  a l l , a c c o r d i n g to the s i l v i c u l t u r e  guide.  Instead the guide recommends the p l a n t i n g  of  Douglas-fir,  stems of such s p e c i e s  on the J/06 our  or  lodgepole p i n e .  The  few  species  spruce  s i t e were growing much b e t t e r than the  preliminary  observations  spruce.  as  If  on host s p e c i f i c i t y are c o r r e c t ,  such a s p e c i e s change would a l s o a v o i d Polyporus rot.  such  tomentosus root  46  V. Polyporus interior 13  tomentosus i s a common root  immature spruce  percent  CONCLUSIONS  of  the  stands.  total  Diseased  area  in  disease  in  central  t r e e s occupied  28.4 .+  the s i x randomly s e l e c t e d  stands. I n f e c t i o n takes place v i a root contact between healthy diseased  trees.  roots, k i l l i n g  The  fungus advances e c t o t r o p h i c a l l y on  them r a p i d l y , but  r e s t r i c t e d to the Polyporus  occur  tomentosus  attack  i s not organized  clear  Douglas-fir,  in  Losses windthrow.  between  i s diseased.  and  There  the extent of i n f e c t i o n  is  in the  from  increment  at  a,ge  55  of a l l spruce  trees,  or  3.4%  this  yields  28.4  x 0.034 =  reduction,  butt  does not appear to be a f a c t o r  type examined.  i n 22.6%  f i r , lodgepole  pine,  r o o t s , remain u n i n f e c t e d .  Mortality  trees  Subalpine  trembling aspen, even though o f t e n in contact  result  of the age and  of d i s e a s e present.  age  the s e v e r i t y of crown symptoms.  with d i s e a s e d spruce  stands  the extent  relationship  Only spruce  plot  Rather,  Obvious crown symptoms are uncommon f o r the  root system and  occurs  affected.  in d i s t i n c t  p a r t s of a stand a p p a r e n t l y healthy and d i s e a s e d t r e e s  together.  diseased  is  inner xylem.  of t r e e s examined and no  small  i n l a r g e roots the pathogen  root rot c e n t r e s in which a l l t r e e s are diseased  and  Losses  in  current  Six of  were windthrown i n one  0.96  the  in  175  season.  and  stands  increment  i s measured to be 20%. trees.  decay  of  Butt decay diseased For whole  47  or a f u r t h e r annual l o s s of 1% of a l l the stems. 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Studies in Forest Pathology. XIV Decay of D o u g l a s - f i r in the C o a s t a l r e g i o n of B r i t i s h Columbia. Can. J . Bot. 32:630-653.  50  Van Groenewoud, H. 1956. A root d i s e a s e complex i n Saskatchewan white spruce. F o r . Chron. 32:11-13. Warren, G.L. 1956. Root i n j u r y to c o n i f e r s i n Canada by s p e c i e s of Hylobius and Hypomolyx ( C o l e o p t e r a : C u r c u l i o n i d a e ) '. For. Chron. 32:7-10. White, L.T. 1953. Studies i n F o r e s t Pathology. X. Decay of White pine i n the Timagami Lake and Ottawa V a l l e y Areas. Can. J . Bot. 31:175-200. Whitney, R.D. 1960. Stand-opening d i s e a s e and the p a t h o g e n i c i t y of Polyporus tomentosus F r . on white spruce ( P i c e a qlauca (Moench) Voss.T! Ph.D. T h e s i s , Queen's U n i v e r s i t y . , Kingston, O n t a r i o . 