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Distribution and abundance of arboreal lichens and their use as forage by blacktailed deer Stevenson, Susan K. 1978

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DISTRIBUTION AND ABUNDANCE OF ARBOREAL LICHENS AND THEIR USE AS FORAGE BY BLACKTAILED DEER  by  SUSAN KINGSBURY STEVENSON B.A., Swarthmore College, 1969  A THESIS SUBMITTED IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE  THE FACULTY OF GRADUATE STUDIES (DEPARTMENT OF FORESTRY)  We accept t h i s thesis as conforming to the required standard  UNIVERSITY OF BRITISH COLUMBIA May, 1978 ~c) Susan K. Stevenson, 1978  In p r e s e n t i n g t h i s  thesis  in p a r t i a l  f u l f i l m e n t o f the requirements f o r  an advanced degree at the U n i v e r s i t y of B r i t i s h Columbia, the L i b r a r y s h a l l I  make i t  freely available  f u r t h e r agree t h a t p e r m i s s i o n  for  I agree  reference and  f o r e x t e n s i v e copying o f  this  that  study. thesis  f o r s c h o l a r l y purposes may be granted by the Head of my Department or by h i s of  this  representatives. thesis  It  is understood that copying or p u b l i c a t i o n  f o r f i n a n c i a l gain shall  written permission.  Department of The U n i v e r s i t y o f B r i t i s h  Columbia  2075 Wesbrook Place Vancouver, Canada V6T 1W5  Date  TJA^JL  //  /f7t?  not be allowed without my  ABSTRACT  Biomass of blacktailed  arboreal  d e e r was  abundance o f  lichens  studied.  used a s  s a r r a e n t o s a and  b i o m a s s was  measured by  sampling the  A s y s t e m of  visual  and  to  predictive  crown, and  0,75)  was  trees  to  infrared  {Y =  estimate of CL  trees.  a i r photography assessed,  use  Biomass o f ranged from  21  {A  using  using  to  1528  of  the  82  productivity  was  lithin  the  s i t e s studied,  abundant on above 500  m,  moderate t o where t r e e  and  negatively  related  was  lichen  have  densitometry strong  14  lichens.  plots  studied  vegetative related  to  lichen  elevation  in lichen  abundance.  to l i c h e n  Alectoria  poor.  sampled  lichen  abundance,  (s.1.) was  steep south-facing slopes, growth  =  colour  neither  the  2  photo  with  a s p e c t , and  variation  Forest  range o f  on  the  376,89; r  inventorying  were measured and  the  of  abundance o f a r b o r e a l  kg/ha. P h y s i c a l  p e r c e n t of  =  x  large-scale  demonstrated t o  abundance. Taken t o g e t h e r , s l o p e , for  for  (sensu l a t o )  plots  lichen  biomass values from  d e n s i t o m e t r y and  in inventorying  A  a portion y  each method, but  was  Alectoria  characteristics  accounted  where Y i s  S ,  trees.  developed  sampling.  n = 40;  value of  a tool  Lichen  abundance was  x CL),  lichen  spp.  assess  from f e l l e d  l i c h e n c o v e r on  The as  photo i n t e r p r e t a t i o n for  lichen  Some p o s i t i v e r e l a t i o n s h i p s  abundance were f o u n d  potential  lichens  i s crown l e n g t h ;  unsampled  interpretation.  nor  158.03  used t o e x t r a p o l a t e  abundance was  Bryoria  b i o m a s s e s t i m a t e s o b t a i n e d by  b i o m a s s , A i s an tree  estimates of  eguation  by  T h r e e methods were used t o  Alectoria  related  winter food  at  most elevations  iii  To  assess  availability  deer,  litterfall  three  sites  was  measured i n s i d e  where l e v e l s  relationships  of d e e r  between l i t t e r  examined. Q u a n t i t i e s o f  poor  winter  winter  (p<0.05) A l e c t o r i a  litter  and  percent  31.9  was  present  o f a v a i l a b l e g u a n t i t i e s on  litter  the  on The  w e a t h e r were  were  151.2  69.9 kg/ha/180  significantly  inside  of A l e c t o r i a  by  kg/ha/180 d a y s i n a  a l l three s i t e s ,  Utilization  were known.  litter  range a r e a ,  exclosures; g u a n t i t i e s of non-lichen significantly.  utilization  outside exclosures  i n winter  (s.1.)  winter  On  their  d e p o s i t i o n r a t e s and  range area,  range area.  and  use  Alectoria  kg/ha/180 d a y s i n a s e v e r e days i n a m i l d  o f l i c h e n s and  was  than  outside  d i d not  differ  37,  and  three  more  53,  52  sites,  respectively. The winter  relationship  h a b i t a t by b l a c k t a i l e d  group c o u n t s ,  t r a c k counts,  A r e a s s e l e c t e d by high  between l i c h e n  in lichen  deer as  abundance.  deer  and  winter  the  abundance and  was  assessed,  data  selection  based  on  of  pellet  of other i n v e s t i g a t o r s .  r a n g e tended  t o be  moderate o r  iv  T A B L E OF CONTENTS  ABSTRACT  i  LIST  OF T A B L E S . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .  LIST  OF F I G U R E S  LIST  OF A P P E N D I C E S . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .  i v i  . . . . v i i i x  ACKNOWLEDGEMENTS  X  I.  1  INTRODUCTION Rationale  ...........................................  Objectives The II.  Study  3 Area  ......................................  EACKGBCUND Dse  Nutritional  L i c h e n s By U n g u l a t e s  V a l u e Of A r b o r e a l  Of A l e c t o r i o i d  I I I . , QUANTIFICATION  Visual  14  L i c h e n s ......................  16  OF B I O M A S S OF AEBOHEAL FORAGE L I C H E N S  E s t i m a t e s Of L i c h e n  Photo  Forage  7 L i c h e n s ........  Methods  Air  Quantification  ..  Computations Computing  26  Abundance  26  Of L i c h e n  Abundance  .....  And R e s u l t s  Biomass T o t a l s  .........................  Quantification  Of Methods  Biomass R e s u l t s RELATIONSHIPS  ............................  ............................  Compared W i t h  BETWEEN L I C H E N  Literature  BIOMASS AND  39 39 45  . . . . . . . . . . . . . . .......... .. . . . . . . . . . . . . . . . .  Evaluation  30 36  R e s u l t s Of A i r P h o t o Discussion  22  . ...  Bicaass Sampling  IV.,  4 7  Of A r b o r e a l  Ecology  1  SITE  Reports  .  50 50 55  V  CHAfiACTERISTICS  ..  ..... . . ..... .  Methods  60  Besults  ..  63  Vegetation Communities  ...........................  63  Measures Of P h y s i c a l Environment .................  68  F o r e s t Measures  75  Discussion V.  58  AVAILABILITY  81  OF ASBOREAL LICHENS AND UTILIZATION  BY DEES  Methods ........................... ,. .. Litterfall  85 87  Measurement ...........................  87  H a b i t a t S e l e c t i o n ................................  90  Besults  .............................................  Total L i t t e r f a l l  Q u a n t i t i e s ......................  L i c h e n L i t t e r f a l l In R e l a t i o n To L i c h e n  92 92  Biomass ,. 94  L i t t e r D e p o s i t i o n I n R e l a t i o n To Time And Heather  94  U t i l i z a t i o n Of L i t t e r f a l l ........................ 102 Deer Use Of L i t t e r f a l l P l o t s ..................... 106 Lichen Abundance And Deer Use .................... 107 D i s c u s s i o n .......................... ... ............. 110 Litterfall  ..................................... .. 110  Deer Use And l i c h e n Abundance VI. , SUMMABY  116  AND MANAGEMENT RECOMMENDATIONS ................. 118  Summary  ...... ........ . .... 118  Eecommendations ..................... ................ 122 LITEEAIUEE CITED  124  vi  LIST OF TABLES  I.  N u t r i t i o n a l value of a r b o r e a l forage l i c h e n s  II.  Biomass o f A l e c t o r i a on sampled  III.  Optical  with high and low l i c h e n biomass  Spearman's rho values f o r p r e d i c t i o n s abundance on i n d i v i d u a l t r e e s  V.  .............. 4 1  d e n s i t y values and red:green f i l t e r  of t r e e s IV.  trees  ....... 1 5  ratios  .......... 4 6  of l i c h e n  ......................,48  Comparison of reported biomass t o t a l s of  epiphytic  lichens  5 6  VI.  Site c h a r a c t e r i s t i c s of plots  6 4  VII.  Litterfall  VIII.  Alectoria  quantities litterfall  i n s i d e e x c l o s u r e s ............ 9 3 i n r e l a t i o n to A l e c t o r i a .....  standing crop  i n s i d e and o u t s i d e e x c l o s u r e s . 1 0 4  IX.  Litterfall  X.  Deer use and snow depths at P l o t s 3, 4, and 5, March 28-29, 1977  quantities  ........................................ 1 0 8 and l i c h e n biomass ............... 1 0 9  XI.  Habitat s e l e c t i o n  XII.  Comparison o f reported q u a n t i t i e s litterfall  9 5  of l i c h e n  ......................................... I l l  «  vii  LIST OF FIGURES  1.  L o c a t i o n of the study area  5  (Jones 1975)  2. , Measurement of crown width p a r a l l e l and p e r p e n d i c u l a r to is ix© sXcp^ • # • * • • * • * * • . « '• 3.  *(•••••«.,•.'•'*•••  •  '• •'  ••  *'-••'•.•••• 29  D i v i s i o n of a t r e e i n t o l a y e r s f o r v i s u a l e s t i m a t e s of 1 xc i i ©n  cii^un d c^n c €  • • • • • • •  «* • • * * • %-•>• • • •  • • • * • ... 29  :  '*» . M f i c t o r i a saraentosa on D o u g l a s - f i r ;  normal c o l o u r  i  film  (A) and c o l o u r i n f r a r e d  film  with Jfratten 12  f i l t e r and two CC20 magenta f i l t e r s 5.  Colour i n f r a r e d a i r photographs o f p l o t with low 1 xch-€ E  6.  31  (B)  JDXGIUCL ss  * • *"• • * • • • • • * • • • "• *./#:» *"*,"• :  • *••#>•»" #• • • • •  Colour i n f r a r e d  a i r photographs o f p l o t with high  1 i c li s n 13 x o sts ss  »• * • « • • * • * *  34  * * * * • • • • • •. • • • • •« * • • • * .  7.  Regression used t o c a l c u l a t e l i c h e n biomass of v i s u a l  8.  Lichen biomass per t r e e based  on independent  estimates by two o b s e r v e r s ...:....... 9.  35  visual  *..... >....... 44  Vegetation community and A l e c t o r i a biomass (A): v e g e t a t i o n community and percent A l e c t o r i a  IB) . . . . . . .  69  ..... 70  10.,  R e l a t i o n s h i p between s l o p e and percent A l e c t o r i a  11.  R e l a t i o n s h i p between aspect and percent A l e c t o r i a  12.  L i n e a r r e g r e s s i o n o f percent A l e c t o r i a on p o t e n t i a l  .... 71  annual r a d i a t i o n . . . . . . . . . . . . . . . . . . v .... . . . v i . . . . . . . . . .  72  13.  L i n e a r r e g r e s s i o n o f percent A l e c t o r i a on e l e v a t i o n ..  73  14.  R e l a t i o n s h i p between crown c l o s u r e and percent A l e c t o r i a i densiometer v i s u a l estimates  (A); moosehorn (B);  (C) ......... .... ........ . . * •. v * . . . « * . . . . .  76  viii  15.  R e l a t i o n s h i p between b a s a l a r e a  16.  Linear r e g r e s s i o n o f percent codcffiinant  17.  tree layer  ACT~tor13^  Alectoria  on h e i g h t o f  * • • • • • •  • •» * » • •  • • •"» •  •*-#-• • • -  19.  Deposition rates of Alectoria  litter  i n early  (E),  winter  (L),  late 21.  i n early  winter  (ft), and l a t e  winter  ( E ) , mid-winter  Precipitation winter  winter  lichen  (H), and 97  (L) , 1976-1977  ( f 4 ) , and l a t e at Port  i n early  winter  Hardy  (E) , m i d - w i n t e r  80  1S76-  {non-Alectorioid)  Deposition rates of a l l l i t t e r mid-winter  22.  plots  Deposition r a t e s of other litter  '  89  Layout o f l i t t e r f a l l  20.  79  l e n g t h / t r e e h e i g h t and  18.  mid-winter  . 78  Alectorla  ................................  R e l a t i o n s h i p between crown pcxcsnt  and p e r c e n t  winter ( E ) ,  ( L ) , 1976-1977.  ......  98  and Hoss Camp i n e a r l y  (M) , and l a t e  winter ( L ) ,  1 S1 € *" 1 S *77 • • * « # • • * * • * • * • • ••«:*•:••.•*••••*••*• • • • • • • • • • m- * • 1 0 0 23.  R e l a t i v e amounts o f wind clearouts  (B) i n e a r l y  (H) , and l a t e 24.  Alectoria  winter  utilization  i n mature  winter  timber  and  (E), mid-winter  (L) , 1976-1977 . on P l o t s  (A)  . . ,  V  . , . . , . . . . . . . . 1 0 1  3, 4, and 5, w i t h 9 5 %  c c n £ icls nc€ 1 x 01 i t s • • * • • • • • « « • • * • * * • * • • • * • •» •• • • *. * # • • * * * • 1 0 5 25.  Deposition r a t e s o f forage conifer  foliage)  (Rochelle 26.  litter  i n the study  ( l i c h e n s and g r e e n  area,  1978)......................  Deposition rates o f three l i t t e r f a l l Northwest  1973-1974  Bay, 1975-1S76  . . . . . . . . . . . . . . .  . . . .  114  components a t  (Kale unpublished  d a t a ) . . . . . 115  ix  L I S T OF  I. , C a l c u l a t i o n  of A l e c t o r i a  APPENDICES  (s.l.)  biomass t o t a l s f o r 131  plots II.  Differentiated  table  o f vegetation  data  146  X  ACKNOWLEDGEMENTS  I  received  personal f i n a n c i a l  from t h e Canadian Ltd.,  and  Wildlife  the U n i v e r s i t y  This study  was  Branch  Eastman,  i/c Wildlife  arrangements;  The of  by t h e B r i t i s h  a g r a n t t o Dr.  Research,  s t u d y was  conducted  Forest  J o n e s , and t h e s t a f f the trees  the  Dr. P.A.  Bunnell, Mr.  h i s cabin.  Regional  support.  Julius  shop.  Mr.  And  assistance  within  K a p i t a n y , Mr.  Mr.  Mrs.  I am  the  Gordon Mr.  the  Greg  Flowerdew  Henry  Shear  T. C a r l y l e h e l p e d  with  Committee, Dr..H.B.  especially  and  g r a t e f u l t o my  Schofield  constructive supervisor,  Dr.  F.L.  f o r h i s g u i d a n c e , e n c o u r a g e m e n t , and p a t i e n c e . Parker W i l l i a m s , of I n t e g r a t e d Resources  wish t o t h a n k  i n the f i e l d  support  Graduate  Murtha p r o v i d e d d i r e c t i o n  provided e x c e l l e n t I  me  Hebert,  the  t o acknowledge i n p a r t i c u l a r  of the f i r e  members o f my  criticism.  Ltd.  D.  f o r b i o m a s s s a m p l i n g , and  t o use  D.S.  f i e l d w o r k i n many ways. The  and  Dr.  P r o d u c t s , L t d . Many i n d i v i d u a l s  S t a n C h e s t e r , Mr.  o f Mr.  and  B u n n e l l . Dr.  w i t h t h e c o o p e r a t i o n and  assistance  me  Fish  the funding  provided a d d i t i o n a l  I would l i k e  allowed  Products  Columbia  F.L.  handled  company a i d e d me;  felled  Forest  study  Columbia.  have been c o m p l e t e d .  Biologist,  Canadian  Canadian  f o r graduate  w i t h o u t h i s a s s i s t a n c e and e n c o u r a g e m e n t ,  s t u d y would n o t Wildlife  through  Service,  of B r i t i s h  funded  Wildlife  assistance  o f my  and  Photography,  a i r photographs.  a l l t h o s e who with the  husband, David,  worked l o n g h o u r s  p r e p a r a t i o n of the encouraged  me  assisting  manuscript.  through  The  a l l staqes  the study.  1  I.  INTRODUCTION  Rationale The use of c l e a r c u t s and young second range  growth as winter  by b l a c k t a i l e d deer 1 0 d o c o i l e u s hemionus  Richardson)  has been documented f o r many areas  Gates  Miller  1968,  cclumbianus (Brown  1961,  1968), but where s n o w f a l l i s h i g h , mature  timber appears important to deer i n winter. Edwards example, found t h a t ungulate numbers i n B r i t i s h  (1956), f o r  Columbia  d e c l i n e d during years of deep snow, but t h a t adeguate cover could o f f s e t the e f f e c t s of snow. Cowan (19 56) r e c o g n i z e d the importance  of stands of c o n i f e r o u s t r e e s i n p r o v i d i n g s h e l t e r  from heavy s n o w f a l l s to b l a c k t a i l e d deer i n the northern p a r t of i t s range. Gates  (1968) noted that e a r l y s e r a i s t a g e s of  v e g e t a t i o n are of l i m i t e d  value to deer when food  i s r e s t r i c t e d by deep snow. Jones  availability  (1975) r e p o r t e d t h a t  on  northern Vancouver I s l a n d deer, i n winter, used mature timber h a b i t a t s more i n t e n s i v e l y than logged h a b i t a t s , e s p e c i a l l y when snow was  deep and s o f t .  However, not a l l f o r e s t types were found  to be of egual value, and f u r t h e r r e s e a r c h on winter h a b i t a t  was  recommended. Much of the area i n B r i t i s h Columbia  t h a t i s occupied by  b l a c k t a i l e d deer i s prime timber-producing land which i s s u b j e c t to e x t e n s i v e l o g g i n g . I t may  be e s s e n t i a l to the maintenance of  b l a c k t a i l e d deer p o p u l a t i o n s that a p p r o p r i a t e areas of mature timber be i d e n t i f i e d and set aside as h a b i t a t f o r w i n t e r i n g  2  animals.  This situation  into  c h a r a c t e r i s t i c s of  the  Because food forage i s l i m i t e d  has  i s often under  created  critical  the  canopy  must c o n s i d e r  northern  r a n g e of  are  Jones  heavily  as  1975;  u s e d as  Brypria. "beard  trees the  the  a major component o f  1968;  and  of  and  lichen"  forest  or  investigate  the  utilization  by  The  the  blacktailed  (Cowan 1945;  lichens  which  genera Dsnea^  the  to  deer.  d e e r as  forage  the  lichens  are  most  Alectoria^  often  referred  was  lichens  to  mature  litterfall  present study  arboreal  In  Gates  man's b e a r d " . They grow on  p r o d u c t i o n of  winter  deer a r b o r e a l  pendulous l i c h e n s  purpose of  and  forage a v a i l a b i l i t y .  most commonly a v a i l a b l e  floor.  range.  deer i n w i n t e r ,  blacktailed  to  research  of a mature f o r e s t ,  1978). The  long, "old  to  deer winter  winter d i e t  forage belong  They are  are  the  Rochelle  u r g e n t need f o r  blacktailed  range i n v e s t i g a t i o n s part  an  on  to and  their  3  Objectives  F o u r o b j e c t i v e s were d e f i n e d f o r t h e  1.  To d e v e l o p and e v a l u a t e methods o f q u a n t i f y i n g b i o m a s s o f  arboreal  2.  project:  forage  To r e l a t e  type,  lichens.  arboreal forage lichens  common f o r e s t  to vegetation  i n v e n t o r y measures, and o t h e r  community  site  characteristics.  3.  To a s s e s s  winter  availability  and u t i l i z a t i o n  of forage  lichens.  4. of  To r e l a t e winter  abundance o f a r b o r e a l f o r a g e l i c h e n s t o s e l e c t i o n  h a b i t a t by b l a c k t a i l e d  deer.  4  The  Study  Area  The study area i s l o c a t e d i n the Nimpkish V a l l e y on n o r t h c e n t r a l Vancouver I s l a n d ( F i g . 1). The s i t e o f most of the f i e l d w o r k was the southeastern  p o r t i o n of the Nimpkish  drainage,  near the confluence of Croman Creek and t h e Davie R i v e r . Geologic h i s t o r y , physiography, were described by Willms  and c l i m a t e of t h e study  area  (1971). The area i s mountainous, with  U-shaped g l a c i a l v a l l e y s a t about 300 m a . s . l . r i s i n g t o peaks of  1500 t o 1800 m. The c l i m a t e i s c h a r a c t e r i z e d by moderate  temperatures  but high p r e c i p i t a t o n i n winter. Snowfall i n the  study area i s g e n e r a l l y g r e a t e r than the 10 2 cm mean annual t o t a l recorded at Boss Camp, l o c a t e d 14 km northwest of the study area at an e l e v a t i o n of 140 m a . s . l .  ( H i l l m s 1971).  Vegetation i n the study area f a l l s within the C o a s t a l Western Hemlock Zone, the Mountain Hemlock Zone, and the A l p i n e Tundra Zone  (Krajina 1965). Most of the f i e l d w o r k was c a r r i e d out  w i t h i n t h e C o a s t a l Western Hemlock Zone. C l e a r c u t l o g g i n g began i n t h e study area i n 1947 (Willms  1971) and, at present, the  area i s a mosaic of r e g e n e r a t i n g stands aged 30 years or l e s s , and  mature stands aged 300 years or more. A l l study  p l o t s were  l o c a t e d i n mature stands. Although  many s p e c i e s of a r b o r e a l l i c h e n are present i n the  study area, the s p e c i e s most used as forage belong  to the  c l o s e l y r e l a t e d genera A l e c t o r i a and B r y o r i a . A l g g t o r i a sarmentosa  (Ach.) Ach. i s the most abundant s p e c i e s o f t h i s  group i n the study area. B r y o r i a g l a b r a (Mot.) Brodo and D. Hawksw. i s the most abundant of the Brioriae.. Other s p e c i e s  Figure .1.  Location of the study area (Jones 1975).  6  occasionally  found i n the study  D. Hawksw., B± o r e g a n a capillaris  from in  The  area  t r i c h o d e s . and B. t r i c h o d e s s u b s p .  i s more  reported  f o r forage, i s extremely rare  end o f t h e N i m p k i s h V a l l e y  more  where t h e  oceanic.  fieldwork  September 1977.  as important  ( o n l y one known s p e c i m e n ) . I t o c c u r s  at the north  Brodo a n d  (Michx.)  (Hot.) Brodo and D. Hawksw. The genus Usnea,  t h e study  climate  friabilis  (Ach.) Brodo and D. Hawksw., B. t r i c h o d e s  other l o c a l i t i e s  commonly  a r e B..  (Tuck.) Brodo a n d D. Hawksw., B.  Brodo and D. Hawksw. s u b s p . americana  area  was c a r r i e d  out during  the period  May 1976 t o  7  II.  Use  of Arboreal Lichens A number o f c e r v i d  In  g e n e r a l , consumption  associated  with  include the fallen  high  lower  trees,  and  Blacktailed litterfall. that  On  deer.  brought trees.  down by On  commonly  use  of a r b o r e a l l i c h e n  identified  as  h i g h w i n d s , and  found  Dsnea b a r b a t a •  1  of  broken  from  as  (1945)  rumen c o n t e n t s  L i c h e n s were consumed f r o m  snow and  forage  naturally  lichens available  V a n c o u v e r I s l a n d , Cowan which he  be  loggers.  by volume o f w i n t e r  by  that arboreal lichens,  logging, constituted  rumen s a m p l e s and item. Jones in  by  forage.  