98 pp. Whitney, R.D. 1961. Root wounds and a s s o c i a t e d root r o t s of white spruce. F o r . Chron. 37:401-411. Whitney, R.D. 1962. Polyporus tomentosus F r . as a major f a c t o r in stand-opening d i s e a s e of white spruce. Can. J . Bot. 4_0 : 1631-1658. Whitney, R.D. 1963. A r t i f i c i a l i n f e c t i o n of small spruce roots with Polyporus tomentosus . Phytopath. 53:441-443. Whitney, R.D. 1964. I n o c u l a t i o n of e i g h t Saskatchewan t r e e s with Polyporus tomentosus . Can. Dep. For., Bimonthly Prog. Rep. 20(5):3. Whitney, R.D. 1965. M y c o r r h i z a - i n f e c t i o n t r i a l s with Polyporus tomentosus and P. tomentosus v a r . c i rc inatus on white spruce and red p i n e . F o r . S c i . 11:265-270. Whitney, R.D. 1966. Germination and I n o c u l a t i o n t e s t s with b a s i d i o s p o r e s of Polyporus tomentosus . Can. J . Bot. 44 :1333-1343. Whitney, R.D. 1972. Root r o t i n white spruce p l a n t e d i n areas formerly h e a v i l y a t t a c k e d by Polyporus tomentosus i n Saskatchewan. Can. Dep. F i s h . F o r . , Bimonthly Res. Notes. 28 (4):24. Whitney, R.D. 1977a. Polyporus tomentosus root r o t of conifers. Dept. F i s h . E n v i r o n . , Can. F o r . Serv., Sault Ste. Marie, Ont., For. Tech. Rep. 18. 12 pp. Whitney, R.D. 1977b. Polyporus tomentosus root r o t on conifers. F o r e s t r y T e c h n i c a l Report, Can. F o r . Ser. (1977). No. 18, 10 pp. {En,27 r e f . , 3 pi.} Great Lakes For. Res. Cent., S a u l t S t e . Marie, Ont.  51  Whitney, R.D. 1977c. V a r i a t i o n of Polyporus tomentosus F r . c u l t u r a l 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 on c o n i f e r seedlings. Can. J . Bot. 55: 1389-1398.  in  Whitney, R.D. 1978a. Polyporus tomentosus root and butt r o t of t r e e s in Canada. Statement Paper f o r 5th I n t e r n a t i o n a l Conference (Kassel,Germany) August 1978 on Problems of Root and Butt Rot i n C o n i f e r s , pp. 283-297. Whitney, R.D. 1978b. Root r o t of spruce and balsam f i r i n northwestern O n t a r i o . I I . Causal fungi and s i t e relationships. Can. F o r . Serv., Sault S t e . Marie, Ont. Rep. O-X-284. 42 pp. Whitney, R.D. and H. Van Groenwoud. 1964. The rate of advance of stand-opening disease over a ten-year p e r i o d i n white spruce at Candle Lake Saskatchewan. F o r . Chron. 40 -.308-312. Whitney, R.D. and W.P. Bohaychuk. 1976. Growth of Polyporus tomentosus c u l t u r e s d e r i v e d from b a s i d i o s p o r e s , sporohore t i s s u e , and decayed wood. Can. J . Bot. 54(22):2597-2602. Wood, C.S., G.A. Van S i c k l e and T.L. Shore. 1984. Forest i n s e c t and disease c o n d i t i o n s , B r i t i s h Columbia & Yukon, 1983. Can. Env., Can. For. Serv., P.F.R.C. BC-X-246. Pp. 27.  