branches  recently  felled  the s o u t h e a s t e r n c o a s t of Vancouver I s l a n d Gates  (1968) o b s e r v e d winds and  a r b o r e a l l i c h e n s as  of a r b o r e a l l i c h e n s seems t o  trees felled  an a r b o r e a l l i c h e n  blacktailed  s p e c i e s use  s n o w f a l l . Sources  southern  36%  Unsulates  b r a n c h e s of t r e e s , l i t t e r f a l l ,  deer  constituted  by  BACKGROUND  were t h e t h i r d  (unpublished  rumen s a m p l e s  13%,  data)  2  of b l a c k t a i l e d  made a v a i l a b l e by  volume, o f  most i m p o r t a n t found deer  10%  and  collected  by  winter  winter  4%  strong  forage  arboreal lichen  during severe  and  I v i s i t e d Cowan's s t u d y a r e a n e a r G o l d s t r e a m L a k e i n J u n e , 1976 and n o t e d t h a t A l e c t o r i a and B r ^ o r i a s p p . were a b o u t a s common as Usnea spp. Although i t i s p o s s i b l e t h a t the animals were s e l e c t i v e l y c o n s u m i n g Dsnea. i t seems l i k e l y t h a t a l l p e n d u l o u s f r u t i c o s e l i c h e n s i n the rumen s a m p l e s were i d e n t i f i e d as Usnea b a r b a t a . 1  x  2  Canadian F o r e s t Products  L t d . , Woss, B r i t i s h  Columbia.  8  mild  winters, respectively,  Vancouver I s l a n d .  His f i e l d  were more h e a v i l y  used  i n the Nimpkish  Valley  o b s e r v a t i o n s suggested  than t h e s e  data i n d i c a t e .  on  that  have been low b e c a u s e  rumens were c o l l e c t e d  i n logged areas  deer  t h e same s t u d y a r e a , B o c h e l l e lichens  were a major d i e t a r y  (1978) f o u n d  most  (Jones  that  lichens  Lichen  g u a n t i t i e s i n rumen s a m p l e s may from  northern  1975). I n  Alectorioid  item f o r deer c o l l e c t e d  in  timbered  s t a n d s d u r i n g w i n t e r . They o c c u r r e d a t a volume o f 35.5%, w i t h 100%  f r e g u e n c y . The  ifiaultheria species  shalloni,  which had  w e l l , but  l i c h e n s a r e e a t e n by  they  may  Schroeder  hemionus hemionus) and feeding  on  Mountains  Alectoria  were o f s e c o n d a r y  mule d e e r and depth that  whitetailed  i n c r e a s e d (Schneewis  component o f t h e mule d e e r  deer  from  deer  as  winter  (Odocqileus  ( O d o c o i l e u s v i r g i n ^ anus)  fallen  t r e e s i n the  Columbia.  importance  deer;  5%.  as  In South  Selkirk  Dakota,  winter food items f o r  more l i c h e n s  were e a t e n  e t a l . 1972). S c o t t e r  as  snow  (1964) r e p o r t e d  d u r i n g p e r i o d s o f deep snow, t r a p p e r s i n w e s t - c e n t r a l  Alberta  c u t down t r e e s s o t h a t  arboreal lichens. indicate  that  decreased midwinter. lichens fall  white t a i l e d  sarmentosa  salal  o t h e r North American  both  of s o u t h e a s t e r n B r i t i s h  Usnea s p p .  than  n o t be a c r i t i c a l  (1974) o b s e r v e d  only to  a volume o f 43.6%; a l l o t h e r  o c c u r r e d i n volumes o f l e s s  Arboreal  diet.  l i c h e n s were second  Data  6.9%  were l e s s I n an  o f l i c h e n s by  mule d e e r n e a r  oak  fall  p r o b a b l y o c c u r r e d because  a b u n d a n t on  the  animals*  woodland h a b i t a t  in  the  e t a l . (1976)  o f rumen c o n t e n t s i n e a r l y  T h i s change  ranges.  p r e s e n t e d by W i l l m s  consumption  from  mule d e e r c o u l d f e e d on  Kamloops  t o 0.6%  in  arboreal  winter ranges northwestern  than  on  9  California, iii  the  lenziesii)  blacktailed t o 24%)  and  deer  use  caribou  of  and  many a u t h o r s . snowfall  Usnea  arboreal reindeer  and  the  changing t h e i r  more on  from  h e r d e r s sometimes f e l l  domesticated for  reindeer  40-50 cm.  entirely  on  has  arboreal  food.  been  that,  areas,  left  to arboreal  and  beard l i c h e n s  until  rest  (Alectoria  and  from f a l l e n  snow. F e l l e d l i c h e n s by  trees at logging  about  one-third  of  sites the  areas forest,  lichens,  and  often  the  (1975) f o u n d  January  Lapp  winter  down by  were used  animals.  that  i n the  when snow  jubata^-A  branches brought  Kola  lichen-bearing  Helie  of the  the  l i c h e n s are  JL. A l J l g x a j , km, sarmentosaJL a v a i l a b l e from t h e trees  alpine  i n F i n n i s h c o n i f e r f o r e s t s dug  the  terrestrial  to descend i n t o the  or break o f f  grasses  on  mountains of  reindeer  terrestrial  trees  iRanjifer  l i c h e n s than  snowfall  by  i n heavy  e s p e c i a l l y i n late winter,  During  (up  trees  documented  reindeer  In the  crusted  animals. Sulkava  g r o u n d l i c h e n s and  reached  spp.)  t h a t i n Sweden, a r b o r e a l  to reindeer,  branches f o r the  l a r g e s t volumes  lower branches c f  Union, w i l d  southern Urals,  (1969) n o t e d  necessary  the  (1955) r e p o r t e d  main w i n t e r  diet  though  (Book e t a l . 1 9 7 2 ) .  (Rangifer  when snow became dense and  Skuncke  year,  l i c h e n s , whereas i n low  were t h e  peninsula  the  Soviet  fed  reticulata  were f o u n d i n rumens o f  l i c h e n s on  Nasimovitch  tarandus tarandus)  lichens  spp.  i n winter  areas i n the  terrestrial  l i c h e n s Bamalina  throughout  occurred  The by  arboreal  snow  depths  they f e d  almost  .freigntji^  f c  lower  branches  storms or as  sources  of  melting of  10  In  North America,  concerning lichens, forest.  the  has  importance of  to c a r i b o u Cringan  winter  food  in  Lake s u p e r i o r .  been some  (Rangifer  He  argued  that  that  arboreal  the  from f a l l e n  t r e e s , independent  of c a r i b o u  who  immature as  that  and  that  guesticned  mature s t a n d s i n  by  c r u s t e d , but  northern  anecdotal  evidence  found only  boreal  forest  et  al.  available information  for  on  small  than i n the  when  amounts o f  ( S c o t t e r 1967).  A h t i and  Hepburn  the  habits  food  where b o t h were a v a i l a b l e . I n t h e f o r e s t , where snow was  d e e p e r and  and  He had  (1967) of  Ontario  p r e f e r r e d to  southern  part of  terrestrial  l i c h e n s were t h o u g h t t o be  north.  of  the  particularly  Ground l i c h e n s seemed t o be  abundant, a r b o r e a l  (1976),  use  l a r g e r g u a n t i t i e s might have been f o u n d i n winter.  His  maintain  i n rumens c o l l e c t e d i n December  lichens  important  levels.  winter.  i n winter.  less  Euler  than  available  made e x t e n s i v e  caribou  boreal  Islands  produced  population  lichens to caribou,  s a m p l e s been t a k e n l a t e r summarized  Slate  supply  forests to  Slate Island caribou  Usnea s p p .  i n the  speculated  later  arboreal  deep and  Alectoria  lichens,  sustained  r e g u i r e climax  (1964) p r e s e n t e d  importance of snow was  was  w e l l as  Scotter  January  caribou  populations  observed  of the  important  f o r e s t s , were p o t e n t i a l l y more i m p o r t a n t  l i c h e n s because of  steady  boreal  l i c h e n s were more  terrestrial  that  arboreal  tarandus c a r i b o u l . of the  t h a n woody browse t o c a r i b o u  i n climax  conclusion  controversy  lichens, including  (1957) r e p o r t e d  as  mainly  there  tree the  lichens  more  11  Bergerud of t h e  (1972) f o u n d t h a t  c o n t e n t s of e i g h t  collected  rumens o f  increasingly  Bergerud  on  considered  that  were a v a i l a b l e , b u t  preferred  required. . Miller  or  l i c h e n s and  grasslike plants  browse i n l a t e  winter  Consumption associated  that  their to  w i t h deep o r  thinner there  lichens  jPinus  was  no  i n bogs u n t i l  where t h e  i t s i n t e r c e p t i o n by  wind c r u s t .  Caribou fed  snow d e p t h s r e a c h e d  d e v e l o p e d . The they  fed  able  to o b t a i n  animals then  m a i n l y on  and  the  snow.  60  m a i n l y on cm  d e p t h s which r e a c h e d 65-70 cm  (1975)  black  and  was  and  on  cratering the  study  ridges  spruce greatest  on  was  arboreal  wind  was  s o f t e r because lichens  crusts  bogs t o  l i c h e n s . Apparently  l i c h e n s by  always  snow c o v e r  trees,  moved from t h e  terrestrial  ground  woody  lichen availability  (Larix laricina)  banksiana) r i d g e s  because o f  and  Stardom's  m a r i a n a ) bogs. Ground l i c h e n a v a i l a b i l i t y  jack pine  in  terrestrial  l i c h e n s i n w i n t e r i s not  i n southeastern Manitoba, a r b o r e a l  (Picea  necessarily  caribou  f o r m e d on  snow. I n  the  snow became  were not  d i e t from  arboreal  crusted  i n open t a m a r a c k  general,  l i c h e n s were used  they  when a h a r d c r u s t  of e p i p h y t i c  In  l i c h e n s as t h e  arboreal  54%  caribou  (1976) n o t e d t h a t  :  n o r t h w e s t e r n Manitoba changed  greatest  represented  Newfoundland  arboreal  because they  area  lichens  i n March under s e v e r e snow c o n d i t i o n s .  animals fed deeper.  arboreal  despite during  ridges, they  where  were  a v e r a g e snow a severe  winter. Arboreal caribou  l i c h e n s are  in British  the  early  part  fed  first  on  of  the  Columbia. the  p r i n c i p a l winter food In  mountain  Wells Gray Park, c a r i b o u  w i n t e r i n low  ground v e g e t a t i o n  of  and  elevation  spent  f o r e s t s where  t h e n , a f t e r snow became  they deep.  12  on  arboreal  seventy the  lichens  (37%)  (Edwards and  feeding  consumption  Bitcey  observations  of a r b o r e a l  1959) . T w e n t y - s i x  made d u r i n g  l i c h e n (Edwards and  January the  snow was  migrated  tim.ber.line f o r e s t s where t h e y  to  (Edwards and represented and  April  Bitcey 11 o f  1959). The (92%)  (Edwards and  Freddy  British  throughout  the  winter,  but  from  October to February  May.  Twenty-three of  during Caribou  the  forests  increased  feeding  the  the  l i c h e n s are  a  i n some a r e a s .  Usnea b a r b a t a  was  Eastman  u s e d by  March  extremely that  10 p e r c e n t Prince  of the  trees,  high  caribou.  valuable  food  mature s p r u c e - f i r survival. diet  Quesnel r e g i o n  in late  to  lichens.  Cowan e t a l . (1950) i m p l i e d  moose i n t h e  f o r e s t near  z o n e more  lower branches of  minor component o f t h e  (1978) n o t e d t h a t  pulmonaria c o n s t i t u t e d a coniferous  concluded  spruce-  observations  of arboreal  e s s e n t i a l to caribou  (Alces alces)  in  lower part of  a r b o r e a l l i c h e n s an and  January  f o r e s t zone  amounts o f l i c h e n a v a i l a b l e t o  were p r o b a b l y  Arboreal  used the  upper p a r t more f r o m  l i c h e n s from  Selkirk caribou  Columbia.  the  April  lichens  between  t r e e s were a l s o u s e d ; deep snowpacks at  Freddy c o n s i d e r e d the  l a s i p e a r pal  t h i r t y - t h r e e (70%)  m a i n l y on  though f a l l e n  for  mountain c a r i b o u  used t h e  and  until  they  S e l k i r k Range i n  p e r i o d October-May c o n s i s t e d  fed  elevations  Abies  a n i m a l s and  winter.  of  moose  that British  Lobaria  rumen c o n t e n t s  George, B r i t i s h  of  of  moose  Columbia.  i  Meules  (1965) f o u n d  no  evidence of  By  *  that i n the  J Pice a enqelmannii -  noted  1960).  of a r b o r e a l  observations  1960}  Columbia,  the  period  Bitcey  remained  consumption  feeding  Ritcey  (1974) f o u n d  southeastern fir  f i r m enough t o s u p p o r t  12  this  of  moose f e e d i n g  on  arboreal  Des  13  l i c h e n s i n Laurentide Park, Quebec, although l i c h e n s were abundant i n some areas s t u d i e d . Importance of a r b o r e a l l i c h e n s t o Eocky  Mountain  elk  l£ervus canadensis n e l s o n i ) i s apparently low. K u f e l d (1973) summarized the r e s u l t s of 48 food h a b i t s s t u d i e s and only one  (DeNio 1938)  i n which l i c h e n consumption  mentioned  was r e p o r t e d .  Hash (1973) found t h a t a r b o r e a l l i c h e n s made up 2,435 of the winter d i e t and occurred i n 35% of 57 e l k rumens c o l l e c t e d i n northern Idaho. Murie  (1951) d e s c r i b e d a r b o r e a l l i c h e n s as "very  p a l a t a b l e " to E o o s e v e l t e l k (Cervus canadensis r o o s e v e l t i ) the Olympic  on  P e n i n s u l a and noted t h a t they were consumed both  from f a l l e n branches and from trunks of standing t r e e s . Packee (1975) r a t e d use of Usnea spp. by E o o s e v e l t e l k as medium i n winter and s p r i n g , but added t h a t the importance  of l i c h e n s i n  the d i e t of e l k i s p o o r l y understood, A r b o r e a l l i c h e n s are important i n the winter d i e t s of some ungulates i n the S o v i e t Union. Nasimovitch  (1955) r e p o r t e d that  they are a p r i n c i p a l winter food of musk deer moschiferus) throughout  i t s range, and  (Moschus  of moose i n p a r t s of i t s  range. A r b o r e a l l i c h e n s are important secondary of the range of red deer (C. nippon) bonasusl  , roe deer  iCervus elaphus)  , sika  foods i n p a r t s deer  (Capreolus capreolus) . b i s o n (Bison  , and chamois (Rupicapra r u p i c a p r a ) -.  14  j M i i i i i 2 i a l Value of A r b o r e a l Forage Lichens  A v a i l a b l e data on the chemical composition l i c h e n s o f the genera  Alectoria  of a r b o r e a l  Isensu l a t o ) and Usnea are  summarized i n Table I . , I n g e n e r a l , crude p r o t e i n l e v e l s are lower than the 6-7% g e n e r a l l y recommended as minimum maintenance l e v e l s f o r deer  (French e t a l . 1956; D i e t z 1972). P r o t e i n l e v e l s  i n t h e study area are the lowest that are r e p o r t e d , although they might have been higher had R o c h e l l e sampled B r v o r i a spp. as well as A l e c t o r i a sarmentosa. The Ca-P r a t i o reported by Bergerud  (1972) i s f a v o u r a b l e . The d i g e s t i b i l i t y  by B o c h e l l e  (1978) are extremely  values reported  high - 75-85% compared to 20-  50% f o r other major f o r a g e s p e c i e s during winter.  Furthermore,  A l e c t o r i a sarmentosa appears t o have a s y n e r g i s t i c e f f e c t on n e t d i g e s t i b i l i t y when combined with ether forage p l a n t s (J.A. Hochelle  1  unpublished d a t a ) . Seme w r i t e r s have suggested  A l e c t o r i a spp. are a good energy source  that  (Scotter 1965, 1972),  but p r e l i m i n a r y r e s u l t s o f a n a l y s i s of v o l a t i l e f a t t y a c i d s i n A l e c t o r i a sarmentosa i n d i c a t e that the l i c h e n s may be r e l a t i v e l y low i n a v a i l a b l e energy  (Bochelle unpublished d a t a ) . The  n u t r i t i o n a l value of A l e c t o r i a Bochelle  1  (s.1.) i s d i s c u s s e d more f u l l y by  (1978).  Weyerhaeuser Co. L t d . , C e n t r a l i a , Washington.  TABLE I.  N u t r i t i o n a l value of arboreal forage  Crude In V i t r o Protein Dry Matter D i g e s t i b i l i t y [%) (%) Rochelle  1978  Fat (*)•  Fi ber (*)  Ash (%)  NFE (%)  Ca (*)  P (*)  0.56  7.47  1.45  84.74  0.11  0.07  4.6  2.34  4.39  1.73  86.75  A. sarmentosa  2.55  7.05  1.42  0.57  88.42  "A. jubata" June September March  6.26 4.51 3.98  1.00 0.41 0.43  6.62 5.72 5.36  1.08 1.23 1.04  85.04 88.10 89.19  Usnea hirta September March  5.45 4.59  4.87 5.70  7.10 6.93  1.41 1.41  81.17 81.37  A. sarmentosa  Sulkava & Helle 1975 Bergerud 1972  Alectoria  Scotter  1974  spp.  <2 4.2 5.78  A. sarmentosa  3.06  subfloridana  "A. rjubata"  1965  1  2  Usnea Schroeder  75 - 85  jubata"  "A.  Packee 1975  3  9.259.38  h  B l a c k t a i l e d deer; winter Mainly B. trichodes Bryoria  subsp. americana (Brodo & Hawksworth 1977:  spp.  B. lanestris  lichens.  and B. simplicior  (Brodo & Hawksworth 1977:  45).  45).  16  Ecolo<jI o f  A l e c t o r i o i d Lichens  Ecological  studies  must be  taxonomy. U n t i l r e c e n t l y , accepted  classification  to e c o l o g i c a l research. Hawksworth*s the  the  of The  - Alectoria, Bryoria, d o e s not  level  the  are  considered  Alectoria  and  most s p e c i e s  (Brodo and  i n the  discussion  are  opportunities,  and  lichens  and  Hawksworth  time.  by  colour  As  the  Factors  is  follows.  more  abundance range are  genera,  of solar  characteristics, dispersal  that  also  Bryoria  species  1977). Some o f t h e  substrate  These  to s p e c i e s  with  the  in  -  or a s s o c i a t e d  t h e i r geographical  animals are  gryoria,,  at  that  North  Pseudephebe i s  by  Ecology  contribute  s u c h as b r e a k a g e t h r o u g h t h e  consumption  laid  authors  essentially cold climate  either northern  energy, water, n u t r i e n t s ,  These  identification  field.  lichens within  and  y e l l o w - g r e e n , whereas  i n the  Bryoria  are  field  important e c o l o g i c a l f a c t o r s determining  of  and  C o l u m b i a , and  i n the  some c a s e s ,  or impossible  Alectorioid  obstacle  Sulcaria.  g e n e r a A j L e c t o r i a and  gray-green to  brownish. In  mountains  an  A l e c t o r i o i d lichens discussed  easily distinguished  i s not  widely  genera w i t h i n  P s e u d e p h e b e , and  in British  genus, a l l the  spp.  difficult  and  occur  t h e s i s belong to  genera are  and  Brodo  f o r m e r genus A l e c t o r i a i n t o f o u r  a terrestrial  are  of  ecological studies.  America  spp.  workable  (1977) monograph d i s p e l l e d much c o n f u s i o n  the  Alectoria  of a  publication  divided  this  lack  a satisfactory  A l e c t o r i o i d l i c h e n s was  groundwork f o r i m p r o v e d  Sulcaria  b a s e d on  action of  important.  to t h e wind  removal and  rain,  17  The importance o f l i g h t and humidity i n determining the abundance of A l e c t o r i a workers  ( s . l . ) has been pointed out by other  (Schroeder 1974)  and may  be i n f e r r e d from the  d i s t r i b u t i o n of these l i c h e n s i n t r e e s . In g e n e r a l , l i g h t i n t e n s i t y i n c r e a s e s and r e l a t i v e humidity decreases with i n c r e a s i n g d i s t a n c e from the ground i n a f o r e s t stand al,  1955;  (Onura e t  Geiger 1966:298). On the b a s i s of p h y s i o l o g i c a l  s t u d i e s of s e v e r a l s p e c i e s of e p i p h y t i c l i c h e n s and mosses at various i l l u m i n a t i o n s and h u m i d i t i e s , Hosokawa e t a l . (1964) concluded t h a t t h e i r e c o l o g i c a l ranges were r e s t r i c t e d lower l i m i t s by l i g h t  and at the upper  l i m i t s by  at the  water.  In the humid west c o a s t f o r e s t the dominant genus of A l e c t o r i o i d l i c h e n s i s A l e c t o r i a . In lower slope p o s i t i o n s where f o r e s t s are denser, A l e c t o r i a i s found nigh i n the crown. On x e r i c s i t e s where stands are more open, A l e c t o r i a  extends  f u r t h e r down the t r e e and B r y o r i a i s present near the top of the t r e e . Both genera are sparse i n the lower p a r t s of t r e e s except at very open s i t e s . T h i s d i s t r i b u t i o n p a t t e r n , documented by the work of Szczawinski (1953) and Pike et a l . (1975) s t r o n g l y suggests that i n the c o a s t a l f o r e s t , l i g h t i s the dominant c l i m a t i c f a c t o r c o n t r o l l i n g the d i s t r i b u t i o n of .A^lectoria ( s . l . ) , and that the B r y o r i a s p e c i e s of t h i s f o r e s t type are more photophilous than the A l e c t o r i a s p e c i e s . In the i n t e r i o r of B r i t i s h Columbia,  with i t s more  c o n t i n e n t a l c l i m a t e , humidity i s more f r e q u e n t l y a l i m i t i n g f a c t o r than i n the moist, c o a s t a l f o r e s t . Edwards e t a l . (1960) noted that i n some dry f o r e s t s o f S e l l s Gray Park, ( s . l . ) was  Alectoria  present i n the lower p a r t s o f the crowns, but absent  18  in  t h e upper  southeastern Bryoria  British  o f t h e S e l k i r k Range o f  Columbia,  increased  areas,  with e l e v a t i o n  1978). A l t h o u g h  part  other  to increased  evaporation The  that  abundance o f A l e c t o r i a  (Ahti  variables  supply with  and  i s p r o b a b l y due higher  and t h e r e f o r e  lower  limiting  i n water  that  lichens and m i n e r a l  nitrogen,  nitrogen  which a r e l e a c h e d  In the study area,  w i t h good l i g h t  penetration  richer sites  seems u n l i k e l y t h a t  often  nutrienthave much  with a denser  i n t h e crowns o f t r e e s by  tree  n u t r i t i o n i s a major  growth o f A l e c t o r i o i d l i c h e n s , e x c e p t  n o t been e n r i c h e d  present  Rochelle  elevation,  d e p e n d e n t on e x t e r n a l  a s n i t r a t e s and o r g a n i c  canopy. I t t h e r e f o r e  Air  with  the t h a l l u s . C o r t i c o l o u s  l i c h e n abundance t h a n  high  (s.l.)  1974). A l e c t o r i o i d l i c h e n s have g r e e n a l g a e a s  poor f o r e s t s i t e s  has  associated  t h e t r e e c a n o p y by r a i n f a l l .  factor  change w i t h  of lichens i s dissolved  p h y c c b i o n t s , and a r e t h e r e f o r e  very  also  a c c u m u l a t e a wide r a n g e o f o r g a n i c  (Hale  s o u r c e s such  a r e a s as  rates.  comes i n c o n t a c t selectively  humidity,  growth  1962, B a r i c h e l l o 1975,  l o w e r summer t e m p e r a t u r e s ,  nutrient  nutrients  excessive lichen  l i c h e n abundance a t h i g h e r e l e v a t i o n s  precipitation,  higher  Thus,  1974). Many a u t h o r s have n o t e d , i n c o a s t a l  as i n t e r i o r  increases  from  o c c u r r e n c e o f A l e c t o r i a and  c a n make f o r e s t s t o o d r y f o r c o n t i n u e d  (Schroeder  in  I n parts  a p p e a r s t o be l i m i t e d b y h u m i d i t y .  thinning  well  parts.  possibly  where t h r o u g h f a l l p r e c i p i t a t i o n  leaching.  p o l l u t i o n i s t h o u g h t t o be an i m p o r t a n t d i s t r i b u t i o n of s e v e r a l  species  f a c t o r i n the  of Bryoria  occurrence of A l e c t o r i o i d l i c h e n s i n c i t i e s  i n Europe,  and i n d u s t r i a l  19  areas i s l i m i t e d  (Brodo and Hawksworth 1977). Sheridan e t a l .  (1976) found that biomass o f A l e c t o r i a  (Bryoria)  fremontii  was  exremely reduced below c o n t r o l s i t e values as much as 16 km from a pulp m i l l i n Montana. The i n f o r m a t i o n i n d i c a t e s that s p e c i e s  currently  of A l e c t o r i a and B r y o r i a  available  differ in their  s e n s i t i v i t y t o a i r p o l l u t i o n , but i t i s not y e t p o s s i b l e to describe  their relative sensitivities  (Brodo and Hawksworth  1977) . The  Best common s u b s t r a t e s  are c o n i f e r o u s such as b i r c h  of corticolous A l e c t o r i a  (s.l.)  t r e e s and t r e e s with s i m i l a r bark c h a r a c t e r i s t i c s (Brodo and Hawksworth 1977). I t i s not c l e a r  whether these l i c h e n s a c t u a l l y p r e f e r a c i d i c bark due to some p h y s i o l o g i c a l requirements, or whether they r e q u i r e and  general environment o f c o n i f e r o u s  the  most a v a i l a b l e s u b s t r a t e .  the  latter possibility  the c l i m a t e  f o r e s t s and simply occupy  According t o Brodo and Hawksworth,  i s more l i k e l y ,  as a wide v a r i e t y of  t r e e s w i t h i n each f o r e s t type are used as s u b s t r a t e s .  Although  s u b s t r a t e - s p e c i f i c i t y o f A l e c t o r i o i d l i c h e n s i s r e l a t i v e l y lew compared to seme c o r t i c o l o u s l i c h e n s , e s p e c i a l l y s p e c i e s are i n c l o s e r c o n t a c t  with the s u b s t r a t e  (Brodo 1973), some  important d i f f e r e n c e s i n abundance of A l e c t o r i a conifer species  which  is.1.)  cn  are apparent. Bark c h a r a c t e r i s t i c s of some  B r i t i s h Columbia t r e e s and t h e i r s u i t a b i l i t y as h a b i t a t s f o r epiphytes have been d i s c u s s e d (1962). Moisture c a p a c i t y , texture,  by Szczawinski  (1953) and A h t i  mineral content, pH, p h y s i c a l  and s t a b i l i t y are bark c h a r a c t e r i s t i c s p o t e n t i a l l y  important i n determining s u i t a b i l i t y o f a t r e e as a (Brodo 1973).  substrate  20  The establishment  of a l i c h e n s p e c i e s i n an area depends on  the a v a i l a b i l i t y of v i a b l e propagules, e i t h e r s e x u a l or vegetative. Alectoria  Spore-producing s t r u c t u r e s are r a t h e r r a r e among  ( s . l . ) and most s p e c i e s  methods of propagation reproduction  depend mainly on v e g e t a t i v e  (Brodo and Hawksworth  1977) .  Vegetative  by windblown t h a l l u s fragments i s o f major  importance i n short-range d i s p e r s a l , but there i s some evidence that s p e c i e s  of A l e c t o r i a and B r y o r i a that produce s o r e d i a -  clumps of a l g a l c e l l s surrounded by f u n g a l hyphae - have an advantage i n long-range d i s p e r s a l over s p e c i e s t h a t do not (Stevenson unpublished d a t a ) . Fragmentation and i s o l a t i o n o f o l d f o r e s t stands by human settlement, severely  a g r i c u l t u r e , and logging  r e s t r i c t the d i s t r i b u t i o n of some l i c h e n s p e c i e s  cannot d i s p e r s e e f f e c t i v e l y over long d i s t a n c e s .  Rose  may  that  (1976)  found t h a t some a r b o r e a l l i c h e n s i n Great B r i t a i n , i n c l u d i n g several species  of A l e c t o r i a ( s . l . ) . a r e p r e s e n t l y  a n c i e n t , undisturbed  f o r e s t s and are apparently  l i m i t e d to  unable t o  c o l o n i z e s u i t a b l e h a b i t a t s at a d i s t a n c e . Brodo and Hawksworth (1977) suggested that t h i s phenomenon may be an important  aspect  of the l o c a l d i s t r i b u t i o n o f some s p e c i e s i n North America. Studies  o f e f f e c t i v e d i s p e r s a l d i s t a n c e s of l i c h e n s a r e needed,  as they have i m p l i c a t i o n s f o r f o r e s t management, p a r t i c u l a r l y with r e s p e c t to s i z e and spacing  of c l e a r c u t s .  The abundance of A l e c t o r i a ( s . l . ) i n a s u i t a b l e f o r e s t s i t e depends on how much time has been a v a i l a b l e f o r establishment and  growth o f t h e l i c h e n s . Growth r a t e s of l i c h e n s are g e n e r a l l y  slew compared t o other  p l a n t groups. There are many t e c h n i c a l 5  d i f f i c u l t i e s i n v o l v e d i n measuring growth r a t e s of f r u t i c o s e  21  a r b o r e a l l i c h e n s , and consequently l i t t l e  useful information i s  a v a i l a b l e on growth r a t e s of a l e c t o r i o i d l i c h e n s . The great abundance of a l e c t o r i a  relatively  ( s . l . ) i n mature f o r e s t stands as  opposed to immature stands has been noted by many authors (Edwards et a l .  1960, A h t i  1962, A h t i and Hepburn  1967). There  i s a need f o r r e s e a r c h on the dynamics of l i c h e n development i n the course of f o r e s t s u c c e s s i o n , and a s s o c i a t e d changes i n l i c h e n bicmass.  22  III.  QUANTIFICATION OF  In  order to study l i c h e n  ecosystem  f a c t o r s , i t was  measuring  lichen  and  BIOMASS OF  forage  p r o d u c t i o n and  necessary  abundance. The  evaluate techniques  ARBOREAL FORAGE LICHENS  to  first  other  have methods f o r o b j e c t i v e was  to  develop  of q u a n t i f y i n g biomass o f a r b o r e a l  lichens.  T h r e e methods o f a s s e s s i n g l i c h e n bicasass was A system related  measured by  sampling  of v i s u a l estimates to  extended  t h e l i c h e n s from  of l i c h e n  to  o t h e r a r e a s a s an a i r p h o t o g r a p h y as  l i c h e n s of the genera  Alectoria  Many t e c h n i c a l and  Pike et a l .  1972,  i n the  The  trees.  developed,  sampling, value  limited  was  to arboreal  p r o b l e m s are on  a s s o c i a t e d with an  areal  basis.  crown o f a l a r g e t r e e i s  the t r e e r e q u i r e s e l a b o r a t e e q u i p m e n t  (e.g.  1977), and f e l l i n g t h e t r e e d e s t r o y s and  s c a t t e r s some o f t h e e p i p h y t e s . I n t e n s i v e b i o m a s s s a m p l i n g p r o d u c e a good e s t i m a t e but  of e p i p h y t e  i s too time-consuminq  Rapid  estimates  purposes actual  t o be  of epiphyte  biomass.  used  quantities i n a small for inventory  can area,  purposes.  abundance s u i t a b l e f o r i n v e n t o r y  a r e o f t e n i m p r e c i s e and  epiphyte  and  of  inventory tool  biomass of e p i p h y t e s  to epiphytes growing  was  Lichen  Bryoria*  statistical  to q u a n t i f y the  climbing  and  was  felled  intensive  a potential  quantification  difficult;  abundance  inventory tool.  t e s t e d . In each c a s e ,  attempts  b i o m a s s were used.  b i o m a s s e s t i m a t e s o b t a i n e d by  large-scale  Access  r e l a t e i t to  are d i f f i c u l t t o r e l a t e  to  23  Some i n v e s t i g a t o r s (Scotter 1962, Andre e t a l . 1975, Hein and  Speer 1975) have determined l i c h e n biomass by removing and  weighing the e n t i r e l i c h e n l o a d from one or more t r e e s w i t h i n a f o r e s t type.  These authors s t u d i e d f a i r l y s m a l l t r e e s  (25 m or  l e s s ) ; the method would be i n e f f i c i e n t i f t r e e s were l a r g e and l i c h e n guantiti.es great.  Various  subsampling schemes have been  used to study l a r g e r t r e e s . Edwards e t a l . (1960) counted the branches i n each 3 m height representative  i n t e r v a l , s e l e c t e d two  branches f o r sampling, and c a l c u l a t e d biomass  w i t h i n each height  i n t e r v a l by m u l t i p l i c a t i o n . Forman  (1975)  s e l e c t e d a r e p r e s e n t a t i v e o n e - t h i r d of the crown of l a r g e  trees  f o r sampling and m u l t i p l i e d the r e s u l t s by t h r e e . Pike e t a l . (1972, 1977) developed a very  i n t e n s i v e method o f c a l c u l a t i n g  t o t a l epiphyte biomass on a l a r g e (Pseudctsuqa menziesii)  (65 m) s t a n d i n g  Douglas-fir  t r e e . Each branch was assigned an  "importance value" on the b a s i s of d e t a i l e d o b s e r v a t i o n s  of i t s  s i z e and epiphyte biomass, which were made from a c l i m b i n g p o s i t i o n on the trunk of the t r e e . Importance values as t h e b a s i s f o r a random p r o p o r t i o n a t e  scheme o f sampling  branches f o r d e t a i l e d study. These values extrapolate  were used  were l a t e r used t o  epiphyte biomass from sample branches t o the e n t i r e  t r e e by means of a l i n e a r r e g r e s s i o n of biomass on epiphyte importance  value.  Most r e s e a r c h e r s  (Edwards e t a l . 1960, S c o t t e r  1962, Andre  et a l . 1975) have extended t h e i r biomass t o t a l s f o r i n d i v i d u a l t r e e s t o an a r e a l b a s i s by m u l t i p l y i n g by t r e e s per u n i t Unless the f o r e s t i s q u i t e uniform or the sample s i z e  area.  very  l a r g e , t h i s method could be expected t o l e a d t o e r r o r s . Forman  24  (1975) f o u n d a s i g n i f i c a n t r e l a t i o n s h i p between guantities quantity  and t r e e  diameter,  often  o f forage l i c h e n s  been r e p o r t e d ,  1975). A n t i  (1962) u s e d  which c l o s e l y  1975, J o n e s  lichen  cover,  a n d numbers o f  method  gave  p a r a l l e l e d the general impressions of  Russell  1  and S. R u s s e l l  (pers.  i n his  field  interior  British  Columbia  dividing  the t r e e  section,  and e s t i m a t i n g  notes.  comm.) d e v e l o p e d  l i c h e n abundance on t h e l o w e r p a r t s  branches. N e i t h e r  of lichen  i n H e l l s Gray P a r k . T h i s  abundance r e c o r d e d by A h t i  estimating  b a s i s has  a " r a n g e i n d e x " o f l i c h e n abundance on  lichen  lichen  (Barichello  estimates o f percent  lower 3 m e t e r s o f t r e e s  results  trees.  ratings  o f abundance o f t w i g s on a 1-5 s c a l e ,  stems/acre t o c a l c u l a t e the  of epiphyte  on an i n v e n t o r y  other than simple  abundance c n a low-raedium-high s c a l e  ratings  regressions  on DBH t o c a l c u l a t e t o t a l s f o r unsampled  Quantification not  and u s e d  epiphyte  a system o f of trees i n  f o r e s t s which i n v o l v e d  into sections, percent  counting c o v e r on  D.  visually  branches i n each representative  c f these systems i s s u i t a b l e f o r  abundance on t h e e n t i r e crown o f t r e e s  estimating  i n t h e coast  forest. A literature inventory although air  arboreal  no p r e v i o u s a t t e m p t s t o  l i c h e n a b u n d a n c e by means o f remote  terrestrial  photographs  research,  search revealed  sensing,  l i c h e n c o m m u n i t i e s have been mapped  (Buttrick  frcm  1978). D e s p i t e t h e a b s e n c e o f p r e v i o u s  i t seemed r e a s o n a b l e t o e x p e c t  that  differences  i n the  a p p e a r a n c e o f t h e f o r e s t c a n o p y due t o v a r i a t i o n s i n l i c h e n  1  Yukon Game B r a n c h , Box 2703, S h i t e h o r s e ,  Yukon T e r r i t o r i e s , ,  25  abundance might Since the used  be d e t e c t a b l e i n l a r g e - s c a l e initiation  quantify  lichen  study area.  He  used  abundance on  23  sites,  which I q u a n t i f i e d  lichen  When d a t a were t r a n s f o r m e d predictions  0.41  present study,  normal c o l o u r o b l i q u e a i r photographs  abuncance i n the  study  of the  was  on l i c h e n  (Harestad  1978) .  to l o g  | 0  to a s s e s s  a dot g r i d  (1978)  lichen  technique  including  9 plots  to on  measurements.  # a r e g r e s s i o n of a i r photo  a s measured on t h e  significant  photographs. Harestad  abundance u s i n g ground  bit-mass  statistically  air  (p < 0.06)  ground i n  and  had  an r  this z  of  26  Methods  V i s u a l estimates  of l i c h e n abundance  Q u a n t i f i c a t i o n of l i c h e n abundance by v i s u a l estimates conducted on 14 p l o t s . Procedures f o r l a y i n g out  p l o t s and  s e l e c t i n g t r e e s f o r l i c h e n estimates v a r i e d s l i g h t l y on whether or not the p l o t was of  used f o r t e s t i n g  was  depending  photo-prediction  l i c h e n abundance. Nine non-photo p l o t s were chosen to  represent areas f o r which i n f o r m a t i o n on winter deer use or lichen l i t t e r f a l l Harestad  was  a v a i l a b l e (Jones  1978). These p l o t s were sguare and  ha i n area, but a f t e r completing s i z e to 0.2 All  1975,  Rochelle  1978,  were i n i t i a l l y  three p l o t s , I i n c r e a s e d  ha t o improve r e p r e s e n t a t i o n of the f o r e s t  t r e e s w i t h i n the p l o t which were 18 cm  or more i n  numbered p l a s t i c cards, and the f o l l o w i n g i n f o r m a t i o n  was  f o r each t r e e :  - DBH  (diameter  at 1.5  m above the p o i n t of  germination); - species; - dominance c l a s s or  (dominant, codominant, i n t e r m e d i a t e  overtopped);  - l i v i n g or dead  (dead t r e e s 10 m or more i n height  were f u r t h e r c l a s s i f i e d as dead (i.e.  potential  merchantable) or dead u s e l e s s (B.C.  S e r v i c e 1973); and  plot  type.  diameter were i n v e n t o r i e d . Trees were t e m p o r a r i l y l a b e l l e d  recorded  0.1  Forest  with  27  - lean  (this  was  approximately  noted  10  i f trees  degrees from  leaned  more  than  vertical).  i  then  Frequencies of i n d i v i d u a l species-dominance c l a s s e s  were  compiled  using  and  15  random p r o p o r t i o n a t e  trees  selected for lichen estimates  scheme  (Cochran  c h a n c e o f any  particular tree  proportionate  to  class.  the  from t h e  In t h e were l a i d  out  the  within  p h o t o g r a p h s , and  inside  the  the  later  In the  and  and  area  of  located  on  the  was  were  the  boundaries  cn  the  ground. A i r photo  plots  topographic  a l l living  t r e e s having  a i r photograph  were  sampling  scheme. A f t e r p l o t b o u n d a r i e s was  the  random  made from the  and  were  proportionate  were l a i d  c a r r i e d out  bases  identified  a dominance c l a s s . F i f t e e n t r e e s  for l i c h e n estimates using  ground, a t r e e i n v e n t o r y  overlap  shape b e c a u s e o f  selected  identifications  sample  trees  plots, plot  stereo  laboratory,  assigned  overtopped  a i r photo  p l o t b o u n d a r i e s on  to s p e c i e s  i n the  process.  five  were v a r i a b l e i n s i z e and irregularities.  trees,  selection  case of  included  the  number o f t r e e s i n i t s s p e c i e s - d o m i n a n c e  Dead t r e e s , l e a n i n g  omitted  being  1963:89), s u c h t h a t  a  out  on  the  species  p h o t o g r a p h were c o r r e c t e d  where  necessary. Standard  f o r e s t m e n s u r a t i o n t e c h n i q u e s were used on a l l  plots  to o b t a i n  width  (Ca^  ) of t r e e s  estimates.  part  heiqht  highest  of the  (TH),  which had  Crown l e n g t h  meters, from the living  tree  was  crown l e n g t h  been s e l e c t e d  defined  living  as  the  diameter, i n meters, of  the  crown  at the  crown  for lichen  vertical  p a r t of the  c r o w n . Crown w i d t h was  ( C L ) , and  distance,  crown t o the  defined  as  widest part,  the and  in  lowest average was  28  determined slope  by a v e r a g i n g  and p a r a l l e l  measurements t a k e n  to the slope  Lichen estimates  (fig.  were made f r o m  perpendicular to the  2).  a viewing  position  which a s much a s p o s s i b l e o f t h e crown c o u l d be s e e n . it  was n e c e s s a r y  portion four  to s h i f t  of the tree.  layers  each l a y e r  t o a second  The l i v e  viewing  crown was v i s u a l l y  (CW» ) :  The crown w i d t h  p o i n t f o r e s t i m a t i n g t h e width  i n meters  - % t r e e cover cover  (TCj):  dimensions  included  were made f o r  trunk,  epiphytes —  as a o f each  (i =  was e s t i m a t e d  having  into  of t h e t r e e ,  which had a l r e a d y been measured, was u s e d  layer,  divided  the a i dof binoculars;  - crown w i d t h  reference  Sometimes,  p o i n t t o see a  ( F i g . 3 ) , and t h e f o l l o w i n g e s t i m a t e s with  from  For each l a y e r ,  percent  w i t h i n an imaginary  rectangle  CWj x CL/4 ( F i g . 3 ) . T r e e  branches,  anything  tree  cover  t w i g s , f o l i a g e , and  w h i c h was p a r t o f t h e t r e e o r  which grew on i t . - % Alectoria consisting each  To  ( A | ) : The p e r c e n t  of Alectoria  a n d B r y o r i a was e s t i m a t e d f o r  assess interobserver r e l i a b i l i t y ,  i n Plot  visual  1969:114) u s e d significantly  a field  estimates of lichen  a s s i s t a n t and  abundance on t h e  1. A r e g r e s s i o n o f one s e t o f o b s e r v a t i o n s on t h e  o t h e r was c a l c u l a t e d ,  from one.  of t r e e cover  layer.  I made i n d e p e n d e n t trees  (s.l.)  and S t u d e n t ' s  t o determine from  t-test  (Dixon a n d Massey  whether t h e i n t e r c e p t  zero, or the s l o p e d i f f e r e d  differed  significantly  29  Figure 3.  Division of a tree into layers for visual estimates of lichen abundance.  30  During the f i r s t summer, a l l t h r e e e s t i m a t e s d e s c r i b e d  here  were made on a l l t r e e s s t u d i e d . During the second summer, only A l e c t o r i a was estimated. The reason i s given i n the d i s c u s s i o n of  Air  computations and r e s u l t s .  photo q u a n t i f i c a t i o n of l i c h e n abundance Eefore f l y i n g the area, I photographed l i c h e n s using  various combinations of f i l m s and f i l t e r s .  C o n t r a s t between  c o n i f e r f o l i a g e and l i c h e n s was best with c o l o u r - i n f r a r e d a Wratten 12 f i l t e r , and two CC20 Magenta f i l t e r s A e r i a l photography  film,  ( F i g . 4) .  was done by I n t e g r a t e d Resources  Photography, L t d . of Vancouver on June 5 and 6, 1976, using Kodak Aercchrcme I n f r a r e d Film 2443, a Wratten 12 f i l t e r , and two CC20 Magenta f i l t e r s .  Two Vinten cameras mounted on the  w i n g - t i p s of a Cessna 180 were exposed s i m u l t a n e o u s l y t o produce s t e r e o images. A l l areas were flown a t a height of approximately 152 m above the tops of the t r e e s t o o b t a i n imagery a t the s c a l e 1:2000; f o r some f l i g h t  l i n e s 1;4000 and 1;8000 photographs were  also obtained. The use of a i r photographs t o d e t e c t l i c h e n abundance was t e s t e d by two methods: densitometry and photo i n t e r p r e t a t i o n . Densitometry i s a technigue that may be used t o q u a n t i f y dye-layer d e n s i t y i n a c o l o u r photograph. C o l o u r - i n f r a r e d p o s i t i v e t r a n s p a r e n c i e s c o n s i s t of three developed dye-layers and a s t a b l e base. The d e n s i t i e s c f these d y e - l a y e r s are i n v e r s e l y r e l a t e d to the s p e c t r a l r e f l e c t a n c e p a t t e r n of the s u b j e c t photographed, although other f a c t o r s i n f l u e n c e dye-layer d e n s i t y as well  (Murtha 1972). In c o l o u r - i n f r a r e d f i l m ,  spectral  A  Figure 4.  Alectoria  savmentosa on D o u g l a s - f i r :  normal colour f i l m (A) and colour i n f r a r e d f i l m with  Wratten 12 f i l t e r and two CC20 Magenta f i l t e r s (B).  32  reflectance the  i n the near-infrared  cyan dye-forming  filters  o f each  spectrum  needles r e f l e c t (Steiner  t h e d y e s used  subtractive,  visually  sarmentosa  reflectance  pattern  i s developed  lichens  with Wratten f i l t e r s  (Fig. 4 ) . In light.  92  densities  into classes  pattern  the  two c l a s s e s The  ability  result i n a  Densitometer  (green),  and 94  was  (blue) t o plots f o r  The sample t r e e s  were  and,low l i c h e n b i o m a s s a s by t h e mean. S t u d e n t s t  1969:114) was used t o compare t h e mean  dye-layer,  and t h e r e d : g r e e n f i l t e r r a t i o , o f  f o r a l l trees  together  o f photo i n t e r p r e t e r s  with v a r y i n g  1966, G a t e s  o f t h e t r e e s i n t h e photo  abundance on i n d i v i d u a l t r e e s subjects  would  Reflection  p l o t s , separated  (Dixon and Massey  reported f o r  and Gutermann  ( r e d ) , 93  of high  from ground  o f each  contrast,  Data on t h e s p e c t r a l  which l i c h e n b i o m a s s had been c a l c u l a t e d .  determined  f i l m are  film.  A Macbeth TR-524 T r a n s m i s s i o n  measure d y e - l a y e r  i n the infrared region of  needles appear r e d t o  (Steiner  image on c o l o u r - i n f r a r e d  density  separately i n  a t a l l w a v e l e n g t h s h a s been  1970). Such a s p e c t r a l r e f l e c t a n c e  test  coloured  of A l e c t o r i a sarmentosa a r e not a v a i l a b l e ,  high reflectance  divided  with  1966, G a t e s 1970, H u r t h a  conifer  appears very  other l i g h t - c o l o u r e d  used  dye l a y e r  i n colour-infrared  green  magenta when t h e f i l m Alectoria  strongly  and Gutermann  1972). S i n c e  light  i s used  affects  transparencies.  Conifer  but  o f the spectrum  A densitometer  t o measure t h e d e n s i t y  positive  the  layer.  region  and f o r s e p a r a t e to p r e d i c t  and on s i t e s  lichen ecclogy  lichen  was t e s t e d . Ten  degrees o f experience with  i n t e r p r e t a t i o n and with  species.  were u s e d .  photo  33  The infrared  i n t e r p r e t e r s were shown normal c o l o u r and  photographs, taken from the ground, of A l e c t o r i a -  covered B o u g l a s - f i r stereo  p a i r s , cne one  trees  ( F i g . 4 ) . They were then shown  representing representing  a p l o t with low  (Fig.  5) and  (Fig.  6), i n which a t o t a l of 34  to l i c h e n abundance. The ones on  colour-  l i c h e n biomass  a p l o t with high l i c h e n biomass t r e e s were l a b e l l e d a c c o r d i n g  t r e e s l a b e l l e d i n the  photographs were  which biomass sampling or v i s u a l e s t i m a t e s had  c a r r i e d out.  They were r a t e d  two  on a 6-point s c a l e T  been  (trace)  through 5 (abundant), corresponding broadly t o k i l o g r a m s of l i c h e n on the t r e e s . I n t e r p r e t e r s a d d i t i o n a l stereo c i r c l e d , and using for  were then shown three  p a i r s of ground p l o t s , each with 15  trees  asked to r a t e l i c h e n abundance on those t r e e s ,  the l a b e l l e d s e t s f o r r e f e r e n c e .  which v i s u a l estimates had  C i r c l e d t r e e s were ones  been made. F i n a l l y , 9 of the  10  i n t e r p r e t e r s were asked t o rate g e n e r a l l i c h e n abundance on eleven stereo using  p a i r s of ground p l o t s as high,, medium, or  four l a b e l l e d photo p a i r s f o r  low,  reference.  In a d d i t i o n to sampling other photo i n t e r p r e t e r s , I made p r e d i c t i o n s of l i c h e n abundance on t r e e s i n photo p l o t s for  v i s u a l estimates before l o c k i n g at the  did  not  make p r e d i c t i o n s  selected  trees i n the f i e l d .  f o r s i t e s because I had  field  I  knowledge  of t h e i r l i c h e n abundance. To determine whether i n t e r p r e t e r r a t i n g s of abundance on i n d i v i d u a l t r e e s were c o r r e l a t e d l i c h e n biomass, Spearman's rank c o r r e l a t o n (Conover 1971:245). Spearman's rho r a t i n g s of each i n t e r p r e t e r on  lichen  with c a l c u l a t e d  c o e f f i c i e n t was  or " r " was  used  calculated for  each of the three p l o t s and  the  used  Figure 5.  Colour i n f r a r e d a i r photographs of p l o t with low l i c h e n biomass.  35  36  to t e s t  for positive  interpreters test  1  (Concver  correlation  of ratings  r h o s c o r e s were u s e d as d a t a  Subjects  o r by f i e l d  f o r t h e Mann-8hitney  b l a c k and w h i t e ,  photo  knowledge o f A l e c t o r i a  who had p r e v i o u s e x p e r i e n c e  with  interpretation  (s.l.)  ecology.  interpretation of  normal c o l o u r , and c o l o u r i n f r a r e d a i r  p h o t o g r a p h s were c o n s i d e r e d t o have a h i g h interpretation  biomass. I h e  1971:224) t o d e t e r m i n e w h e t h e r , a b i l i t y t o j u d g e  l i c h e n abundance was i n f l u e n c e d by e i t h e r experience  with  with  only  n o r m a l c o l o u r and b l a c k and w h i t e a i r p h o t o g r a p h s , o r l e s s ,  were  considered with  experience;  l e v e l of p h o t o  inexperienced  me i n t h e s t u d y  s u b j e c t s who had e x p e r i e n c e  interpreters.  area during the f i e l d  t o have knowledge o f A l e c t o r i a For lichen  statistical  Subjects  analysis,  work were  (s.l.) ecology; interpreters'  as 1, 2, o r 3. The mean r a t i n g  calculated  and Spearman's r a n k c o r r e l a t i o n  t o d e t e r m i n e whether mean r a t i n g s calculated  Biomass  considered  others  were c o t .  judgements o f  abundance on s i t e s a s l o w , medium, o r h i g h  guantified  were  f o r each s i t e  was  c o e f f i c i e n t was used  f o r s i t e s were c o r r e l a t e d  with  l i c h e n biomass t c t a l s f o r s i t e s .  sampling  Biomass s a m p l i n g fieldwork and  who had been  the f i r s t  abundance. I n e a c h p h o t o p l o t ,  t o be f e l l e d  chosen from t h o s e were s e l e c t e d dominance  out during  i n two photo p l o t s , c h o s e n t o r e p r e s e n t a r e a s  low l i c h e n  selected  was c a r r i e d  f o r biomass sampling.  f o r which v i s u a l e s t i m a t e s  to represent  classes.  summer o f of high  f i v e t r e e s were  The t r e e s were had been made, and  t h e major l i c h e n - b e a r i n g , s p e c i e s -  37  The f o u r compass d i r e c t i o n s were s p r a y - p a i n t e d on t h e h o l e s of  the t e n t r e e s . They were then f e l l e d by a p r o f e s s i o n a l  f a l l e r . The t h r e e D o u g l a s - f i r broke up b a d l y when they f e l l , but the cedars and t h e hemlock were r e a s o n a b l y i n t a c t .  Different  sampling methods were used i n the two c a s e s . The branch was t h e b a s i c s a m p l i n g u n i t f o r i n t a c t  trees.  Branches cn the a c c e s s i b l e upper 180 degrees of the f a l l e n t r e e s were sampled; l i c h e n abundance on t h e lower s i d e s of t h e t r e e s was assumed e g u i v a l e n t t o t h a t on t h e upper s i d e s . A l l branches g r e a t e r than 0.5 meters l o n g p r o j e c t i n g from the upper 180 degrees o f t h e t r u n k were numbered and t h e f o l l o w i n g data recorded f o r each  branch:  - l a y e r i n which branch o c c u r r e d ; - branch l e n g t h ( B L ) ; - density factor  (DF) (a s u b j e c t i v e e s t i m a t e o f branch  s u r f a c e a r e a , independent  of branch l e n g t h , r a t e d on a  1 to 5 scale) ; - e s t i m a t e d percent A l e c t o r i a s u r f a c e area  ( s . l . ) cover of branch  (%A); and  - average l e n g t h o f A l e c t o r i a  ( s . l . ) clumps (LA).  These data were used i n t h e f i e l d t o c a l c u l a t e a L i c h e n Estimate  (LE) f o r each branch, u s i n g t h e f o r m u l a : LE = BL x DF x 3SA/10 0 x LA  (1)  The l i c h e n E s t i m a t e s were then used as t h e b a s i s f o r a randcm p r o p o r t i o n a t e s a m p l i n g scheme. Twenty t o t h i r t y branches  frcm  each t r e e were randomly s e l e c t e d f o r biomass s a m p l i n g , such t h a t the p r o b a b i l i t y o f any g i v e n branch b e i n g s e l e c t e d was p r o p o r t i o n a t e t o i t s L i c h e n E s t i m a t e . T h i s sampling scheme, l i k e  38  that  proposed by Pike e t a l . (1972, 1977), ensured  with high l i c h e n biomass were sampled  t h a t branches  intensively.  Lichen biomass was sampled on the trunks of f o u r of these t r e e s . A l l A l e c t o r i a and B r y o r i a was c o l l e c t e d from 10 x 30 cm guadrats  at 3 m i n t e r v a l s along the s i d e s o f the trunk  p r e v i o u s l y l a b e l l e d as n o r t h - f a c i n g and s o u t h - f a c i n g . Some adjustment i n the aspect of the sample l i n e was necessary  when  the north or south s i d e was f a c i n g d i r e c t l y down. T h i s p a r t i c u l a r sampling  scheme was part of a separate study o f  v e r t i c a l d i s t r i b u t i o n of epiphytes on the study t r e e s . Lichen l i t t e r from the immediate area of the f a l l e n  trees  was c o l l e c t e d s e p a r a t e l y f o r each l a y e r of the crown. The  three D o u g l a s - f i r were so badly broken up that branches  could not be used as sampling  u n i t s . Two of these t r e e s were  sampled by running e i g h t one-meter-wide t r a n s e c t s a c r o s s the area occupied  by f a l l e n t r e e s , p e r p e n d i c u l a r to the trunk. A l l  A l e c t o r i o i d l i c h e n s were c o l l e c t e d from w i t h i n these t r a n s e c t s , i  r e g a r d l e s s of whether they o c c u r r e d on branches or on the ground. Lichens c o l l e c t e d from the trunk were bagged s e p a r a t e l y . The  t h i r d D o u g l a s - f i r had a very s m a l l crown and a l l the l i c h e n s  were c o l l e c t e d from the area surrounding The  tree.  sample t r e e s were aged by counting annual  stump h e i g h t , approximately  r i n g s at  0.6 m.  Lichens were hand-picked, and  this  bagged and l a b e l l e d i n the f i e l d ,  d r i e d i n a hose dryer f o r s t o r a g e . In the l a b o r a t o r y , they  were cleaned of d e b r i s , even-dried at 60-65°C f o r 24 hours, and weighed with a M e t t l e r P1210 e l e c t r i c s c a l e t o t h e nearest 0.01 g.  39  Computations and  Computing  Results  biomass  totals  Hand-picking the A l e c t o r i a and B r y o r i a from the t e n sample t r e e s r e p r e s e n t e d two seeks o f work f o r a crew o f f o u r . Time r e g u i r e d f o r c l e a n i n g , d r y i n g , and weighing was weights f o r the sample  54 hours. L i c h e n  branches were then e x t r a p o l a t e d to a r r i v e  at biomass t o t a l s f o r t r e e s and f o r s i t e s . D e s c r i p t i v e data f o r branches  (BL, DF, %&,  LA) were used t o  estimate l i c h e n weights f o r branches that had not been  sampled.  A r e g r e s s i o n of l i c h e n weights f o r the sample branches of a l l t r e e s on L i c h e n Estimates (Equation  1) gave an r  2  value of  C.61.  To o b t a i n the best p o s s i b l e l i c h e n weight e s t i m a t e s f o r branches that were not sampled, s e v e r a l d i f f e r e n t r e g r e s s i o n equations were evaluated f o r each t r e e , and the eguation y i e l d i n g the hiqhest r unsampled  2  value was used to c a l c u l a t e l i c h e n bicmass f o r branches on that t r e e . In most c a s e s , the best  r e q r e s s i o n equation was o f the form y =  b  0  +  (bjX,)  •  (biX ) z  •  (b^Xj)  where x, throuqh x represent BL, DF, %h, 0  lh x ) H  H  + fa, (x, x x x ) 2  3  (2)  LA, but i n one case  4  y = b  *  (3)  v  accounted f o r more of the v a r i a t i o n i n weights. In g e n e r a l , r  2  values i n c r e a s e d with the order i n which t r e e s were d e s c r i b e d (namely, 0.34, that my a b i l i t y practice.  0.30,  0.59,  0.53,  0.74,  0.65,  0.69), s u g g e s t i n g  to make c o n s i s t e n t e s t i m a t e s improved with  40  T o t a l l i c h e n biomass f o r each l a y e r of each t r e e obtained two  was  by summing l i c h e n weights f o r branches, m u l t i p l y i n g  to account f o r the lower, unsampled s i d e of the t r e e ,  adding the weight of the l i t t e r  associated  by  and  with t h a t l a y e r .  Layer weights were summed to obtain t o t a l s f o r t r e e s . Trunk t o t a l s were c a l c u l a t e d and with high  added to the t r e e t o t a l s . For  l i c h e n l o a d s , l i c h e n biomass on the trunk  c o n s i s t e n t l y 8% of the t o t a l l i c h e n biomass, but lower l i c h e n loads, percent higher  and  trees  was  f o r t r e e s with  l i c h e n biomass of the trunk  was  more v a r i a b l e (Table I I ) . To estimate trunk biomass  f o r t r e e s from which no trunk samples had t o t a l l i c h e n biomass was of l i c h e n , and  been taken, 8%  of  added f o r t r e e s having more than 2C00 g  171 f o r t r e e s having l e s s that 2000 g. A l l  biomass t o t a l s and  percentages o c c u r r i n g on the trunks of  sampled t r e e s are summarized i n Table I I . Ages of the sampled t r e e s , i n c l u d e d i n T a b l e I I , showed little 400  v a r i a t i o n . Seven of the ten t r e e s were between 350  years o l d . To o b t a i n l i c h e n bio mass t o t a l s f o r each s i t e , i t  necessary to e x t r a p o l a t e using  to t r e e s that had  to determine which had  p r e d i c t i v e c a p a c i t y . Lichen  (CL/4),  1,...,4), percent (A^)  crown length  crown width f o r l a y e r s t r e e cover  the  regression  greatest  weight f o r l a y e r s of sampled  the dependent v a r i a b l e and  for layers  was  not been sampled,  v i s u a l estimates of l i c h e n abundance. S e v e r a l  models were evaluated  was  and  (TCj) , and  (CL)  (CH^;  percent  trees  or crown l e n g t h where i = Alectoria (s.l.)  were the independent v a r i a b l e s . A s i n g l e eguation was  used  f o r a l l t r e e s , r e g a r d l e s s of s p e c i e s or dominance c l a s s because  41  TABLE Biomass of Alectoria  II. on sampled trees  Tree  Species  Dominance Class  Age  1  112  WH  D  370  1  13  WH  C  350  1  22  WH  I  1  119  WRC  1  101  2  Alectoria Biomass (g)  % of Biomass Occurring on Trunk  1  5693  8%  1  7036  a  350  1851  a  C  260  436  15%  DF  C  395  1507  14%  5  WH  D  275  4786  8%  2  13  WH  C  181  621  a  2  49  WRC  C  360  1035  22%  2  61  DF  D  394  3233  8%  2  50  DF  D  395  1809  a  Plot  Symbols used i n Table: WH WRC DF AF D C I a  -  western hemlock western red cedar Douglas-fir amabilis f i r dominant codominant intermediate not sampled  Center of trunk r o t t e n ; age estimated.  42  the  sample s i z e  was  too  small to  warrant separate equations.  The  regression y =  81.617 + (n =  had  an  r  expect  a  v a l u e of  2  problem  of  the  and  a  positive  il§£ifiria and  0.75.  (s.l.)  used t o  lichen  had  improve the the  did  not  the  slope  between a measure  was  (A i x  CL)  for  to cf  s u b s t r a t e . To  regression  r  of  2  the  second  8.  158. 03  S  Y  >  X  lichen  estimates  =  376.89; r  biomass of  the  The  of CW  avoid  trees  forced  the  with  through  no  zero,  r  2  test of  and  not  systematically  that  high  biomass f o r  field  for  TC  did  or  0  (t =  class  on  set  of  7.  significantly omitted  0.85.  was  1.07;  to  weights f o r computed. was  each  measurements, t o t a l  1  t  0  os  the 15  Alectoria  shown  intercept 2.10),  1.22;  t  0 0 5  and =  was  other.  t r e e s on  calculate  Using  are  - =  (t =  Mean l i c h e n  used t o plot.  The  observations  in relation  lichen  sites  not  visual  Figure  reliability  r e g r e s s i o n was  neither low  which  t h e y were  interobserver  each s p e c i e s - d o m i n a n c e c l a s s that  trees  d i f f e r s i g n i f i c a n t l y from  Using c a l c u l a t e d  biomass f o r  0.75)  tested,  d i f f e r s i g n i f i c a n t l y from did  =  2  fieldwork.  of  the  (5)  5 is illustrated in  regressions  summer c f  2.10), i n d i c a t i n g  from  376.57)  biomass p r e d i c t i o n s  been made. E g u a t i o n  B e s u l t s of Figure  40;  calculate  B e c a u s e the  lichen  (4)  eguation  estimates  in  =  X  x CL)  measure c f a v a i l a b l e  cover, the  (n =  during  Y  relationship  Y =  was  S .  (A£  B i o l o g i c a l l y , i t i s reasonable  multiplicative  percent cover  40;  148.58  plot  each  plot,  biomass total  areas  for  lichen  computed  ( s . l . ) biomass  (kg/ha)  4000-n  3000H  rt 2 0 0 0 -  —  cn  1000-  l 4  PERCENT  Figure 7.  1 — i 6  1  ALECTORIA  i  8  X  i  r  1©  12  CROWN  14  LENGTH  Regression used to c a l c u l a t e lichen biomass of visual estimate trees (y =• 158.03 x; n = 4.0; S . y  r  2  - 0.75).  x  = 376.89;,  44  Figure 8.  Lichen biomass per tree based on independent v i s u a l estimates by two observers (y = 0.379 + 0.89 x; n = 19; S . ' = 733.51; r y  x  2  = 0.85).  45  was  calculated  f o r each s i t e .  and  s u p p o r t i n g c a l c u l a t i o n s a r e given i n Appendix I ,  R e s u l t s o f a i r photo  Results a r e presented i n Table VI,  guantification  T a b l e I I I summarizes mean o p t i c a l d e n s i t y red:green f i l t e r lichen  r a t i o s o f t h e photo i m a g e s o f h i g h  biomass t r e e s .  a l l trees  optical  taken  density  with high  together  one t r e e  and f o r D o u g l a s - f i r ,  no s i g n i f i c a n t d i f f e r e n c e s .  w e s t e r n hemlock, d i f f e r e n c e s  trees  to trees  reports  Gutermann 1966, G a t e s further  as a p r e d i c t o r  highly  for  cnly  approached  ratio  statistical  lichens  i s consistent  reflectance  patterns of  and o f c o n i f e r f o l i a g e  ( S t e i n e r and  1970).  assess t h e value o f t h e red:green f i l t e r o f l i c h e n abundance I r e g r e s s e d r e d : g r e e n  Results  i n the  of t h e spectrum,  biomass. T h i s  on t h e s p e c t r a l  r a t i o s f o r w e s t e r n hemlock t r e e s  was  In the case of  i n the r e d region  w i t h low l i c h e n  other l i g h t - c o l o u r e d  those t r e e s .  low l i c h e n  w i t h h i g h l i c h e n b i o m a s s had  high r e f l e c t a n c e  published  To  with  (p<0.07). The d i r e c t i o n o f t h e d i f f e r e n c e s  r a t i o suggests that  with  biomass.  comparison o f  i n the red:green f i l t e r  between h i g h and low l i c h e n t r e e s  relatively  lichen  v a l u e s and r e d : g r e e n . f i l t e r r a t i o s f o r t r e e s  biomass r e v e a l e d  compared  had h i g h  l i c h e n biomass with t h o s e f o r t r e e s  significance  and low  S e p a r a t e c o m p a r i s o n s were n o t made f o r  western red cedar because only For  v a l u e s and  ratio filter  cn l i c h e n biomass v a l u e s o f  were d i s a p p o i n t i n g ;  s i g n i f i c a n t ( t = 3.0497;  t  Q  Q  5  although the regression = 2.021), i t a c c o u n t e d  18% o f t h e v a r i a t i o n i n t h e l i c h e n  biomass.  46 TABLE  III.  Optical density values and red:green f i l t e r r a t i o s of trees with high and low l i c h e n biomass (x ± S.D.)  Red Fi 1 t e r  Student s t 1  Green Fi1ter  Student s t 1  Blue Fi1ter  Student's t  Red:green ratio  Student's t  A l l Trees High l i c h e n n = 34 3? = 3917 ± 1268 g  0.529 ± 0.144  Low l i c h e n n = 45 x = 910 ± 726 g  0.537 ± 0.185  0.208  0.976 ±0.290 1  0.563  1.507 ±0.369 1  0.936 + 0.325  0.642  0.559 ± 0.127 1  1.455 ±0.356  1.288  1  1.906  2  0.182  3  0.602± 0.163  Western hemlock High l i c h e n n = 28 x = 3996 ± 1260 g  0.526 ± 0.129  Low l i c h e n n = 15 x = 1084 ± 764 g  0.502± 0.128  1.089 ± 0.252  1.576 ± 0.301  0.470 ± 0.118  High l i c h e n n - 5 x = 3736 ± 1424  0.560 ±0.238  0.954 ±0.512  1.508 ±0.626  0.615 + 0.116  Low l i c h e n n = 10 x = 1379 ± 589.5  0.518 ±0.088  0.999 ± 0.231 0.574  1.527 ±0.310 1.190  2  0.532 ± 0.092 0.498  2  2  Douglas-fir  1  2  3  t  0 5  = 2.000  t  0 5  = 2.021  t  0 5  = 2.160  0.507  0.330  3  0.895 ± 0.192  0.302  3  1.444 ±0.203  3  0.601 ± 0.153  47  Besults individual  of i n t e r p r e t e r s '  trees  compares t h e internal  lichen  are presented  internal  ranking  ranking of the  interpreter's  ratings  ability  biomass w i t h i n  i n T a b l e IV, of  actual  predicted  30  o f t h e 33  correlated {39%)  was  the c o r r e l a t i o n  results  c f the  lichen  his a b i l i t y trees.  to  In a l l  were p o s i t i v e l y 13  cases  s i g n i f i c a n t at the  cases.  The  significant  ecology  :  stronger  The  influence. i n the  individuals  who  to  judge l i c h e n  effect  (photo  T = 218, with  used t o  0.05  evaluate  with photo i n t e r p r e t a t i o n  ability  previous experience  reflected  low  biomass, but i n o n l y  statistically  experience  i n both  statistically 89;  not  predictions  Hann-Whitney t e s t ,  ecology affected  positive  lichen  the  level.  whether p r e v i o u s lichen  but  test  the  o f h i g h and  to i n d i v i d u a l  (91S), l i c h e n  with c a l c u l a t e d  confidence The  cases  on  Spearman's rho v a l u e s with  trees  i s tested,  a s s i g n a b s o l u t e amounts o f l i c h e n but  abundance  biomass v a l u e s . Thus,  to distinguish a plot  of lichen  T  lichen  0  0  was T =  155,  = 83) . Of t h e two  5  ecology  appeared  with  a b u n d a n c e , were  of each f a c t o r  interpretation:  or  T*  were most f a m i l i a r  3 and  t o have  7 - the  with the ecology  =  factors, the  importance of a p r i o r i experience  scores of Interpreters  o o 5  is  two  of the  study  area. Success i n r a t i n g plot  i s of  general  little  level  abundance on correlation  abundance on  p r a c t i c a l value  cf lichen  Spearman's r h o  lichen  test  plots between  on  u n l e s s one  abundance on t h e interpreters'  were n e g a t i v e ; predicted  individual can  rate  p l o t , R e s u l t s of  predictions  t h e r e was  l e v e l of  also  trees  of  a weak  lichen  lichen negative  abundance  and  in a the  48  TABLE IV. Spearman's rho values f o r predictions of l i c h e n abundance on i n d i v i d u a l trees  Interpreter  1  2  Photo I n t e r p r e t a t i o n Experience  L  L  3  L  4  L  5  6  L  L  7  H  8  9  H  H  10  11  H H  Knowledge of l i c h e n Ecology Plot 9  .20  .16  Plot 10  .38  P l o t 11  Spearman's rho  .59  .53  .45  .27  .55  .04  .20 -.07  .62  .44  1  .16  .53  .70  .31  .85  .59  .33  .66  1  = 0.44; n = 15.  Underlined values are s t a t i s t i c a l l y s i g n i f i c a n t . ( p < 1  n = 14; Spearman's rho = 0.46.  0.05)  . l l  .25  .34  .25  -.03  .66  .39  .57  .59  .42  -.30  .71  1  49  actual  level  of l i c h e n  photo i n t e r p r e t e r s a plot  with  between  low l i c h e n  results  c a n compare  with reasonable  distinguish  abundance  {xho = - 0 . 3 5 ) . A p p a r e n t l y ,  l i c h e n abundance  s u c c e s s , but t h e y  areas  on t r e e s  a r e unable  with high l i c h e n  within  to  abundance and  areas  abundance. P o s s i b l e e x p l a n a t i o n s f o r t h e s e  are d i s c u s s e d i n the f o l l o w i n g  section.  some  50  Discussion  E v a l u a t i o n of methods The  methods of determining l i c h e n biomass t h a t were  developed  i n t h i s study r e p r e s e n t a compromise between the need  to q u a n t i f y biomass on a v a r i e t y of s i t e s , and t h e d e s i r e f o r reasonable c o n f i d e n c e i n the numbers obtained. The study was not designed  i n such a way t h a t i t i s p o s s i b l e to put c o n f i d e n c e  l i m i t s on the f i n a l biomass t o t a l s , but the methods which produced The  the biomass t o t a l s can be assessed. use of f e l l e d t r e e s f o r epiphyte sampling  criticized  has been  by workers c a r r y i n g out d e t a i l e d r e s e a r c h on e p i p h y t e  d i s t r i b u t i o n w i t h i n a s i n g l e tree not b e l i e v e i t caused  (Pike e t a l . 