52  APPENDIX A - ECOLOGICAL SITE CLASSIFICATION AND LISTINGS  VEGETATION  The f o l l o w i n g s i t e s p l o t s have been c l a s s i f i e d i n the b i o g e o c l i m a t i c u n i t of the Sub-Boreal Spruce J . Information from each p l o t i n c l u d e s a s i t e d e s c r i p t i o n , a v e g e t a t i o n l i s t i n g , some of these based on 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 a b b r e v i a t i o n s , and the accepted subzonal d i v i s i o n . A l l s i t e s are f u r t h e r g r a d a t i o n s of the SBSj/01, or the Mesic Oak Fern. S i t e j_ : Beaver F o r e s t Road, kilometer 44. Taken from p l o t B1. T h i s s i t e has been c l a s s i f i e d as the SBSj/01.2, a Mesic Oak Fern, Thimbleberry s u b a s s o c i a t i o n . The s i t e i s f l a t , s l i g h t l y u n d u l a t i n g , with no aspect. I t i s a f l u v i a l t e r r a c e , with s o i l s that are t y p i c a l l y Humo-ferric p o d z o l s . The s i t e i s submesicsubmesotrophic. A2 A3 B1 B2 C D  75% 20% 10% 05% 15% 97%  Piceene, A b i e l a n A b i e l a s , Piceene Abielas Abielas Aralnud, Corncan Pleusch, P t i l c r i ,  Rhyttri  Species l i s t : Abies l a s i o c a r p a P i c e a exg Pseudotsuga menziesi i Rubus p a r v i f l o r u s Aralia nudicaulis Cornus canadensis Lonicera involucrata Pleurozium s c h r e b e r i Ptilium crista-castrensis Maianthemum canadense O r t h i l i a secunda Rhytidiadelphus triquetrus Lycopodium annotinum Chimaphila umbellata Gymnocarpium d r y o p t e r i s Ribes l a c u s t r e M i t e l l a nuda Rubus pubescens Viburnum edule Rosa a c i c u l a r i s Lycopodium obscurum  53  S i t e Two : J e r r y Creek, 500 road. Taken from p l o t J567. T h i s s i t e i s l o c a t e d i n the J e r r y Creek v a l l e y , at the toe of a 20m s l o p e . The s i t e i s t y p i c a l of the SBSj1/01.1, the Mesic Oak Fern Black Huckleberry s u b a s s o c i a t i o n . The s o i l s are t y p i c a l of a leached b r u n i s o l developed from g l a c i a l - f l u v i a l m a t e r i a l . The s o i l p i t had c h a r c o a l , and had no seepage at 25cm depth. The s o i l was t y p i c a l l y a medium-textured,sandy loam with coarse fragments of 40.-50%, and some l a r g e r cobbles to 20cm diameter. The humus form i s a mor. The s i t e i s mesic-submesotrophic. A2 35% Piceene, A b i e l a s A3 15% Piceene B1 8% A b i e l a s B2 5% Rubupar, Vaccmem, L o n i i n v C 90% Gymndry, Rubuped, Corncan, D 99% Pleusch, P t i l c r i Species l i s t : Picea exq. Aralia nudicaulis Gymnocarpium d r y o p t e r i s CIintonia uniflora Smilacina racemosa Ribes l a c u s t r e Streptopus roseus Abies l a s i o c a r p a Cornus canadensis Rubus pedatus Lycopodium annotinum Viburnum edule O r t h i 1 i a secunda Pleurozium s c h r e b e r i Ptilium crista-castrensis Rhytidiadelphus triquetrus Tiarella trifoliata Vaccinium membranaceum Lonicera involucrata Heracleum sphondylium Veratrum v i r i d e Oryzopsis a s p e r i f o l i a Rubus p a r v i f l o r u s Spiraea b e t u l i f o l i a  Aralnud  54  S i t e three : Highway 16, 20 k i l o m e t e r s east of P r i n c e George. Taken from survey stand. This s i t e has s i l t y c l a y l a c u s t r i n e l u v i s o l s o i l s with a moder humus. I t i s a f l a t area c h a r a c t e r i z e d by p e r i o d s of high water. The s i t e i s t y p i c a l of the SBSj1/01.1, the Mesic Oak Fern, Black Huckleberry s u b a s s o c i a t i o n . A1 A2 A3 B1 B2 C D  5% 15% 20% 03% 02% 10% 99%  Piceene Piceene Piceene, Abielas, Sorbsco, Corncan, Ptilcri,  Species l i s t : Spirbet Loni inv Vibuedu Vaccmem Petapal Achimil Pleusch Ptilcri Rosaac i Poputre Piceene Actarub Corncan Astecon Rubupub Goodpbl Fragvir Astecil Linnbor Sorbsco Galitrf  Abielas Piceene, Corncan, Rubupub Vibuedu Rubupub Pleusch  55  S i t e four : Hixon 3800 road, Kilometer 12. Taken from Hixon study p l o t . T h i s s i t e i s t y p i c a l of the SBSJ1/06, the Submesic Queen's Cup s u b a s s o c i a t i o n . The s o i l s are c l a s s i f i e d as shallow (3cm) mor humus forms on a ferro-humic p o d z o l . The s i t e i s t y p i c a l l y submesic to submesotrophic . The texture of the s o i l i s a loamy sand with 10% coarse fragments, probably f l u v i a l . The s i t e has about a 4% s l o p e , with a south a s p e c t . The s i t e occurs on the lower s l o p e . A2 40% Poputre, Betupap, Piceene, Pseumen A3 40% Piceene, A b i e l a s B1 15% A b i e l a s , Piceene B2 2% A b i e l a s C v a r i a b l e 5%-80% Corncan, Rubuped D 100% Pleusch, P t i l c r i Species l i s t : Picea exg. Pseudotsuga menziesi i Pinus c o n t o r t a Betula p a p y r i f e r a Cornus s e r i c e a Lycopodium obscurum Goodyera o b l o n g i f o l i a Rubus pedatus Lycopodium annotinum Veratrum v i r i d e O r t h i 1 i a secunda Clintonia uniflora Streptopus roseus Tiarella trifoliata Sphagnum c a p i l l a c e u m Ptilium crista-castrensis Pleurozium s c h r e b e r i P o l y t r i c h u m juniperinum Abies l a s i o c a r p a Linnaea b o r e a l i s Lonicera involucrata Streptopus a m p l e x i f o l i s Spiraea b e t u l i f o l i a Gymnocarpium d r y o p t e r i s Rosa a c i c u l a r i s Vaccinium membranaceum Rhytidiadelphus triquetrus Aralia nudicaulis Populus tremuloides  56  S i t e f i v e : Tabor Mountain Northern s l o p e . Taken from survey stand. T h i s s i t e i s a t y p i c a l i n v e g e t a t i v e nature. I t appears to be i n a s e r a i stage. The s o i l s are t y p i c a l l y a morainal blanket mor, the s i t e i s north aspect on a 20% s l o p e . A mid-slope s i t e with such a slope and morainal parent m a t e r i a l would p l a c e t h i s i n the SBSj1/01. A2 A3 B1 B2 C D  80% 20% 2% 5%  Piceene, Betupap, A b i e l a s , Piceene  Poputre  Streamp, Actarub, S t r e r o s Brachyl  Species l i s t : Betupap Piceene Abielas Streamp Actarub Streros CIinuni Braculy Poptre  57  APPENDIX B - STEM MAPS OF THE TEN STUDY PLOTS Ten maps are presented from the p l o t work i n the P r i n c e George East Forest D i s t r i c t . B1 to B4 are from four s i t e s at the Bowron R i v e r . DC1 to DC3 are from three s i t e s northwest of the latter. Two s i t e s , J567 and J568 are i n the J e r r y Creek drainage, and PLOTHIXON i s from the Hixon area. A l l maps are s c a l e d 1cm = 1m The codes f o r the maps are d e s c r i b e d as f o l l o w s ; Number 1 through 4 correspond to r e l a t i v e amounts of Polyporus tomentosus on white spruce, such that 1 = healthy 2 = d i s e a s e at roots only 3 = d i s e a s e at roots and root c o l l a r 4 = d i s e a s e at breast height An "x" r e p r e s e n t s a dead t r e e , and i f the dead t r e e had any sign of P. tomentosus i t was denoted as X . An  "a" r e p r e s e n t s a non-spruce t r e e .  