1972), but I do  any s e r i o u s problems i n t h e present  study.  Despite some breakage o f tranches and s c a t t e r i n g of e p i p h y t e s , little difficulty  was experienced i n determining  which of the  f a l l e n l i c h e n had come from t h e sample t r e e . Some e r r o r was caused  probably  by twigs knocked o f f adjacent t r e e s o f t h e same  s p e c i e s by the f a l l i n g  sample t r e e . T h i s p o t e n t i a l source of  e r r o r i s a f a c t o r t h a t should be taken i n t o c o n s i d e r a t i o n when d i r e c t i o n of f e l l i n g i s decided. Seme of the r e g r e s s i o n s r e l a t i n g L i c h e n Estimates t o l i c h e n biomass on branches  d i d not have as high r  2  v a l u e s as expected;  those of the f i r s t two t r e e s sampled were o n l y 0.34 and 0.30. F o r t u n a t e l y , the r a t i o of sampled branches  to unsampled  was high  sampled tended  (1:2 c r b e t t e r ) , and t h e branches  branches to be  51  those  with  high  lichen  abundance. T h e r e f o r e ,  c a l c u l a t i o n s f o r t r e e s were b a s e d a c t u a l samples.  Despite  better to extrapolate unsampled have  be  improved  (probability  the  Cochran  cases,  on  i t seems  t h a n t o assume t h a t  sampled  ones,  as some w o r k e r s  g u a n t i f i c a t i o n o f l i c h e n b i o m a s s on  by  using  a method a n a l a g o u s t o  proportionate  1963:251). T h i s  necessarily  accurate)  population,  and  to prediction)  size  how  with  weighed, and predicted  clumps o f  each branch.  a probability  the  results  used t o  biomass e s t i m a t e s ,  confidence In the  limits  present  v i s u a l estimates  proportionate to t h e i r  study, of  sampling  that  and  the  the  (but  sample  t o the  from  would consistent) be  p r e d i c t i o n of  s a m p l e b r a n c h e s would  be  determine a c o r r e c t i o n r a t i o An  advantage to  confidence  i f the  limits  p r e d i c t i o n s are  each p l o t  this  around  the  consistent,  were s e l e c t e d  abundance a c c o r d i n g  scheme. T r e e s  of  narrow.  15 t r e e s on  lichen  not  Sample b r a n c h e s would  are r e l a t i v e l y  representation  scheme e n s u r e d  to put  1967;  the  researcher  arbitrary  b i o m a s s t o measured b i o m a s s .  method i s t h a t i t i s e a s y final  an  proportionate  b i o m a s s on  scheme u s e d  (but  unit i n  branches  3P  error of a small  t h i s approach, the  l i c h e n biomass. L i c h e n  sampling  method r e q u i r e s a p r e c i s e  many l i c h e n on  the  (Grosenbaugh  p r e d i c t i o n f o r each  Using  were p r e s e n t  selected  volume  measurement w i t h o u t  population.  predict  for  same as  f o r e s t e r s to determine timber  the  i n a few  extent  done.  could  the  values  2  biomass  to a considerable  from r e g r e s s i o n s  branches are  I believe that  by  poor r  the  to a  were s e l e c t e d i n  random  proportion  i n each species-dominance c l a s s .  The  species-dominance c l a s s e s having  the  52  largest  numbers were most h e a v i l y s a m p l e d .  9 species-dcminance class  was  sometimes r e p r e s e n t e d  Variability great.  relatively  i n forests  biomass i n a s t a n d  on  as t h e  scales  strata  The be  system  w e i g h t s on classes. trees was  visual  to t e s t  and  more than  the  present  range, although one  researcher  system  p o s i t i o n s from  the  most o f i t , c a n this  mainly  on  bottcm s i t e s assess.  the  may  not  sidehill  by  abundance,  of this of  within  monitored  which s h o u l d  be  i f  noted.  of the researcher to f i n d entire  an  estimates.  crown o f t h e  viewed. In v e r y  be  p o s s i b l e . The  w i t h i n the study  set  seems t o be  be c a r e f u l l y  be  sites;  a second  because of l i m i t a t i o n s  makes v i s u a l  which t h e  lichen  f o r a l l species-dominance  i t should  ability  of  o f t h e r e g r e s s i o n , but  study  used  l i c h e n a b u n d a n c e seems t o  regression equation  value  be  stand.  of  0.75  of  as the b a s i s f o r  has s e v e r a l l i m i t a t i o n s  or two  out  was  predictive  i t d e p e n d s on  forests,  and  estimates  estimates  i n the  a  sometimes  abundance c l a s s e s would t h e n  value of the  First,  least  to l i c h e n  money. I n t e r - o b s e r v e r r e l i a b i l i t y  acceptable  The  2  as  trees.  improved  I t would have been d e s i r a b l e t o f e l l  impossible  time  r  c o u l d be  biomass t o  of v i s u a l  w o r k a b l e . The  o r two  plot,  l i c h e n abundance, e s t i m a t e s  according  Lichen  lichen  one  within a  w i t h i n a c l a s s was  for tree s e l e c t i o n ,  e x t r a p o l a t i o n of  only  probably  trees directly  a 1-5  present  s i n c e a s many  where s p e c i e s - d o m i n a n c e c l a s s e s a r e  homogenous i n t h e i r  stratifying e.g.  by  of l i c h e n abundance  Except  lichen  c l a s s e s might be  But  tall  or  tree,  very  present study  one or  at  dense was  I b e l i e v e t h a t some o f t h e  carried valley  a r e a w o u l d have b e e n d i f f i c u l t  to  53  Second, the system  was designed to d i s c r i m i n a t e among t r e e s  which vary g r e a t l y i n t h e i r l i c h e n l o a d s . Estimates of a l e c t o r i a ( s . l . ) cover f o r l a y e r s of t r e e s encountered i n the study area ranged from 0 t o 80%. In i t s present form, the system  would not  be a p p r o p r i a t e f o r making f i n e r d i s t i n c t i o n s as, f o r example, d i s c r i m i n a t i n g among t r e e s i n young stands i n which A l e c t o r i a cover never exceeded  5%.  T h i r d , i t must not be assumed that the r e g r e s s i o n s used i n the present study t o e x t r a p o l a t e from sampled t r e e s t o unsampled t r e e s a r e general i n t h e i r a p p l i c a b i l i t y . I f l i c h e n biomass i s to be assessed i n other areas, f u r t h e r biomass sampling should be undertaken i n those f o r e s t types t o determine between v i s u a l estimates and l i c h e n  relationships  biomass.  Results of the densitometry i n d i c a t e some p o t e n t i a l f o r d e t e c t i o n of l i c h e n abundance using red:green f i l t e r r a t i o s . Use of r a t i o s r a t h e r than a b s o l u t e d e n s i t y values e l i m i n a t e s much o f the v a r i a t i o n i n o p t i c a l d e n s i t y readings caused by d i f f e r e n t i a l lighting  c f t r e e s . I t i s l i k e l y t h a t a l a r g e r sample s i z e would  show that red:green r a t i o s o f t r e e s with high l i c h e n biomass d i f f e r s i g n i f i c a n t l y from those of t r e e s with low l i c h e n biomass,  at l e a s t f o r some s p e c i e s . However, the present r  value of the r e g r e s s i o n  2  ( r = 0.18) suggests that the p r e d i c t i v e 2  value of the r a t i o i s low. Photo i n t e r p r e t a t i o n r e s u l t s inability  were d i s a p p o i n t i n g . The  of i n t e r p r e t e r s to d i s t i n g u i s h areas o f high l i c h e n  abundance from areas o f low l i c h e n abundance suggests that the p o t e n t i a l u s e f u l n e s s of a i r photography  as a l i c h e n i n v e n t o r y  t o o l i s low. S e v e r a l f a c t o r s may e x p l a i n these poor  results.  54  First,  the  presence of  crowns o f  trees  to d e t e c t  abundance o f  spp.  are  lichen two  factors  differences  flying  from an  and  may  occurred.  were more  I had  1:2000, 1:4000, and  lower f l y i n g  height  reason, scale. large this  the  scale  among  account  The results and  be  i s the  growth  habit  species  tend t o  form  may  that  by  three  over  photographs taken at in scale.,For  actual  to  the  this  also  varied in  in  the  abundance,  have c o n t r i b u t e d  of  of  the  lichens.  Bryoria  and  distinguish  be  degree t o which they  are  visible  good r e p r e s e n t a t i o n remarked t h a t  visible  the  the  photograph  study  area,  l i c h e n abundance can  sarmentosa  growth  directly  f r o m above  to  may  above,  not  be  but  a  interpreter  a s s e s s on  trees  b e c a u s e t h e y were s e e n  I have observed be  poor  which hang f r o m  from  easier  the  Alectpria  t h e i r a c t u a l b i o m a s s . One  l i c h e n abundance was  edge o f In  of  to  have a p e n d e n t  e l o n g a t e d clumps may  low  of  difficulties  level  for interpreters' i n a b i l i t y  that  obliguely.  the  more n o t i c e a b l e of  t w i g s . They  near t h e  believe  planned f o r  interpreters  regardless  factor  b r a n c h e s and  and  the  by  tended t o  third  they  Bryoria  ;  most a r b o r e a l  habit;  the  lichens  anility  t h e y were c o n f u s e d  1:8000. But  were v a r i a b l e  photographs,  sites.  However, I  above g r o u n d  photographs assessed  Arboreal  may  heights  in general.  originally  m o u n t a i n o u s t e r r a i n were s u c h t h a t two  interpreter's  the  important.,  i n photo s c a l e .  at a constant  high i n  have c a u s e d u n d e r e s t i m a t e s  i n t e r p r e t e r s remarked t h a t  photo s c a l e s :  Alectoria  Alectorioid lichens  abundance where t h e y  other  mixed w i t h  have d e t r a c t e d  dark i n c o l o u r  Several  of  may  Bryoria  distinguished  that with  areas of  high  binoculars  55  from  a viewpoint  that  oblique  across a valley.  aerial  p h o t o g r a p h y m i g h t be  l i c h e n abundance, a t l e a s t  Biomass r e s u l t s The compared  lichen  compared  at the  with  in this  with r e s u l t s o b t a i n e d  i n other  north  Alectorioid  biomass t o t a l  in this  totals reported,  but  most o t h e r r e p o r t e d the  ttaximum  reported  but  study the  f i g u r e s are  provide  an  study  are  temperate not  a l l l i c h e n s and  they  production  others include  i d e a of the  t h a t o c c u r s . The  (1528  values. I t i s only  and  strictly  any  kg/ha) i s g r e a t e r about h a l f as  v a l u e - 3289 k g / h a , r e p o r t e d f o r f i r f o r e s t i n the  range  lowest  (21 kg/ha) i s s m a l l e r t h a n  highest  Englemann s p r u c e - s u b a l p i n e Columbia,  The  b a s e d on  lichens,  magnitude o f l i c h e n  V.  stage.  reports  obtained  c o m p a r a b l e , as some a r e  in  reconnaissance  literature  suggest  useful i n inventorying  biomass t o t a l s  b o r e a l ecosystems i n Table  only  These o b s e r v a t i o n s  interior  other  than  great  as  an of  British  TABLE V. Comparison of reported biomass t o t a l s of epiphytic lichens  Source  Area  Edwards et a l . 1960  Wells Gray Park, B.C.  Scotter 1962  Black Lake, northern Saskatchewan  Forest Type  Mi xed conifers  282  Picea Abies  engelmanniilasiocarpa  754  Picea Abies  engelmanniilasiocarpa  3289  Picea  mariana  Pinus  banksiana  Pinus  excelsa  B.iazrov 1969  U.S.S.R.  Picea  W.D. Boehm 1972 unpublished data  Ross Lake, Washington  Pinus  Tsuga  All  lichens  Bryopogon  abies-mixed  deciduous  contorta  Pseudotsuga  Oregon  837  sarmentosa spp.  contorta  Archangel U.S.S.R.  Pike et a l . 1972  Alectoria and Bryoria  Pinus  Rudnova, Tonkonogov, & Dorokhova 1964 ( c i t e d i n Rodin & B a z i l e v i c h 1967)  Province,  Oven-dried weight (kg/ha)  Epiphytes  menziesii  All  rl  implexus  menziesii  3  sarmentosa spp.  2  2  700  600 282 47-233 527  heterophylla  Pseudotsuga  2051  lichens  Alectoria Bryoria  198  1  All  lichens * 1  564  Ln ON  TABLE V. (Continued)  Source  Area  Schroeder 1974  Forest Type  S e l k i r k Mountains, B.C. & Washington  Picea Abies  engelmanniilasiocarpa  Picea Abies  engelmanniilasiocarpa  Larix  occidentalis  Andre et a l . 1975  France  Abies  Wein & Speer 1975  Cape Breton Island, Nova Scotia  Turner & Singer 1976 This study  Oven-dried weiqht (kg/ha)  Epiphytes  430 A l l macrolichens  5  103 429  All  lichens  1040  Abies balsamea-Picea mariana  All  lichens  47-280  Western Washington  Abies amabilis-Tsuga mertensiana  All  lichens  1900  Northern Vancouver Island  Tsuga heterophyllaPseudotsuga menziesiiThuja plicata  1  P r i m a r i l y Bryoria  2  A i r - d r i e d weight.  3  Probably Bryoria  4  Only epiphytes on overstory D o u g l a s - f i r included.  5  P r i m a r i l y Alectoria  alba  spp ., Evernia mesomorpha. Usnea hirta.  implexa.  sarmentosa  and Bryoria  spp.  Alectoria sarmentosa and Bryoria spp.  21-1528  58  IV.  RELATIONSHIPS  BETHEEN LICHEN BIOMASS AND  SITE  CHARACTERISTICS  The  second  o b j e c t i v e of the study  was  to r e l a t e  of f c r a g e l i c h e n s t o p h y s i c a l  and  sites.  p o r t i o n of the study  The  rationale  information  gathered  of A l e c t o r i o i d areas  with  abundant  and  would a s s i s t  s h o u l d be  that  r e a c h i n g the  should  lichen  a s p e c t s and  n e g a t i v e l y w i t h crown c l o s u r e  o f crown c l o s u r e ,  positively  with the r a t i o  a measure which has  crown c o m p e t i t i o n . The and  inadequate  low  and  I a l s o expected  die that  elevation.  study  ( B o c h e l l e 1978)  1975), and  lichen  of l i v e  proposed  t e n d s t o be  (Hard  1964,  lichen  and  an  limiting),  penetrating the  crown  a s an  to  Chiam  tree  indicator  branches  of  adjacent  receive  1967). would  i n c r e a s e with  a l r e a d y been n o t e d  i s probably a s s o c i a t e d with  be  h i g h i n open-grown  abundance  T h i s t r e n d had  at  abundance would  s t a n d s , where t h e l o w e r  increasing area  been  ratio  i n dense  light  high  radiation  (except  where s u b s t r a t e becomes  that  canopy. I a l s o expected  levels  L i c h e n abundance  tree  stands  forest  abundance  p o t e n t i a l amount o f s o l a r  t h e amount o f l i g h t  height,  ecology  s t e e p s l o p e s because  b e c a u s e crown c l o s u r e l i m i t s  correlated  the  correlated  a s s o c i a t e d with high  a surface at a given l a t i t u d e .  be c o r r e l a t e d  levels  in identifying  t r e e canopy. High  a s s o c i a t e d with south  on  that  of the  h i g h l i c h e n abundance would be  these f a c t o r s i n c r e a s e the incident  was  of  lichen.  measures which a r e t h e m s e l v e s  of s u n l i g h t  low  vegetative characteristics  would i m p r o v e u n d e r s t a n d i n g  lichens,  I expected with  for this  abundance  watershed  increasing  i n the  (Barichello  humidity.  59  although crown closure also tends to be more favourable f o r i  lichen growth at higher elevations (Jones 1975, Barichello 1975) . Data collected by Bochelle (197.8) suggested that high lichen abundance would be negatively correlated with basal area and with stand height.  60  Methods  Data were c o l l e c t e d abundance was  at the f o u r t e e n p l o t s on which l i c h e n  estimated. A complete  floristic list  of v a s c u l a r  p l a n t s and major bryophyte s p e c i e s of the f o r e s t f l o o r was made at each p l o t . A percent cover value was estimated f o r each s p e c i e s , and each s p e c i e s except those i n the t r e e l a y e r assiened a d i s t r i b u t i o n c l a s s . D i s t r i b u t i o n c l a s s e s were: 1.  rare i n d i v i d u a l  2.  few s c a t t e r e d  3.  s i n g l e patch  4.  several scattered  5.  a few s m a l l patches  6.  s e v e r a l well-spaced  7.  continuous cover of well-spaced  8.  continuous dense cover with a few  9.  continuous dense cover u n i n t e r r u p t e d  individuals  patches  Topographic  individuals openings  using the method of t a b u l a r comparison  by the Braun-Blanquet  (Mueller-Dcmbois  1  individuals  Vegetation was c l a s s i f i e d developed  was  School of P h y t o s c c i o l o q y  and E l l e n b e r g 1974: 177-193);. measurements taken were e l e v a t i o n , measured  with an a l t i m e t e r ; s l o p e , measured with a c l i n o m e t e r ; and aspect, measured with a compass. For computer a n a l y s i s , aspect  Developed by the Vegetation F u n c t i o n a l Subcommittee of the B r i t i s h Columbia Land Resources Committee, Environment and Land Use' Committee, M i n i s t r y of Environment, B r i t i s h Columbia. P r e s e n t l y the Resource A n a l y s i s Branch, M i n i s t r y of Environment, B r i t i s h Columbia. 1  61  readings  were c o n v e r t e d  Measurements o f s l o p e tables  (Buffo  potential The  et  i n a narrrow topography  (Garrison  not  1949)  canopy was  was  and  except it is  Plot  12.  located by  the  "mocsehorn"  data  (see C h a p t e r 3)  trees  and  met  16  (Lemmon  over  3 m or  the  technigue  1956).Mean  readings of  located  the  main  tree  visually.  18 cm  were a l l dead p o t e n t i a l  i f they  were u s e d  to c a l c u l a t e  i n diameter  were  (merchantable) t r e e s  more i n h e i g h t .  above c r i t e r i a  18  Down t r e e s  (British  cm  were  Columbia  Forest  1973). height the  of the  measured  codcminant heights  o f crown l e n g t h  calculated,  b a s e d on  t r e e l a y e r was  of the  to t r e e  estimated,  v i s u a l estimate trees.  height  measurements o f  (CL/TH)  The  was  v i s u a l estimate trees  (see  3). variables described  above were r e l a t e d t o a  representing  l i c h e n abundance:  per  Tsuga s p e c i e s .  1  annual  i s restricted  e a c h p l o t . Crown c l o s u r e  A l l living  as  mean r a t i o  The  site  because  were c a l c u l a t e d from  within  o r more i n d i a m e t e r  Chapter  12  length  measured u s i n g  also estimated  area.  b a s e d on  Plot  each  a s p h e r i c a l densiometer  values  Tree inventory  included,  with s o l a r r a d i a t i o n  were u s e d t o d e t e r m i n e  applicable to  south.  ( B u f f o e t a l . 1972).  systematically  The  i n d e g r e e s f r o m due  together  v a l l e y i n w h i c h day  crown c l o s u r e  Service  aspect  a l . 1972)  Crown c l o s u r e  included  and  departure  s o l a r r a d i a t i o n i n c i d e n t on  t a b l e s are  basal  to  l a y e r on  Hereafter  r e f e r r e d t o as  1  mean p e r c e n t  This  alectoria  v a r i a b l e was  "percent  variable  Alectoria"  c h o s e n as  (s.l.) a  62  measure o f t h e  q u a l i t y of  of the  s i z e or  number o f  rather  than a l l tree  species  was  suggesting than  site  tree  t r e e s on  species  f o r l i c h e n growth, the  site.  that as  on  other  a limiting of  the  Alectoria l a y e r was  species  Hemlocks were used  t h a n on  substrate  may  other be  Tsuga  species,  more  important  factor.  vegetation  (s.l.)  regardless  b e c a u s e l i c h e n b i o m a s s on  almost i n v a r i a b l y greater  Besults total  a site  a n a l y s i s were a l s o r e l a t e d  biomass.  related to  1  The  height  e l e v a t i o n as  of  the  w e l l as t o  to  codcminant percent  Alectoria,. Relationships reqression Statistical  1  Hereafter  model o f  amonq v a r i a b l e s were t e s t e d u s i n g the  computer  proqram MIDAS o f  Research Laboratory, U n i v e r s i t y of  r e f e r r e d to  as  " A l e c t o r i a biomass"  the  the  Michigan.  linear  63  Besults  Table VI summarizes the data used i n the a n a l y s i s of l i c h e n abundance i n r e l a t i o n t o s i t e  characteristics.  V e g e t a t i o n communities The v e g e t a t i o n c l a s s i f i c a t i o n r e s u l t e d i n the d e l i n e a t i o n of three communities  (Appendix  I I ) . Because my data base  was  i n s u f f i c i e n t f o r a comprehensive c l a s s i f i c a t i o n of the v e g e t a t i o n i n the study area, the f o l l o w i n g comments are on f i e l d  impressions and  based  p u b l i s h e d c l a s s i f i c a t i o n s of s i m i l a r  areas, as w e l l as on data c o l l e c t e d at p l o t s . P l o t s 1, 3, and  10 formed a group which was  D o u g l a s - f i r / S a l a l Community combination  (DF/S). The  labelled  the  characteristic  of s p e c i e s f o r t h i s community i s Pseudotsuqa  menzijasii^ G a u l t h e r i a s h a l l o n , Hemitomes congestum, Stokesie11a oregana  A  V i o l ^ sempervirens„ and Boschniakia h o o k e r i . The  community develops on x e r i c s i t e s , most f r e q u e n t l y on  south-  f a c i n g s l o p e s , r i d g e s , and k n o l l s . In the study area, i t may occur from  the v a l l e y bottom to about 700  elevations i t i s limited The  to very x e r i c  m, although a t high  sites.  D o u g l a s - f i r / S a l a l Community i s s i m i l a r to the  (splendentis) - Eurhynchio  Hyloccmio  (oregani) - G a u l t h e r i o ( s h a l l o n i s ) -  Pseudotsugetum m e n z i e s i i (the G a u l t h e r i a shalIon a s s o c i a t i o n ) of Kojima and  K r a j i n a (1975) and t o the cjrthic G a u l t h e r i a f o r e s t  type of O r l o c i  (1961). Kojima and  K r a j i n a (1975) d e s c r i b e d t h e i r  Gaulthjgria s h a l l on a s s o c i a t i o n as o c c u r r i n g on x e r i c s i t e s with extremely  w e l l - d r a i n e d and shallow s o i l s , and a r e l a t i v e l y  high  64 TABLE V I .  . Plot ^ectorza Number 7, ?u , < 9/ ) 7  A  0 n  S S  k  P  e  r  c  e  n  t  Alectoria  Elevation (m)  . . aspect  (  Slope D e g r e e s  ha  )  Site c h a r a c t e r i s t i c s of plots  Potential Annual Radiation (cal/cm /year)  Densiometer  Moosehorn  2  Crown Closure {%)  Crown ; Height , Length/ of Co- j ? " * ... , , Tree Dominant , , Height Tree's (m) R  1  L  V  i  s  u  a  l  2  ( m  /  /  h  a  Vegetation Community  )  1  1528  35.3  686  S15E  32  222029  79  43  50  79.0  25  84.4  DF/S  2  175  01.0  442  S40E  15  193182  88  68  60  63.0  40  109:3  MOSS  3  665  26.7  543  S30E  54  215244  77  74  60  72.9  34  87.1  DF/S  4  942  18.1  709  S60E  22  188467  87  90  75  61.4  32  112.5  5  303  11.5  573  N75E  23  147791  86  64  70  73.5  44  89.7  AF/0H  6  975  23.3  669  S60E  30  192113  79  61  65  68.9  22  79.0  TRANS  7  140  04.5  282  N80W  20  154933  88  87  80  72.4 '  47  84.2  MOSS  8  289  16.0  707  S05W  5  181201  79  67  50  70.4  37  75.3  AF/0H  9  800  18.8  646  N82W  32  149169  86  79  70  66.1  40  105.6  TRANS  10  521  15.4  469  S10W  20  209424  87  78  75  65.3  36  93.8  DF/S  11  1516  25.9  792  S60E  14  184887  90  89  80  71.7  32  100.7  TRANS  12  21  00.1  487  S22W  32  -  94  83  60  71 .0  40  90.7  UNCLASSIFIED  13  107  04.0  426  N87W  25  161065  90  83  75  63.3  46  120.3  TRANS  14  176  06.0  603  N35W  24  108470  87  83  60  71.1  47  89.4  AF/OH  UNCLASSIFIED  65  slope g r a d i e n t . Crown c l o s u r e was  r e l a t i v e l y low  (x = 65%)  l i g h t i n t e n s i t y under the t r e e canopy the highest reported  reported years  (x = 3239 lux)  i n t h e i r study. F o r e s t p r o d u c t i v i t y of the  s h a l l c n a s s o c i a t i o n i s low; as 33 m/100  (Kojima and P l o t s 5, 8,  the Amabilis  and  Gaultheria  the s i t e index f o r D o u g l a s - f i r  years and  was  f o r western hemlock, 26 m/1-,00  K r a j i n a 1975). 14,  and  15  1  formed a group which was  F i r / O v a l - l e a v e d Huckleberry  Community  labelled  (AF/OH).The  c h a r a c t e r i s t i c combination of s p e c i e s f o r t h i s community i n c l u d e s Abies a m a b i l i s , Ccrnus c a n a d e n s i s R u b u - s pedatus^ vaccinium  o v a l i f o l i u m , Menziesia  ferruqjnea, Listera  C l i n t e n i a u n i f l o r a , Streptopus roseus, and  Tsucja mertensiana. The  Plagiothecium  community develops on  caurina, undulatorn,  mesic  sites,  most f r e q u e n t l y on g e n t l e or moderate n o r t h - , e a s t - , or westf a c i n g s l o p e s above 500 The  Amabilis  m in elevation.  F i r / O v a l - l e a v e d Huckleberry  comparable to the R h y t i d i a d e l p h o  (lorei) - Plagiothecio  (undulati) - Rubo (pedati) - V a c c i p i o (amabilis) - Tsuqetum h e t e r o p h y l l a e a s s o c i a t i o n ) of Kojima and  Community i s  (alaskaensis) -  Abieto  (the Vaccinium alaskaense  K r a j i n a (1975), and the  clintonicsum  v a r i a n t of the Abieto - Tsuqetum h e t e r o p h y l l a e o f O r l o c i (1961). The  Vaccinium alaskaense a s s o c i a t i o n (Kojima and  Krajina  occurs on mesic-subhygric s i t e s i n the wetter subzone of C o a s t a l Western Hemlock Zone c h a r a c t e r i s t i c of gentle of v a l l e y s , and  1  the  ( K r a j i n a 1965). I t i s  slopes and  of t e r r a c e s near the bottom  i t s d i s t r i b u t i o n appears to be r e l a t e d to  No l i c h e n estimates  1975)  were performed on P l o t  15.  the  66  distribution  of snow-pack. Crown c l o s u r e was  the  Vaccinium  the  canopy  was  intermediate; years  and  alaskaense rather the  a s s o c i a t i o n , and  low  site  (x =  index  1943  high  light  80%)  intensity  lux). Productivity  f o r w e s t e r n hemlock  f o r a m a b i l i s f i r , 37  (x =  m/100  years  was  in under  was  40  m/100  (Kojima and  Krajina  group  included  1975). Plots  6,  9,  11,  and  13  species c h a r a c t e r i s t i c of and  the A m a b i l i s  and  Amabilis  between t y p i c a l  on  i s too  very  but  high  xeric  a t an  f o r the  sites  (792  Plot  that are  steep  and  Four p l o t s c o u l d not approach, these  probably  f o u r , two  floristically Hyocomio  sites.  (669  1 ) . P l o t 11 outside  i s on  the range of 13 a r e  on  by  K o j i m a and  occurring  be  classified  using  sample s i z e are  the  was  too  t o the  hyloccmiosum s p l e n d e n t i s  Eurhynchio  structurally  Zone  {Krajina  small.  similar  v a r i a n t of the  (oregani)  -  1965),  on  subzone of the C o a s t a l  Of  and of  the  Mahonio (the  moss a s s o c i a t i o n )  ( 1 9 7 5 ) . T h i s v a r i a n t was  drier  the  Braun-Blanguet  7)  i n the  site,  well-drained.  (2 and  Krajina  except  west-facing  plots  splendens  a  m a.s.l.)  a xeric  (nervosae) - Pseudotsugo - Tsugetum h e t e r o p h y l l a e Hyloconiium  sites  P l o t 6 i s on  elevation  . P l o t s 9 and  because the  (splendentis) -  on  Douglas-fir/Salal sites  a t an  m a.s.l.)  D o u g l a s - f i r / S a l a l Community slopes  This  pure D o u g l a s - f i r / S a l a l Community  (e.g.  elevation  slope  Community.  (THANS), o c c u r s  Fir/Oval-leaved Huckleberry  subxeric, southeast-facing that  Huckleberry  T r a n s i t i o n Community  are intermediate  which  b o t h t h e p o u g l a s - f i r / S a l a l Community  Fir/O val-leaved  group, termed the that  formed a t h i r d  described  described Western  s e l l - d r a i n e d mesic s i t e s  where  as  Hemlock slopes  67  are moderate and s o i l s r e l a t i v e l y (x = 85%) and l i g h t i n t e n s i t y  deep. Crown c l o s u r e was high  under the f o r e s t canopy low (x =  1750 l u x ) . P r o d u c t i v i t y was i n t e r m e d i a t e ; the s i t e index was 40 m/100 years f o r D o u g l a s - f i r and 37 m/100 hemlock. The Hyloccmium splendens a s s o c i a t i o n has Pseudotsuga  years f o r western  v a r i a n t of the moss  menziesii  f  Tsuga h e t e r o p h y l l a , and  Thuja p l i c a t a i n the t r e e canopy, low cover i n t h e shrub and herb l a y e r s , and a well-developed mess l a y e r i n c l u d i n g HyjLocomium splendens, R h y t i d i a d e l p h u s l o r e u s  A  and S t o k e s i e l l a  2Iia2Bli Because P l o t s 2 and 7 occurred on mesic,  moderately  s l o p e d , low e l e v a t i o n s i t e s and were s t r u c t u r a l l y and floristically  s i m i l a r t o the Hyloccmium s p l e n d e n s s i t e s  d e s c r i b e d by Kojima  and K r a j i n a  (1S75), they were given the  t e n t a t i v e l a b e l of "Moss Community" {BOSS) . The two remaining  p l o t s were not c l a s s i f i e d . I had expected  that P l o t 4 would belong with t h e T r a n s i t i o n Community, but the r e s u l t s of the a n a l y s i s dc not support c l a s s i f y i n g i t with that group. The p l o t has few s p e c i e s c h a r a c t e r i s t i c o f the Douglasf i r / S a l a l , A o a b i l i s F i r / o v a l - l e a v e d Huckleberry, or T r a n s i t i o n Communities, but d i f f e r s of  from Moss s i t e s i n i t s high cover (40%)  Vaccinium alaskaense . P l o t  12 d i f f e r s from a l l other p l o t s  i n i t s v e g e t a t i o n and i t s t o p o g r a p h i c p o s i t i o n . I t has very low cover and s p e c i e s d i v e r s i t y i n a l l l a y e r s beneath  the t r e e  canopy. The p l o t i s l o c a t e d on a steep s o u t h - f a c i n g s l o p e near the bottom of a narrow east-west r e s t r i c t e d by topography. P l o t s 4 and 12.  v a l l e y , where i n s o l a t i o n i s  More data would be needed t o c l a s s i f y  68  The  relationship  between v e g e t a t i o n community and  abundance i s shown i n F i g u r e 9. The  v e g e t a t i o n communities which  appear to have the g r e a t e s t p o t e n t i a l the D o u g l a s - f i r / S a l a l  for lichen  ( F i g . 9E)  production  Community and the T r a n s i t i o n  r e g a r d l e s s of whether A l e c t o r i a biomass Alectoria  lichen  are  Community,  ( F i g , 9A)  or  i s measured, w i t h i n these two  percent  communities  there i s a s t r o n g tendency f o r l i c h e n abundance to i n c r e a s e with elevation.  P l o t U, the u n c l a s s i f i e d  group with the T r a n s i t i o n l i c h e n abundance by both The  p l o t which I expected  Community, i s a l s o r e l a t i v e l y  high i n  measures.  Amabilis F i r / O v a l - l e a v e d Community i s low in- A l e c t o r i a  biomass, but i n two  of the t h r e e p l o t s  moderate r a t h e r than  percent A l e c t o r i a i s  low, These r e s u l t s suggest  t h a t some s i t e s  of t h i s type are moderate i n s u i t a b i l i t y f o r l i c h e n but that amount of a v a i l a b l e Community i s low only two  to  in lichen  s u b s t r a t e i s lew. The  production, Moss  production by both measures.  s i t e s were sampled, f i e l d impressions support  Although the  c o n c l u s i o n that Moss s i t e s are c h a r a c t e r i s t i c a l l y low i n production of forage  lichens.  Measures of p h y s i c a l  environment  Scattergrams potential  of percent  annual r a d i a t i o n ,  are given i n F i g u r e s 10 to Variation  i n slope  and  on s l o p e ,  and e l e v a t i o n  aspect,  on percent  Alectoria  13.  ( F i g . 10}  the v a r i a t i o n i n percent value c f only 0.18  Alectoria  accounts  f o r very l i t t l e  A l e c t o r i a ; • the r e g r e s s i o n has an  i s 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 .  of r  2  1500H  ® cn  CO <  1000"  50(H  n PLOT 10 3 1 DF/S  13 9  6 11  5  TRANS  14 8  7  AF/OH  2  MOSS  12 4 U  30H  20H i I  10H  I  PLOT 10 3 1 DF/S  13 9  6 11  TRANS  Figure 9. Vegetation community and Aleetoria  5  14 8  AF/OH  7  2  MOSS  biomass (kg/ha) ( A ) ;  vegetation community and percent Alectoria  ( B ) . Within  communities, p l o t s are arranged i n order o f increasing elevation.  U = unclassified.  12 4 U  40-  <  30H  rr  O — i o UJ <  20-  O  F-  z  LU O  v  DC LU  0- 10-  _ _  I 50  T  30  20  10  40  SLOPE degrees I  DF/S community | A  TRANS community  V  AF/OH community | MOSS community unclassified  Figure 10.  I  Relationship between slope and percent (y = 7.6012 + 0.33442; n = 14; S . V  Y  Alectoria  = 9.75; r  2  = 0.18).  71  40-1  30H o o LU  20H LU  V  O  CC LU  10-  _Q_  "3o~  180  30 60 90 120 150 ASPECT (degrees from s o u t h )  • DF/S community A TRANS community v AF/OH community B MOSS community o unclassified Figure 11.  Relationship between aspect and percent (y = 23.376 - 0.120 x; n = 14; S . y  x  Alectoria  = 9.31; r  2  = 0.25)  72 (  40  -I  16 P O T E N T I A L  A N N U A L  10,000 x  18  r 20  22  R A D I A T I O N  cal/cm /yr 2  © DF/S community A  TRANS community  v AF/OH community a MOSS community o  Figure 12.  unclassified  Linear regression of percent Alectoria  on potential annual  radiation (y = -19.081 + 0.00197 x; n. = 13; S . y  r  2  = 0.37).  x  = 8.56;  40-i  < 30H  200  400  600  800  ELEVATION meters © DF/S communi ty A TRANS communi ty v AF/OH community a MOSS communi ty o unclassified  Figure 13.  Linear regression of percent Alectoria (y = -15.465 + 0.0527 x; n = 14; S . y  x  on elevation = 8.14; r  2  = 0.47).  74  The degrees  r  v a l u e of  2  frcm  due  percent  south,  Alectoria  i s 0.25,  slightly  s l o p e , but  a l s o i s not s t a t i s t i c a l l y  triangular  shape c f the s c a t t e r g r a m  i s a necessary  lichen  abundance. P o t e n t i a l f o r h i g h  not always r e a l i z e d . appear t o e x p l a i n the p l o t s , although than  expected  illustrates  the  low  levels  south,  of l i c h e n cn  both  b a s i s o f aspect.. The  but  and  Transition  sites  Potential the  variation  statistically  annual  scattergram  a triangular  higher than on  The  2. 522,  t  upper l i m i t  0  O  5  The greatest  on  Transition  lichen sites  which has  v a l u e s , whereas t h a t cn environmental  variation an r  2  i n percent  value  sites  sites  cn  on  south  north —  of 0.47.  e x p l a i n s 37%  percent  Alectoria  that  have amounts o f  sites  11,  this  solar  Moss s i t e s  Alectoria Alectcria  i s c l o s e to is  lower.  f a c t o r which accounts  The  on  p r o d u c t i o n . Most  regression. Percent  Alectoria  of  regression indicates a  shape which s u g g e s t s  t h a t p r e d i c t e d by t h e  single  lower  scattergram  =2.201). L i k e F i g u r e  Amabilis F i r / O v a l - l e a v e d Huckleberry  predicted  not  some  Moss s i t e s a r e  ( F i g . 12)  dependence o f  (t =  D o u g l a s - F i r / S a l a l and  do  a s l o p e o f only 5 degrees  Alectoria.  significant  s e t s an  has  solar radiation  solar radiation  radiation  potential i s  on i n t e r m e d i a t e a s p e c t s .  i n percent  has  the  appears  abundance on  of Douglas-fir/Salal  aberrant s i t e  The  c o n d i t i o n f o r high  D i f f e r e n c e s i n v e g e t a t i o n community  the occurrence  the  { F i g . 11).  that southerly  aspects, Amabilis F i r / O v a l - l e a v e d Huckleberry aspects —  that, c f  percent alectoria  due  lichen guantities  on  than  significant  but not s u f f i c i e n t  approaches  higher  suggests  aspect  t o i n c r e a s e as a s p e c t  on a s p e c t , measured i n  i s elevation  dependence of  f o r the (Fig.,  percent  13),  75  Alectoria  cn  elevation  3 . 0 5 5 ) . Much o f explainable fir/Salal than  by  significant  (t =  3.267, line  differences i n vegetation  community.  Douglas-  are  c o n s i s t e n t l y higher  i n percent  is  regression;  sites  are  c o n s i s t e n t l y l o w e r . Amounts o f A l e c t o r i a  i n percent  to  Fir/Oval-leaved  predicted  values. more o f  A l e c t o r i a than s i n g l e v a r i a b l e s .  The  -38.639 • 0.043 (x ) + 0.00016658 m ) (  (n = where y i s p e r c e n t  13;  Y  A l e c t o r i a , x,  5. 99)  i s elevation  i n meters,  i s potential solar radiation i n cal/cm /year,  has  2  The  (6)  z  S . / =  an  and r  (  (n =  13;  -  0.0903 ( x )  S y. x  2  =  * 0.3715 (x  3  )  (7)  5.06)  where y i s p e r c e n t  A l e c t o r i a , x , i s e l e v a t i o n i n meters, x  d e p a r t u r e f r o m due  s o u t h i n d e g r e e s , and  an  Forest  r  2  cf  2  regression  y = -14.043 + 0.0455 ( x )  has  the  regression y =  0.72.  =  Alectoria  Environmental v a r i a b l e s i n combination e x p l a i n  xz  y  the  close  Amabilis  o  by  T r a n s i t i o n s i t e s are  multiple  a  regression  predicted  variation  t  v a r i a t i o n around the  sites  Huckleberry on  the  i s highly  of  x  3  i s slope  in  2  is  degrees,  0.82.  measures  S c a t t e r g r a m s and measured  by  linear  densiometer,  percent  A l e c t o r i a are  percent  A l e c t o r i a on  significant  {t =  measured by  the  regressions  moosehorn, and  given  i n Figure  of c r o w n  v i s u a l estimate  14,  densiometer readings  3.3174, t  0  0  5  closure  Only the is  regression  of  statistically  = 3 . 0 5 5 ) . Crown c l o s u r e  d e n s i o m e t e r a c c o u n t s f o r 48%  on  of  the  as  variation in  40-1  40-1  40-i  ®  <  ©  ®  DC  30-  301  O 3 0  ©  I-  o  LU  _1  20H  <20-|  20-  •  A  h-  O ©  z  LU  v  o  1<H  10H  LX10'  v  LU  A  CL  «  70  CL  1 100  90  "80"  50  40  70  — i  80  90  CROWN CLOSURE Moosehorn  CROWN CLOSURE Densiometer  A TRANS  DF/S community  Figure 14.  60  v  AF/OH  a MOSS  Relationship between crown closure and percent Alectoria: n = 14; S . y  r  2  x  = 8.08 r  2  -9-  50  60  70  80  CROWN CLOSURE Visual o Unclassified  densiometer (y = 140.7 - 1.473 x;  = 0.48) (A); moosehorn (y = 42.859 - 0.375 x; n = 14; S . y  = 0.21) (B); v i s u a l estimates (y = 27.382 - 0.19005 x; n = 14; S . y  x  = 9.95  x  = 11.01 r  2  = 0.03) (C)  -i 90  77  percent  alectoria.  The  regression  statistically codcniinant Alectoria  (r  =  2  {Fig.  ( F i g , 16)  elevation  regression  and  (r  2  = 0.18), e x c e p t  tolerant  shade-intolerant  hemlock and regression  due  The  to the  height  i s also  of  0 0 5  not  the percent  significantly =  0,5324).  height  indicates  little  p o s s i b l y on J D o u q l a s - f i r / S a l a l I suspected  different  t r e e s on  separately.  f u r t h e r reduced.  Alectoria i s  i s a good p r e d i c t o r o f  crown l e n g t h / t r e e  Douglas-fir was  15),  ( F i g . 17).  might be  regression using  percent  (r = -.6180, r  relationship  the  on  o f crown l e n g t h / t r e e  Transition sites  and  area  0.64). T r e e h e i g h t  with  relationship sites  basal  significant  tree layer  correlated The  of  In  that the  combination  each s i t e , height  so  ratios  each c a s e ,  the  of  I  ccor shade-  repeated  of r  western 2  of  the  <  30-  o r— O  UJ _l  <  20-  \LU O  DC LU  °-  10-  80  90  100  120.  110  BASAL AREA m /ha 2  © DF/S community A TRANS community v AF/OH community B MOSS community o unclassified  Figure 15.  Relationship between basal area and percent (y = 36.907 - 0.235 x; ri = 14; S . y  Alectoria  = 10.71) r  2  x  = 0.08).  79  40n  20  30  40  HEIGHT OF CODOMINANT TREE LAYER  (M)  © DF/S community *  TRANS community  v AF/OH community B MOSS community o unclassified Figure 16.  Linear regression of percent Alectoria  on height of  codominant tree layer (y = 55.918 - 1.104 x; n = 14; S . y  x  = 6.75; r  2  = 0.64).  40  ©  30 H  DC  o o LU  -!20H I—  O  z  LU  O  DC LU  10 H  _—, .65  [—^ .70  C R O W N  L E N G T H / T R E E  —  r .75  H E I G H T  o DF/S community A TRANS community v AF/OH community a MOSS community o unclassified Figure 17.  Relationship between crown length/tree height and percent Alectoria S . y  x  = 10.14; r  (y = -49.68 + 0.93 x; n = 14; 2  = 0.18).  .80  81  Discussion  Tests of r e l a t i o n s h i p s v a r i a b l e s and solar  percent  radiation  quality  of sites  regression  of  s l o p e and  The  high r  are not  the importance of e l e v a t i o n ,  and  determining site.  For  each  of these  and  i s not  radiation aspect  of these  the The  variables i s  surprising, d e p e n d s on  considered  v a r i a b l e s . The  s l o p e , and  aspect  i n percent  6)  than  other.  that  the  accounts  t h a t of  suggests t h a t slope  solar  simply radiation  on  a  i n determining  s u b s t r a t e w i t h i n t h e t r e e canopy w e l l a s how  the  elevation  ( E q u a t i o n 7)  manner t h a n  incident  as  the  with fact  Alectoria  {Eguation  amounts o f  as  of  production.  instance, slope i s probably important  to s u n l i g h t ,  that  fact that separate regressions of  i n a more complex  potential  much l i c h e n  exposed  on  solar  solar radiation  aspect operate  for lichen  significant  variability  the h y p o t h e s i s  major d e t e r m i n a n t s  area  The  (0.82) o f s l o p e  2  more of t h e  elevation  how  significant.  physical.environment  support  are  i n the study  aspect  regression  and  elevation  o f e a c h v a r i a b l e on  indicates  for  alectoria  percent a l e c t o r i a  statistically  effect  and  between  at a s i t e i s  much s u n l i g h t  reaches  the  site. Solar radiation for lichen tends  productivity.  t o be  lower  than  because o f decreased the  effects  t o s e t an  At l o w e r  upper l i m i t  p r e d i c t e d by s o l a r  moisture,  a site-specific  may  on  elevations, lichen radiation,  potential abundance probably  v e g e t a t i o n community i n t e g r a t e s  o f s l o p e , a s p e c t , e l e v a t i o n , and  v a r i a b l e s on vegetation  appears  level.  The  enhance i t s p r e d i c t i v e  other  environmental  siterspecificity  v a l u e . For example,  of Plot  82  10 i s g e o g r a p h i c a l l y potential  developed  site;  draw and p r o b a b l y  t h a n e x p e c t e d on t h e b a s i s Douglas-fir/Salal  have t h e b e s t p o t e n t i a l higher elevations. solar  where s o l a r  radiation  high  abundance.  lichen like  vegetation  sites  i s high.  Conversely,  i s relatively  community,  forest  The r e l a t i o n s h i p s  that  would be more c l o s e l y  related  included  relationship lichen  After  visual  carrying  develop  to lichen  readings  abundance may  visual  out the  understory  trees.  The a p p a r e n t  relationship  considering  to explain  and s h o u l d  t h e poor r e s u l t s  either  poor and  of  considering between  a s measured by t h e d e n s i o m e t e r and p e r c e n t  difficult  closure  of understory  to the d i f f i c u l t y  c o v e r o f t h e main canopy w i t h o u t  the  mixed  e s t i m a t e s o f crown c l o s u r e  have been due  have  measures and  i n t h e i n s t r u m e n t r e a d i n g s . The  between  develop  relationships  abundance t h a n  because c o v e r  at  where  never  e s t i m a t e s o f crown  estimating  closure  especially  Tests of  In  appear t o  Moss s i t e s  between crown c l o s u r e  f i e l d w o r k , I expected  was  sites  percent A l e c t o r i a p r o d u c e d  are p u z z l i n g .  d e n s i c m e t e r o r moosehorn  solar  measures i n t e g r a t e  forest  trees  less  l o w , and p r o b a b l y  between  Alectoria  receives  productivity,  o f many e n v i r o n m e n t a l v a r i a b l e s .  percent  2, a B o s s s i t e , i s  of i t s gross p o s i t i o n .  effects  results.  