N O R T H  8S  N O R T H  69  N O R T H  ID  09  61  « i  a o  N  N O R T H  39  N O R T H  n  £9  N O R T H  f9  N O R T H  o  1*5  N O R T H  3 1  99  N O R T H  Cr,  •44  LS  68  APPENDIX C - RELATIONSHIP BETWEEN TREE SIZE AND BASAL AREA INCREMENT. T h i s c o n t a i n s four graphs showing the general shape of data points. t r e e volume i s r e p r e s e n t e d by DBH squared times h e i g h t . M u l t i p l e occurrences of p o i n t s i n the same p o s i t i o n are represented by numerical v a l u e s which count the number of such overlaps. Rotcod 2 s p e c i f i e s a l l t r e e s with any amount of evident P. tomentosus , while rotcod 1 s p e c i f i e s a l l h e a l t h y t r e e s . The t h i r d and f o u r t h graph show the r e s u l t i n g l i n e a r i t y of p o i n t s when logs are taken f o r both the x and y axes. It i s of note that four p o i n t s from the f i r s t graph, two from the second, and one p o i n t from the f o u r t h graph are missing due to the upper l i m i t of the y a x i s .  SCATTER  PLOT N=  6AI 8000.O  +  7 111.1  +  6222.2  +  5333.3  +  4444.4  +  3555.6  +  2666.7  +  1777.8  +  888.89  +  + 0.  <\> ROTCOD : 1 BASAL AREA INCREMEN1 VS. 121 OUT OF 125 2 7 . B A I VS. 29.D2H  *  2* * * * ** **** 2***** 3  3**2 * •3524522  * *  *  **  (DBH + * 2 )+HEIGHT  * •  **  + 3333.3 1666.7  6666.7 5000.0  10000. 8333.3  13333. 11667.  D2H 15000.  SCATTER  PLOT <2> ROTCOD:2 LN(BASAL AREA N=, 169 OUT OF 170 3 3 . L B A I VS.  LBAI 8.5369  +  7.9918  +  INCREMENT) VS. 34.LD2H  LN((OBH**2)*HEIGHT)  *  * *  + *  7.44G7  +  **  +  6.3566  +  *  *  *  *  2  **  *  ** * * * * * * * * * * *  *  *  *  *  +  2  * *  *  * * *  *  * +  i  *  * • * * < *  * *  *  *  + * * * *  * *  2  *  **  +  2  *  *  *  +  5.8115  *  •  + 6.9016  •  2  •  *  *  *  *  *  2  +  *  ** * A  5.2664  *  +-  *  * ** * * *  4.7213  +  4.1762  +  3.6312  +  + 5.1156  +  +  *  5.6240  +  + 6.1325  +  + 6.6410  +  + 7.1494  +  + 7.6579  +  + 8.1663  +  + 8.6748  +  + 9.1832  +  + LD2H 9.6917  SCATTER BAI 8000.0  PLOT N=  <2> ROTCOD:2 BASAL AREA INCREMENT VS. 168 OUT OF 170 2 7 . B A I VS. 29.D2H  (DBH**2)+HEIGHT  +  +  7111.1  +  +  6222.2  +  +  5333.3  + *  4444.4  +  3555.6  +  2666.7  +  * * 1777.8  * *  ** *  + * +  *  * 2  2 *  +  0.  * **2 2 322 * 3 * * * 223** 2 2 •32X62*42* * * + 22 ** +  *  *2** * *2 * * *2  888.89  ***  *  **  *  «  2 2*  *  **  *  3333.3 1666.7  6666.7 5000.0  1000O. 8333.3  13333. 11667.  D2H 15000  <SCATTER BYSTRA TA VAR = L B A I . L D 2 H C A S E S = N E G B A I : 0 STRAT = ROICOD HEAD=1 1 LN(BASAL SCATTER PLOT <1> ROTCOD:1 LN(BASAL AREA INCREMENT) VS. L N ( ( D B H * + 2 ) ' H E IGHT ) N= 125 OUT OF 125 33.1.BAI V S . 34 . LD2H LBA I 9.9882 +  9.3744  +  8.7606  +  8.1469  + *  +  •*  *  *  * 6.9193  *2  *  «  INCREMENT) 'VS . LN((DBH +* 2)+ H E I G H T ) >  *  + t 2  7.5331  AREA  2 2 * *  -i  *  +  * 4' : * *  * 6.3055  +  5.6918  +  5.0780  +  4.4642  +  * *  4.7957  6.0492 5.4224  *  *  +  ++  +  2  2  *  *  *  7.3027 6.6759  8.5562 7.9294  9.8097 T.I830  LD2H 10.436  

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