Plot  and T r a n s i t i o n  for lichen  m e a s u r e s and  acre  lichen  T h i s i s probably because they  radiation  slightly  on a c o n v e x s l o p e and h a s t h u s  into a Douglas-fir/Salal  radiation  incident  2 and has o n l y  b u t has much h i g h e r  10 i s l o c a t e d  i n an a d j a c e n t  general,  to Plot  annual r a d i a t i o n ,  abundance. P l o t  located  close  be i n t e r p r e t e d  o f t h e o t h e r crown  crown Alectoria i s  cautiously, closure  83  measures. The height ratio  poor r e l a t i o n s h i p c f t h e  with percent t o be  penetrates solar  A l e c t o r i a was  ratio  surprising. I  a good i n d i c a t o r o f t h e the  o f crown l e n g t h  amount  and  expected  of s u n l i g h t  t r e e canopy, i n t e g r a t i n g the  radation  to  incident species  separately,  I expected the  Douglas-fir  t h a n f o r w e s t e r n hemlock, b e c a u s e D o u g l a s - f i r  shade-intolerant 1965). As trees  cn  exists  the  conditions survival the  of  be  data,  the  of the  probably  was  i t i s possible  lower part  small  that  for  (Krajina (usually  i t . Alternatively, light  of the  canopy  poor i n d i c a t o r c f l i g h t  as  measured by  conditions  f o r e s t measures s u g g e s t e d i s not  height  significantly  of  variability  the  higher  by  Bochelle•s  r e l a t e d to  codominant  tree  lichen  l a y e r accounts  fact  that  tree  growth,  like  many p h y s i c a l  The  the  Gaultheria  shallon association  -  was  noted  slopes,  index of  by  Douglas-fir K o j i m a and  which a r e  m o i s t u r e and  lichen  S a l a l Community Krajina  generally  (Krajina  growth. I n c r e a s i n g  of t h e  (1975). S t e e p ,  x e r i c or s u b x e r i c ,  nutrient conditions  w e s t e r n Hemlock z o n e  for  vegetation  e f f e c t s of  the  in  (1978)  the  siffiilar to  branch  i n l i c h e n abundance. T h i s r e l a t i o n s h i p  r e s u l t s from the  site  2-4  a relationship  community, i n t e g r a t e s low  is  r e f l e c t e d i n l i c h e n abundance.  area  abundance, but 64%  a  two  basal  plots),  better  W e s t e r n Hemlock Zone  unable to detect  i n the  c a n o p y , as Cf  10  I was  may  Coastal  sample s i z e f o r D o u g l a s - f i r  each  and  i n the  r e l a t i o n s h i p to hold  tree  the  that  e f f e c t s of  canopy s t r u c t u r e . C o n s i d e r i n g  tree  present  study  south-facing  provide  f o r t r e e growth  1969), b u t  variables.  adverse  i n the  good c o n d i t i o n s  elevation i s significantly  Coastal for  associated  84  with is  decreasing t r e e height  alsc  and i n c r e a s i n g l i c h e n a b u n d a n c e . I t  p o s s i b l e that slow-growing t r e e s provide  s u b s t r a t e s f o r l i c h e n s than hypothesis an  in  has n o t been e v a l u a t e d .  indicator  should this  area  rapidly  of l i c h e n  abundance  better  growing t r e e s , but Use o f f o r e s t  should  productivity  Bog s i t e s ,  f o r example, which o c c u r  useful  have been n o t e d  in identifying  the  study  not  permit  high  forest  a r e a and s i m i l a r testing  sites  areas.  for significant  which a r e p o t e n t i a l l y  with  high  Although  lichen  vegetation  communities i n order  for  production  lichen  community,  the Amabilis  size did  differences i n lichen  of t h e i r  i s suggested  abundance i n  t h e sample  abundance among v e g e t a t i o n c o m m u n i t i e s , a p r o b a b l e  ranking c f  increasing potential  by F i g u r e 9;  t h e Moss  Fir/Oval-leaved Huckleberry  Community,  T r a n s i t i o n Community, and t h e D o u g l a s - f i r / S a l a l Community .  V e g e t a t i o n community when c o n s i d e r e d  i s a useful indicator  i n conjunction  f e a t u r e s a s s o c i a t e d with lichen  abundance:  productivity  may  high i n s o l a t i o n  a s s o c i a t e d with  may  xeric  abundance  indicate  slopes, south  features that block  indicate  of l i c h e n  with e l e v a t i o n . T o p o g r a p h i c  moderate t o s t e e p  absence of t o p o g r a p h i c  is  i n the study  abundance.  Several factors  the  represented  have p o o r t r e e g r o w t h b u t would n o t n e c e s s a r i l y have  lichen  as  be done c a u t i o u s l y , and  n o t be e x t e n d e d t o v e g e t a t i o n c o m m u n i t i e s n o t study.  this  high  aspect,  s u n l i g h t . Low  good s i t e s f o r l i c h e n or s u b x e r i c c o n d i t i o n s .  growth  and  forest  when i t  85  V.  AVAILABILITY  The  third  together assess  OF  and  as t h e y  ABBOBEAL LICHENS AND  f o u r t h o b j e c t i v e s of the  are c l o s e l y r e l a t e d .  winter a v a i l a b i l i t y  measuring l i t t e r f a l l wished  and  inside  t o examine t h e  wind and  The  and  The  deer.  related  by  locating on  monitoring "visual  winter  deer  Bochelle litterfall found of  use  was  of the  conifer  {Jones  represented  1975, and  deposited during a  kg/ha on site. and  a low  Little  seasonal  in l i t t e r f a l l  site  to  p a t t e r n was  litter  sample s i z e  effective  i n e x c l u d i n g deer,  i n s i d e than  and  by  the f a c t  1978).  1973-1974.  important  source (lichens  approached or  exceeded  (s.l.)  period ranged  i n the  from  2.04  a mid-elevation monthly  litterfall  could  data, be  f e n c e s were c o n s t r u c t e d  p l o t s on  two  sites.  t h a t f e n c e s were n o t  q u a n t i f i e s of A l e c t o r i a  o u t s i d e the f e n c e d  He  forage  apparent  collection  the s a a l l  greater  and  two  monthly r a t e s o f  115.35 kg/ha on  ( B o c h e l l e 1978). C h i c k e n w i r e  a r o u n d some o f t h e  plots,  ftlectprla  no r e l a t i o n s h i p between s n o w f a l l and  determined  to  relate  Harestad  a relatively  180-day w i n t e r  elevation  to  winter of  amounts of r o o t e d f o r a g e . Q u a n t i t i e s o f spp.  also  abundance i n  amounts o f p o t e n t i a l  foliage)  by  o f w i n t e r h a b i t a t by  litterfall  area d u r i n g the  f o r deer:  to  p l o t s i n a r e a s f o r which i n f o r m a t i o n  available  i n the study  of f o r a g e l i c h e n s  to lichen  (1978) s t u d i e d c o m p o s i t i o n  food  green  use  estimate"  that l i t t e r f a l l  winter  and  deer  treated  o b j e c t i v e was  f o u r t h o b j e c t i v e was  was  are  DEES  deposition i n relation  to s e l e c t i o n  Habitat selection  BY  outside exclosures. I  t i m i n g of l i t t e r  precipitation.  study  third  utilization  abundance o f f o r a g e l i c h e n s  ways;  UTILIZATION  a r e a s , and  the  Despite entirely  ts.l.)  were  difference  86  was  statistically  significant  (p  < 0.05)  on  one  site.  87  Methods  Litterfall  measurement  litterfall p l o t s 3,  4, and  were a l s o  7,  and  measured  5.  measured d u r i n g t h e w i n t e r o f Because v i s u a l  performed  litterfall 6,  was  a t these  11)  three  by  under severe  mild  w i n t e r c o n d i t i o n s , and  Plot  5 was  l o c a t e d i n an  d u r i n g the f a l l  identified range.  by  G.H.  I t i s an  of  a t 573  in  f a l l , t h e doe  area  of Plot  basis  cn  m a.s.l.  4,  Plot  site.  s l o p e . The  doe  Plot  (unpublished  used  used  area  as  used  winter  on  (Harestad  an  1 9 7 8 ) , and  ENE-facing  near  as poor  km  area  t o the area  3 i s l o c a t e d i n an  data)  as good  Douglas-fir/Salal site  winter  a t 543  m  range.  an  2.8  km  position,  m a.s.l.  site  to the  on  on  t o be an  of P l o t  identified  range.  a.s.l.  3 at times  It i s a  by  fell  general the a  ESl-facing  d u r i n g much o f t h e  area  area  winter  which I expected,  topographic  remained i n t h i s  under  s l o p e . When snow  moved a p p r o x i m a t e l y 4 i s the s i t e  area  by a r a d i o s - c o l l a r e d , a d u l t  I t i s l o c a t e d a t 709  moved a b o u t 0.6  snowfall.  c o n d i t i o n s , an  a r e a not  (2,  1973-1974.  ( u n p u b l i s h e d data)  i t s v e g e t a t i o n and  Transition  but  area  plots  (1978)  Amabilis Fir/Oval-leaved Huckleberry  located late  an  of  relate  were c h o s e n t o r e p r e s e n t an  winter  1975  Jones  winter  plots  used  doe  deer  d u r i n g the  litterfall  p o s s i b l e to  where B o c h e l l e  on  abundance  four a d d i t i o n a l estimate  were l o c a t e d a t s i t e s  litterfall  The  estimates of l i c h e n  p l o t s , i t was  to standing crop,  1S76-1977  winter,  of heavy Jones  SE-facing  88  at  each l i t t e r f a l l  4 and o n e - h a l f  inch  plot  (11.4. mm)  were 3 t o 3.5 m e t e r s h i g h Twenty l i t t e r f a l l  a deer e x c l o s u r e  traps  nylon  were c o n s t r u c t e d  s e i n e n e t . The  and a p p r o x i m a t e l y  of nylon  screen  digging  i n t o t h e s l o p e on t h e u p h i l l  plan  Litterfall  i s given  each s i t e  i n Figure  s t a p l e d t o a frame o f  with  gussets.  soil  long.  litterfall  ;  plywood  side with  18 x 21 m e t e r s  T h € o n e meter, s g u a r e  1x2*s and r e i n f o r c e d a t t h e c o r n e r s  downhill  exclosures  were s y s t e m a t i c a l l y l o c a t e d i n s i d e and  twenty o u t s i d e each e x c l o s u r e , traps  was c o n s t r u c t e d o f  1/4 i n c h  t r a p s were p l a c e d  and p i e c e s  (6.3  mm}  h o r i z o n t a l l y by  s i d e and b u i l d i n g up t h e  o f wood, a g e n e r a l i z e d  18. P l o t l a y o u t  because of microtopographic  was s l i g h t l y  plot  modified  at  f e a t u r e s and p o s i t i o n s o f  trees. Litterfall  p l o t s were e s t a b l i s h e d i n November  determine the temporal litterfall each s i t e  traps  pattern of l i t t e r  were p l a c e d  i n January  and F e b r u a r y  were r a n d o m l y s e l e c t e d f r o m In from  the f i r s t  over  13 o f t h e o r i g i n a l  and l a b e l l e d ,  Later, the l i t t e r  was  x  including alectoria  E r y o r i a spp. - other  l i c h e n s ; and  - non-lichen  covered  collected  and d r i e d i n a  was s o r t e d  categories: - alectoria  t r a p s cn  1977. The t r a p s t o be  1977, t h e l i t t e r  t r a p s , bagged  hose d r y e r f o r s t o r a g e .  deposition, additional  t h e 20 i n s i d e t h e e x c l o s u r e .  week o f Hay  the l i t t e r f a l l  1976. To  litterfall.  s a r m e n t o s a and  into  three  O o o o o  O. O o o o o o o o o  O o o o o  O o o o o  y Anemometer  '  a • • •  • • •  • • a a  • a ri • a • • • • n a n  • • • •  • • • a a' • • •  •  •  • • •  Exclosure  O o o o o-  O o o o o  O o o 0 o  O .0 o o o o o o o o  Pellet group plot  Q •  10 meters  Litterfall trap  Figure 18".  Layout of l i t t e r f a l l plots, CO VD  90  The  litter  was  oven-dried  at 60-65°C f o r 24 hours and  weighed with a M e t t l e r P201Q e l e c t r i c s c a l e to the 0.1  nearest  g. The K r u s k a l - W a l l i s t e s t  (Conover 1971:  256)  was  used t o  compare l i t t e r f a l l g u a n t i t i e s on the t h r e e p l o t s . The  Mann-  Bhitney  litterfall  0 test  ( S i e g e l 1956;  q u a n t i t i e s i n s i d e and  116)  was  used to compare  outside exclosures.  To measure wind c o n d i t i o n s , t o t a l i z i n g anemometers were s e t up approximately  2.5  the e x c l o s u r e s and,  meters above the ground i n the timber where p o s s i b l e , i n adjacent c l e a r c u t s .  Anemometers were read during January, February, v i s i t s to the  measured during winter  present. Measurements were taken  snow depth f o r the  v i s i t s i f snow  was  was  used as a measure of  plot.,  P r e c i p i t a t i o n data were obtained from  study  May  at 10 s y s t e m a t i c a l l y l o c a t e d  the mean of these r e a d i n g s  Woss Camp and  March, and  plots.  Snow depth was  p o i n t s , and  near  weather s t a t i o n s at  at Port Hardy, approximately  85 km  north of the  area on the east c o a s t of Vancouver I s l a n d .  Habitat s e l e c t i o n Deer use of l i t t e r f a l l s i t e s during the winter of was  monitored by p e l l e t group counts and,  t r a c k counts. subplots  At each s i t e f i f t y  (comparable to 100  c l e a r e d i n November and  ft  2  when p o s s i b l e , by  9 m* c i r c u l a r p e l l e t group subplots)  were counted  again i n e a r l y May.  were t e s t e d f o r goodness of f i t with s e v e r a l d i s t r i b u t i o n s , and  normalized  1976-1977  and  P e l l e t group data freguency  by t r a n s f o r m a t i o n  (Sokal and E o h l f  91  1969:  380).  A n a l y s i s of v a r i a n c e  Duncan's New  (Sokal and Rohlf 1969:  M u l t i p l e Range T e s t  on the transformed  ( L i 1964:  data. Ihen snow was  p l o t s , t r a c k counts  270)  were performed  intersecting  p a r a l l e l to the contours and  and  the  counted.  In a d d i t i o n to P l o t s 3-5, p l o t s 8-10  100-  one  p e r p e n d i c u l a r to the contours. A l l deer t r a c k s c r o s s e d by t r a n s e c t s were  and  present d u r i n g v i s i t s to  were made by walking two  meter t r a n s e c t s , one  204)  12-14  l o c a t i o n s of v i s u a l  estimate  were chosen on the b a s i s of known deer  use  during p r e v i o u s winters. P l o t 8, an A m a b i l i s F i r / O v a l - l e a v e d Huckleberry  s i t e , was  used by a r a d i o - c o l l a r e d a d u l t buck during  mild winter c o n d i t i o n s (Harestad 1S78). P l o t 9, a T r a n s i t i o n s i t e , was  used by the same animal  severe. P l o t s 13 and Huckleberry  when winter weather  14, T r a n s i t i o n and  was  Amabilis F i r / O v a l - l e a v e d  s i t e s r e s p e c t i v e l y , represented adjacent h a b i t a t s  which the buck used l i t t l e  or not at a l l during winter,  he apparently t r a v e l l e d through  them. P l o t  although  10 i s a Douglas-  f i r / S a l a l area which a r a d i o - c o l l a r e d a d u l t doe  moved i n t o at  the time of an autumn s n o w f a l l . I t i s l o c a t e d i n an area i d e n t i f i e d by Jones (unpublished data) as good winter range which r e c e i v e s high deer use. P l o t was  12, which was  used as winter range by a two-year-old  unclassified,  r a d i o - c o l l a r e d buck.  92  Jesuits  Total l i t t e r f a l l  guantities  Q u a n t i t i e s of the v a r i o u s i n s i d e exclosures Because l i t t e r  on the  d i s t r i b u t i o n s of the (e.g.  Q=  components  deposited  s i t e s are given i n Table  c o l l e c t i o n periods  r e s u l t s were adjusted  right  three  litter  v a r i e d s l i g h t l y among s i t e s ,  f o r a 180-day p e r i o d . Host of litterfall  the  samples were s t r o n g l y skewed  3.364). For t h i s reason confidence i n t e r v a l s  were computed around the median using the nonparametric method  (Conover 1971:110). The  from the  medians d i f f e r e d  on the three  considerably  256)  ( s . l . ) g u a n t i t i e s ranged from 31.9  s i t e s ; r e s u l t s of the  151.2  Kruskal-Hallis test  (T = 47.59; T  o o s  kg/ha  (Conover  = 5.99) . Non-lichen  litterfall  ranged from 867.5 to  litterfall  from 978.4 to 1071.1 kg/ha. D i f f e r e n c e s  ncn-lichen  litterfall  s i g n i f i c a n t but total:  to  i n d i c a t e d highly s i g n i f i c a n t differences i n A l e c t o r i a  l e v e l s among s i t e s  in  guantile  means, as i s expected i n skewed d i s t r i b u t i o n s .  Alectoria  1971:  VII.  T = 6.96;  and  1005.0 kg/ha, and  total l i t t e r f a l l  at a much lower l e v e l T  0 O 5  =  l i c h e n s ranged from 34.2  to 45.5  were not s i g n i f i c a n t (T = 0.71;  of  among s i t e s  were a l s o  (non-lichen:  5,99). Q u a n t i t i e s  total  T = 10.33;  non-Alectorioid  kg/faa; d i f f e r e n c e s among p l o t s T  0 0 5  =  5.99).  93  TABLE VII. L i t t e r f a l l q u a n t i t i e s i n s i d e exclosures  L i t t e r f a l l Components Aleotovia  Other Lichen  Non-Lichen  Total  P l o t 3 (DF/S) Mean f o r 180-day period (kg/ha) Median f o r 180-day period (kg/ha) 95% confidence i n t e r v a l around median  69.9  41.0  867.5  978.4  62.9  28.5  662.9  786.9  60.0-81.9  22.6-40.3  556.7-759.4  46.5  868.0  1065.7  31.5  737.7  890.2  22.6-55.1  537.1-784.9  31.9  34.2  1005.0  1071.1  25.9  26.9  1019.4  1086.5  629.5-908.9  Plot 4 ( U n c l a s s i f i e d ) Mean f o r 180-day period (kg/ha) 151.2 Median f o r 180-day period (kg/ha) 130.8 95% confidence i n t e r v a l around median 116.1-166.2  694.4-1012.1  P l o t 5 (AF/OH) Mean f o r 180-day period (kg/ha) Median f o r 180-day period (kg/ha) 95% confidence i n t e r v a l around median  22.5-40.5  20.9-43.8  762.8-1197.3  800.9-1254.6  94  L i c h e n l i t t e r f a l l i n r e l a t i o n to l i c h e n biomass Table V I I I presents l i c h e n l i t t e r f a l l data  f o r seven s i t e s . The  Bochelle  1973-1974 data were c o l l e c t e d  (1978), Data from the two  s u b j e c t to g r a z i n g  by  s t u d i e s must be compared  c a u t i o u s l y , as d i f f e r e n c e s i n sampling between years probably  and l i c h e n biomass  methods and  i n weather  i n f l u e n c e d r e s u l t s , B o c h e l l e * s p l o t s were  (with the e x c e p t i o n of ten l i t t e r f a l l  on P l o t 7) whereas the 1976-1977 data are based on p l o t s . Despite these  traps  exclosed  d i f f e r e n c e s , ranges of A l e c t o r i a  litterfall  values are comparable. A l e c t o r i a l i t t e r f a l l q u a n t i t i e s appear t o be r e l a t e d t o A l e c t o r i a biomass i n the canopy, as i n d i c a t e d by the narrow range o f r a t i o s of l i t t e r f a l l to standing Percentages of l i t t e r f a l l from 7,6  to 9,4%,  ungulates. higher:  a  percentages  (1978) data range  based on the present  study  are  - 16.1%.  L i t t e r d e p o s i t i o n i n r e l a t i o n to time and D a i l y d e p o s i t i o n of A l e c t o r i a l i t t e r , and  crop.  but f a l l e n l i c h e n s were s u b j e c t t o removal by  Litterfall  10.5  based on B o c h e l l e s  relatively  total l i t t e r  weather other l i c h e n  litter,  are given i n F i g u r e s 19-21. The e a r l y winter  p e r i o d i n c l u d e s November to mid-January; mid-winter, mid-January to l a t e February; week of May.  and  l a t e winter,  l a t e February  to the  first  On a l l t h r e e p l o t s , d e p o s i t i o n r a t e s of both  components and of t o t a l l i t t e r  litter  i n c r e a s e d c o n s i s t e n t l y during  winter. D e p o s i t i o n r a t e s of A l e c t o r i a i n c r e a s e d from e a r l y to l a t e winter by f a c t o r s o f 12.7,  9.5,  and 25,6  on the t h r e e  p l o t s , whereas d e p o s i t i o n r a t e s of t o t a l l i t t e r f a l l  increased  the  95  TABLE V I I I . litterfall  Alectoria  Plot  Alectoria standi ng crop (kg/ha)  in r e l a t i o n to Alectoria  Alectoria litterfall 1976-77 (kg/ha)  standing crop  Alectoria litterfall 1973-74 (kg/ha)  Litterfall/ Standing Crop (%)  14.2  8.1  1  2  175  3  665  69.9  10.5  4  942  151.2  16.1  5  303  31.9  10.5  6  975  74.7  7.7  7  140  13.2  9.4  11  1516  115.3  7.6  From Rochelle 1978.  96  > TJ 05 JZ  2000-  cn i1  _i _j  1500-  < LL DC LU  h-  1000-  h-  _1  < 500-  DC  o ho  LU  _J  [fl  <  M PLOT  J5L  L  3  M 4  L  M 5  mean  median and 95% confidence l i m i t s  Figure 19.  Deposition rates of Alectoria  l i t t e r i n e a r l y winter  mid-winter (M), and l a t e winter ( L ) , 1976-1977.  (E),  97  1500-1  < LL DC  LU  Z LU X  o  ^IOOOH CO  •a cc sz  cn 500H  cc LU  0  X  o  PLOT  M  M  M  .3  4  5  L  mean  median and 95% confidence l i m i t s  Figure 20;.  Deposition rates of other ( n o n - A l e c t o r i o i d ) l i c h e n l i t t e r i n e a r l y winter (E), mid-winter (M), and l a t e winter ( L ) , 1976-1977.  98 !  tOOOOi  8000 CO CO  0  6000H  D)  < ^ 4C00LU \z  _J _ j 2000H _l  0  < E  P  L  O  T  M  L  3  E  M  L  E  4  M  L  5  mean  median and 95% confidence  Figure 21.  limits  Deposition rates of a l l l i t t e r i n e a r l y winter ( E ) , midwinter (M), and l a t e winter ( L ) , 1976-1977.  99  2.2,  only  followed of  3.7,  and 2.7 t i m e s .  an i n t e r m e d i a t e  Deposition  rates o f other  rate of increase,  lichens  i n c r e a s i n g hy f a c t o r s  7 . 7 , 8.a, and 9.3. average p r e c i p i t a t i o n  r a t e s a t B o s s Camp and P o r t  d u r i n g e a r l y , mid- a n d l a t e  winter  Hardy  o f 1976-1977 a r e g i v e n i n  F i g u r e 22. P r e c i p i t a t i o n  was g r e a t e s t d u r i n g e a r l y w i n t e r and  least  No r e l a t i o n s h i p  and  during  mid-winter.  litterfall The  winter  snowfall period. period  o f 1976-1977 was u n u s u a l l y i n late  March  was r e l a t e d  t o that storm, but that  the exclosures  the  The o n l y  the l a t e  litterfall  during  possibility  this  c a n n o t be  and i n a d j a c e n t  clearcuts during to Plot  4.  the three Unfortunately,  anemometer l o c a t e d i n t h e c l e a r c u t by P l o t 5 d i d n o t properly,  readings  probably  Hind  as t h e c u p s d i d n o t s p i n f r e e l y , underestimated  c o n d i t i o n s near  ground  the recorded  t r u e amounts o f wind., level  i n c l e a r c u t s should  a p p r o x i m a t e wind c o n d i t i o n s a t t h e t o p o f t h e f o r e s t although  the  winter  amounts o f wind i n t h e t i m b e r  T h e r e i s no c l e a r c u t a d j a c e n t  function  116,  major  a v a i l a b l e data.  F i g u r e 23 s u m m a r i z e s r e l a t i v e  periods.  mild.  1977, d u r i n g  I t i s possible that the increased  with  precipitation  i s apparent.  occurred  evaluated  by  between  more t u r b u l e n c e  274, 312). timber  clearcut indices reliable  i s expected  i n t h e canopy  canopy,  (Geiger  I had a n t i c i p a t e d t h a t anemometer r e a d i n g s  1966: from  would be g e n e r a l l y p r o p o r t i o n a t e t o c o r r e s p o n d i n g  readings,  so t h a t timber  readings  could  be c o n s i d e r e d  t o wind c o n d i t i o n s a t t h e t o p o f t h e c a n o p y . readings  Without  f r o m t h e c l e a r c u t a t P l o t 5, t h e a s s u m p t i o n  c a n n o t be j u s t i f i e d .  100  o  10  — E O E  5H  UJ  DC  0_  Figure 22.  E M L Port Hardy  E M Woss  P r e c i p i t a t i o n at Port Hardy and Woss Camp i n e a r l y winter (E), mid-winter (M), and l a t e winter ( L ) , 1976-1977.  101 I  DC  LU LU  o LU  zz. <  40CO CO  20H  c  DC  M  £ PLOT  3  LU  L  M  M  4  5  LU  \— LU  o  80-  w CO  < ~CO  D  O  1  40  DC  <  LU _J  O  E  M  PLOT  Figure 23.  E  L  3  M  L  5  R e l a t i v e amounts of wind i n mature timber (A) and c l e a r c u t s (B) i n e a r l y winter (E), mid-winter (M), and l a t e winter  ( L ) ,  1 9 7 6 - 1 9 7 7 .  102  Anemometer r a t e s on a l l s i t e s tended t o be lower i n e a r l y winter than i n mid- o r l a t e winter. In the c l e a r c u t by P l o t 3, anemometer r a t e s i n c r e a s e d as winter progressed, but changes i n litterfall  r a t e s were much g r e a t e r than changes i n wind r a t e s as  measured by anemometers. The data do not preclude a r e l a t i o n s h i p between l i t t e r f a l l but  r a t e s and wind as measured by anemometers,  they do not demonstrate a d e f i n i t e  relationship.  U t i l i z a t i o n of l i t t e r f a l l The l i t t e r  samples from 15 of the 20 l i t t e r f a l l  traps  o u t s i d e the e x c l o s u r e on P l o t 5 were i n a d v e r t a n t l y exposed to moisture during s t o r a g e , and began t o decompose. P a r t i a l decomposition a f f e c t e d t h e A l e c t o r i a sarmentosa and B r y o r i a spp. more than the other l i c h e n s , and made i t i m p o s s i b l e to separate them adequately from the r e s t of the sample. The MannWhitney  U test  ( S i e g e l 1956:116) was used t o determine  weights of the p a r t i a l l y  whether  decomposed samples were s i g n i f i c a n t l y  d i f f e r e n t from weights of the u n a f f e c t e d samples., R e s u l t s i n d i c a t e d t h a t d i f f e r e n c e s i n weights approached only i n the A l e c t o r i a category <U = 16.5; U  0 o S  significance  = 14.0).*  T h e r e f o r e , the p a r t i a l l y decomposed samples o f A l e c t o r i a . but not of  of other l i t t e r litterfall  components were e l i m i n a t e d from comparisons  i n s i d e and o u t s i d e the e x c l o s u r e .  In the Mann-Whitney U t e s t , the n u l l hypothesis i s r e j e c t e d i f the c a l c u l a t e d value f o r U i s s m a l l e r than the t a b u l a t e d value. 1  103  Q u a n t i t i e s of l i t t e r f a l l  i n s i d e and outside e x c l o s u r e s  compared i n Table IX. R e s u l t s of t h e Mann-Whitney U t e s t 1956:  116)  i n d i c a t e that q u a n t i t i e s of a l e c t o r i a  significantly  greater i n s i d e than  three s i t e s , d e s p i t e the reduced  were  sample s i z e at P l o t  e x c l o s u r e s d i d not d i f f e r s i g n i f i c a n t l y  5.  outside  at any s i t e s , and  i n one  were g r e a t e r o u t s i d e than i n s i d e . Amounts of n o n - l i c h e n  litterfall in  (Siegel  outside e x c l o s u r e s on a l l  Q u a n t i t i e s of n o n - A l e c t o r i o i d l i c h e n s i n s i d e and  case  are  were a l s o s i m i l a r i n s i d e and o u t s i d e e x c l o s u r e s  and,  cne case, g u a n t i t i e s outside were g r e a t e r than q u a n t i t i e s  i n s i d e . The  s i g n i f i c a n t d i f f e r e n c e between t o t a l  i n s i d e and o u t s i d e the e x c l o s u r e at P l o t 4 was  litterfall  due  p a r t l y to the  larqe difference i n A l e c t o r i a q u a n t i t i e s . A comparison of A l e c t o r i a with the other components suggests  litterfall  that differences i n A l e c t o r i a g u a n t i t i e s  i n s i d e and o u t s i d e e x c l o s u r e s were due  to removal by  and not t o f o r t u i t o u s s p a t i a l p a t t e r n s of l i t t e r  herbivores,  deposition.  B e s u l t s a l s o i n d i c a t e t h a t consumption of n o n - A l e c t o r i o i d l i c h e n s was Bochelle  minimal. These r e s u l t s are c o n s i s t e n t with those  (1978)., who  l i c h e n s , was litterfall  found t h a t A l e c t o r i a  (s. 1.) , but not  a major forage item i n winter. As  was  not separated  other  non-lichen  i n t o components, no c o n c l u s i o n s  be drawn about consumption of l i t t e r  of  components other  may  than  lichens. The  d i f f e r e n c e between A l e c t o r i a q u a n t i t i e s i n s i d e  o u t s i d e each e x c l o s u r e p r o v i d e s an approximation u t i l i z a t i o n at each s i t e  of  and  Alectoria  ( F i g . 24). Using the Mann-Whitney  s t a t i s t i c t o c a l c u l a t e 95% confidence l i m i t s f o r the d i f f e r e n c e  TABLE IX.  L i t t e r f a l l quantities i n s i d e and outside exclosures  Alectoria kg/ha/180 days  Other Lichen kg/ha/180 days  Non-Lichen kg/ha/180 days  Total kg/ha/180 days  Plot 3 Inside exclosure  69.9  41.0  867.5  878.4  Outside exclosure  44.2  37.3  1108.8  1190.7  Mann-Whitney U  47  140  151  186  1  1  1  1  Plot 4 Inside exclosure  151.2  Outside exclosure  71.2  Mann-Whitney U  20.5  868.0  46.5  793.6  43.6 1  186.5  1065.7  1  141.5  908.5 1  127  1  Plot 5 Inside exclosure  31.9  34.2  1005.0  1071 .1  Outside exclosure  15.2  55.4  924.1  994.9  Mann-Whitney U  182  155  146  1  U.Q25  =127  2  U.025 = 20  S  denotes s t a t i s t i c a l s i g n i f i c a n c e at p < .025.  163  1  1  1  105 |  100 - i  80  z: O  H  N co 3] TJ 60 H  o  3  03  <  5  cr \  4o-|  (~\ CD  o LU _J <  20H  PLOT  Figure 24.  Alectoria confidence  u t i l i z a t i o n on Plots 3, 4, and 5, with 95% limits.  106  between two means (Conover  1971:238), u t i l i z a t i o n  at Plot  25.7  (13.5 t o 35.3) kg/ha/180 d a y s ;  utilization  80.0  (46.3 t o 92.5) kg/ha/180 d a y s ;  and u t i l i z a t i o n  was  16.7 (15,.8 t o 28.1) kg/ha/180 d a y s .  53%, and 52% u t i l i z a t i o n  Deer u s e o f l i t t e r f a l l Results  of p e l l e t  winter range than mild per  winter.  at Plot  31%,  is. 1,) .  group c o u n t s  o f non-winter  indicate  range,  g r e a t e r use o f  d e s p i t e the extremely  Mean v a l u e s o f 1.12, 0.80, and 0.40 p e l l e t  (Plot  5  plots  3), the mild winter range s i t e  the negative  4 was  This represented  o f a v a i l a b l e Jklectoria,  s u b p l o t were p r e s e n t on t h e s e v e r e  range s i t e  at Plot  3 was  (Plot  5), respectively.  winter range s i t e  4 ) , and t h e p o o r  The d a t a  binomial d i s t r i b u t i o n ,  groups (Plot  winter  were f o u n d  to f i t  and a l o g a r i t h m i c  t r a n s f o r m a t i o n was a p p l i e d . R e s u l t s o f t h e a n a l y s i s o f v a r i a n c e indicated  a highly significant  d i f f e r e n c e among p l o t s  B e s u l t s o f Duncan's New M u l t i p l e  Range T e s t showed t h a t t h e  means o f P l o t s 3 and 4 d i d n o t d i f f e r mean o f P l o t  5 differed  Two e l k p e l l e t  from  g r o u p s were f o u n d  some w i n t e r  elk  g r o u p s were a l s o n o t e d  pellet  though  none a p p e a r e d  observed any  at Plot  lichen  probably  significantly,  t h e ethers  indicating  use o f t h e area  t o be f r e s h  but the  (p<0.05).  inside subplots at Plot by R o o s e v e l t  i n the v i c i n i t y  5. T h i s low l e v e l  4,  e l k . A few  of Plot  3,  i n s p r i n g 1977. No e l k s i g n o f e l k use i n d i c a t e s  r e m o v a l was due t o e l k r a t h e r t h a n  minor.  (p<0.01).  to deer,  was  that i f i t was  107  The  most s i g n i f i c a n t s n o w f a l l o f the winter  March 27tfc, 1977, Track  occurred cn  counts were made and snow depths  measured on March 28 and 29. R e s u l t s  (Table X) demonstrate that  during t h i s b r i e f period of snow, deer were concentrated  i n the  area thought t o r e p r e s e n t severe winter range, average snow depth a t the s i t e was only 7 cm. P l o t #, t h e m i l d winter  range  area, had much deeper snow (53 cm) and no s i g n s of deer use a f t e r the s n o w f a l l . The area around P l o t 5, where snow depth was intermediate  (26 cm), was e v i d e n t l y being used by one animal.  Lichen abundance and deer use A v a i l a b l e i n f o r m a t i o n on the s t a t u s of l i c h e n p l o t s as winter deer h a b i t a t i s summarized i n Table XI. I t i s d i f f i c u l t to  draw f i r m c o n c l u s i o n s from t h e data a v a i l a b l e here because  a r b o r e a l l i c h e n s r e p r e s e n t only p a r t of the a v a i l a b l e f o r a g e . A v a i l a b i l i t y of rooted f o r a g e , p a r t i c u l a r l y shrubs,  was not  measured as part o f t h i s study, but i s undoubtedly an important f a c t o r i n winter  habitat selection.,  With the exception o f P l o t s 8 and 12, areas used i n winter were moderate or high i n l i c h e n abundance, whereas areas not s e l e c t e d i n winter were low i n l i c h e n abundance. Shrubs are high and  dense at P l o t 8, and are probably the major source of forage  for  wintering deer. S e l e c t i o n o f P l o t  difficult  12 as winter h a b i t a t i s  t o e x p l a i n , as n e i t h e r l i c h e n s nor shrubs are  abundant. The tendency f o r deer t o use moderate or high areas i n winter, coupled  lichen  with the data on u t i l i z a t i o n of  l i c h e n s , s t r o n g l y suggests  that l i c h e n a v a i l a b i l i t y i s a factor  i n f l u e n c i n g winter h a b i t a t s e l e c t i o n .  TABLE X.  Deer use and snow depths at Plots 3, 4, and 5, March 28-29, 1977  Plot  4  Date  T r a c k  Count  D e e r  Seen  S n o w  Depth (cm)  March 28  30  March 28  0  0  53  March 29  0  1  26  109  TABLE' XI. Habitat s e l e c t i o n and l i c h e n biomass  Plot  Summary of Habitat Selection Information  Lichen Biomass (kg/ha)  3  665  4  942  Used i n severe winter by OFL 6 0 , good winter range . 1  2  Used i n mild winter by OFL 60 . 1  5  303  Used i n f a l l by OFL 6 0 , poor winter range . 1  2  9  800  Used in severe winter by OFL 62 . 1  8  289  Used i n mild winter by OFL 6 2 .  14  176  Used o c c a s i o n a l l y i n mild winter by OFL 6 2 .  1  1  13  107  Not used i n winter by OFL 62^.  10  521  Used i n f a l l when snow present by OFL 68 , good winter range . 2  21  12  Used i n Winter by OFL 6 1 .  1  Harestad 1978.  2  Jones 1975 and unpublished data.  l  110  Discussion  Litterfall The l i c h e n l i t t e r f a l l  t o t a l s obtained  i n t h i s study a r e  compared with r e s u l t s of other s t u d i e s i n Table XII. The range of values r e p o r t e d here i s somewhat higher than.that r e p o r t e d by Eochelle  (1976) and Kale  (unpublished data) V i n t h e i r s t u d i e s on  Vancouver I s l a n d , but i n both o f t h e l a t t e r cases l i t t e r f a l l was s u b j e c t t o consumption by h e r b i v o r e s . The l i t t e r f a l l reported by Denison  values  (1973) f o r L o b a r i a oregana and Andre e t a l .  (1975) f o r a l l a r b o r e a l l i c h e n s f a l l w i t h i n the range of values reported here. The maximum value f o r a l l a r b o r e a l l i c h e n s r e p o r t e d by Schroeder  (1974) f o r a subalpine f o r e s t i n i n t e r i o r  B r i t i s h Columbia exceeds t h e maximum value found area by more than 75 kg/ha. As Schroeder's litterfall  were based  underestimate  i n the study  annual r a t e s o f  on summer measurements, they may  true v a l u e s .  L i t t e r f a l l taken as a percentage o f standing c r o p  ranged  from a minimum of 1.2% (Andre e t a l . 1975) to a maximum of 22,9% (Denison  1973). The v a l u e s r e p o r t e d i n the present study a r e  intermediate.  * I am g r a t e f u l t o W. Kale and D. Hebert of the B r i t i s h Columbia F i s h and W i l d l i f e Branch f o r permission to use these data.  Ill TABLE XII.  Area  Source  Comparison of reported quantities of lichen l i t t e r f a l l  Forest Type  Lichens  Denison 1973  Western Oregon  Pseudotsuga menziesii  Lobaria  Schroeder 1974  S e l k i r k Mountains, B.C. and U.S.A.  Picea Abies Larix  A l l arboreal lichens  Andre et a l . 1975  France  Abies alba  A l l arboreal l i c h e n s ; mainly Pseudevernia furfuracea  W. Kale (unpubl i s h e d data)  Northwest Bay, Vancouver Island  Pseudotsuga menziesii  A l l arboreal lichens  engelmanniilasiocarpa; occidentalis  Nimpkish V a l l e y , Vancouver Island  Tsuga heterophyllaPseudotsuga menziesiiThuja plicata  This study  1  Based on summer rates extended to 1 year.  2  Based on 180-day winter p e r i o d .  Tsuga heterophyllaPseudotsuga menziesiiThuja plicata  89.7  392.3-504.4  17.8-22.!  ^  ^  1  gg -j  1040.0  7.2- 9.5%  1.4-105.92  and 0.3- 99.4  2  and Bryoria  spp,  A l l arboreal lichens Alectoria  Litterfall As % of Standi ng Crop  7.8-273.9  A l l arboreal lichens Alectoria  Standing Crop (kg/ha)  oregana  Alectoria, Bryoria, Usnea spp. Rochelle (1976) and t h i s study  Lichen Litterfall kg/ha/year  and Bryoria  spp.  40.5-162.6  2  13.2-115.3  2  66.1-197.7  2  31.9-151.2  2  7.6- 9.4%  10.5-16.1%  112  Seme workers have assumed t h a t epiphyte  biomass i s i n a  steady s t a t e i n an old-growth f o r e s t ; growth i s balanced litterfall,  i n - s i t u decomposition,  and consumption by  (Pike et a l . 1972). I f t h i s i s t r u e , then annual, provides an estimate  To my  herbivores  litterfall  of annual turnover and thereby  growth, s i n c e the other  by  annual  pathways are probably r e l a t i v e l y  minor.  knowledge, i t has not been demonstrated e m p i r i c a l l y t h a t  there i s no net change i n epiphyte  biomass i n old-growth  forests. The r e s u l t s of t h i s study p r o v i d e i n f o r m a t i o n about a v a i l a b i l i t y , u t i l i z a t i o n , and Alectoria  (s.l.)  litterfall  i s provided on v a r i a t i o n increase in l i t t e r f a l l  r e l a t i o n s h i p to s t a n d i n g crop of  i n a s i n g l e winter. No  in l i t t e r f a l l  among years. A r e g u l a r  r a t e s d u r i n g the winter was  the present study, but i t i s not  information  apparent i n  known whether t h a t i s a general  p a t t e r n . F i n a l l y , no r e l a t i o n s h i p between weather and litterfall  r a t e s of  could be e s t a b l i s h e d .  Other s t u d i e s provide i n f o r m a t i o n on v a r i a t i o n i n litterfall  g u a n t i t i e s between years. The  value f o r l i c h e n  litterfall  reported by Andre e t a l . (1975) f o r 1970-1971 was  of  t h a t measured at the same s i t e i n 1972-1973; t o t a l  in  1970-1971 was  91% of t o t a l l i t t e r f a l l  study a r e a , A l e c t o r i a  (s.l.) l i t t e r f a l l  c o n s i s t e n t l y lower d u r i n g the extremely than i n 1975-1976, averaging only 3H% Other l i c h e n s and t o t a l l i t t e r f a l l  litterfall  i n 1972-1973. In  Kale's  ( i n c l u d i n g : Usnea )  was  mild winter on  1976-77  of the 1975-1976 values.  were both lower i n 1S76-1S77  on 4 of the 5 s i t e s s t u d i e d , averaging r e s p e c t i v e l y , of the  76%  U7% and  55%,  1975-1976 v a l u e s . E v i d e n t l y , v a r i a t i o n i n  113  l i c h e n l i t t e r f a l l from one I f the lower l i t t e r f a l l were a s s o c i a t e d with  year t o another can be  values reported by Kale f o r 1976-1977  the unusually  p o s s i b l e t h a t the values reported i s usual f o r the study Timing of l i t t e r important  mild winter, then i t i s i n t h i s study  are lower than  area. d e p o s i t i o n , as w e l l as q u a n t i t y , i s  to w i n t e r i n g deer. I f l i t t e r f a l l  l a t e winter,  considerable.  rates are h i g h e s t i n  as observed i n the present study, then maximum  a v a i l a b i l i t y of l i t t e r f a l l  probably  maximum s t r e s s f o r the animals. (unpublished)  presented  c o i n c i d e s with the  Data of B o c h e l l e  period of  (1978) and  f o r comparison i n F i g u r e s 25 and  i n d i c a t e that the p a t t e r n may  not  not only f l u c t u a t e d e r r a t i c a l l y  be general. L i t t e r f a l l  during the course  temporal patterns v a r i e d from one  Kale  26 rates  of winter,  s i t e to another w i t h i n a  but  study  area. It  seems l i k e l y t h a t l i c h e n l i t t e r f a l l r a t e s are a f u n c t i o n  of s t a n d i n g crop and site-specific,  of weather f a c t o r s , which may  Abee and  Lavender  (1972) reported  be h i g h l y  that most  l i t t e r i n a mature D o u g l a s - f i r stand i n Oregon f e l l  during  winter and a t t r i b u t e d t h i s p a t t e r n to breakage of branches to snow. Pike et a l . (1972) suggested t h a t the i n c r e a s e d of epiphytes  when wet  might be important  breakage. Other authors  (Cowan 1945,  l i c h e n l i t t e r f a l l t o snow and that l i t t e r f a l l i s probably  i n causing  Gates 1968)  wind. B o c h e l l e  due  weight  branch  attributed  (1978) observed  i n f l u e n c e d most by  intense  i n d i v i d u a l storms r a t h e r than o v e r a l l weather p a t t e r n s .  5900  150CH CO  •a CO  O)  ioooH  cc LU I—  LU  o <  500-  cc  o LL  r; N-D-J-F-M-A SITE  1  CIIDL  N-D-J-F-M-A 2  N-D-J-F-M-A 3  D-J-F-M-A  J-F-M-A  4  5  Figure 25. Monthly deposition rates of forage l i t t e r (lichens and green c o n i f e r f o l i a g e ) i n the study area, 1973-1974 (Rochelle 1978).  O T H E R  A L E C T O R I A  g/ha /day  800H  600H  j - 400-j  400-  6 0  kg/ha/day  g/ha/day  800H  * °-  6  20H 10H  200H  200A  O  N  D  J  F  O  M  800-  800-  600H  600-  UJ I— 400-]  400-  N  D  J  F  O  M  N  D  J  F  M  30-1  20-  CO  10-  200-  200-  O  N  D  J  F  O  M  800-  800  600-  600-  ^ 40000 200-  400-  T-  4  30H  CO  CO  N O N - L I C H E N  L I C H E N  J O N  F  D  J  F  800-  800-  co 600-  600-  1—  N  D  J  F M  F  30H  20H 10H  O  M  O  M  200-  L  D J  N  N  D  J  F  M  O  N  D  J  O  N  D  J  M  30H 20-  400-  £ 400-  ion  200-  200 O  N  Figure 26.  D  J  .F  M  O  N  D  J  F  M  Monthly deposition rates of three l i t t e r f a l l  F  components  at Northwest Bay, 1975-1976 (Kale, unpublished data).  M  116  The i n t e r a c t i o n of c l i m a t i c f a c t o r s i s probably in  producing l i c h e n l i t t e r f a l l .  Usnea t h a l l i and had about  were about  Von Schrenk  important  (1898) found  3 times heavier when wet  that  than when dry,  25% as much t e n s i l e s t r e n g t h . In a 20 f o o t  (6.1  m)  wind t u n n e l dry t h a l l i d i d not break at v e l o c i t i e s of 124 km/hr, but wet wind may  t h a l l i began to fragment  at wind v e l o c i t i e s of 80 km/hr.  be more e f f e c t i v e i n producing l i c h e n l i t t e r f a l l  in a  rainstorm or snowstorm than under dry c o n d i t i o n s . I suspect  that  d e t a i l e d , f r e q u e n t , and s i t e - s p e c i f i c monitoring of weather c o n d i t i o n s would be necessary t o demonstrate litterfall  t h e i r i n f l u e n c e on  rates.  Deer use and l i c h e n abundance assessment of deer use of the l i t t e r f a l l  plots indicated  higher deer use and higher l i c h e n abundance i n t h e areas  thought  to  poor  represent winter range  winter range.  than i n the area thought  to be  E v a l u a t i o n of l i c h e n abundance i n areas  selected  by r a d i o - c o l l a r e d deer i n winter i n d i c a t e d a tendency f o r animals to use areas having r e l a t i v e l y There i s no one-to-one correspondence  high l i c h e n abundance. between winter range  a v a i l a b i l i t y of forage l i c h e n s . Rather, I suggest t h a t s e l e c t areas where food a v a i l a b i l i t y  and  deer  i s high, and t h a t a r b o r e a l  l i c h e n s are an important component of winter food. available alectoria for 80.0  ( s . l . ) ranged  from 31.9  a 180-day winter p e r i o d . U t i l i z a t i o n ranged  to 151.2 from  kg/ha  16.7  to  kg/ha f o r the same p e r i o d . Q u a n t i t i e s of a v a i l a b l e rooted  forage were not measured as part o f t h i s study, but R o c h e l l e (1978) r e p o r t e d an average of 42 kg/ha on low e l e v a t i o n  sites  117  and  149 kg/ha  on m i d - e l e v a t i o n  s i t e s i n t h e same s t u d y  R e s u l t s o f t h i s study  support  of  available  forage  potentially  available  rooted  forage,  available  Alectoria  B o c h e l l e * s c o n c l u s i o n t h a t amounts in litterfall  and d e m o n s t r a t e  litterfall  area.  a r e comparable t o  t h a t as much a s 53% o f  may be u t i l i z e d .  118  VI.  SUMMARY AND MANAGEMENT  RECOMMENDATIONS  Summary  1.  Alectoria  436  t o 7036 g.  2.  The  (s.l.)  biomass on t e n sampled  trees  ranged  from  regression y = 158.03{A x C L ) ,  where y i s l i c h e n cover had  on a p o r t i o n  an r  trees that  3.  4.  of  frcm  Optical  with  of A l e c t o r i a  filter  crown, and CL i s crown l e n g t h , t o estimate  lichen  b i o m a s s on  (s.l.)  b i o m a s s on t h e 14 s i t e s  studied  21-1528 k g / h a . d e n s i t y v a l u e s and r e d r g r e e n  high l i c h e n  trees  of the tree  (s.l.)  were n o t s a m p l e d .  Estimates  ranged  b i o m a s s , A i s an e s t i m a t e o f A l e c t o r i a  o f 0.75 and was used  2  (8)  with  ratios  biomass d i d n o t d i f f e r  low l i c h e n  filter  ratios  s i g n i f i c a n t l y from  biomass, but d i f f e r e n c e s  f o r western  for trees  hemlock a p p r o a c h e d  those  i n red:green  statistical  significance.  5.  Photo i n t e r p r e t e r s *  rankings  of trees according t o lichen  abundance were s i g n i f i c a n t l y c o r r e l a t e d lichen  6.  with v i s u a l  estimates of  abundance i n 3 9 % o f c a s e s .  Previous experience  with  photo i n t e r p r e t a t i o n  and knowledge  119  of  lichen  judging  7.  lichen  positively  Of t h e f o u r  A regression  accounted  10.  in  potential  radiation  was  elevation, Alectoria  12.  (s.l.)  ( s . l . ) abundance  on e l e v a t i o n of the  variation  was  variation appeared  community.  and a s p e c t  accounted  were n o t s i g n i f i c a n t l y with  f o r 82% o f t h e v a r i a t i o n i n  abundance.  method  as measured  were  by v i s u a l e s t i m a t e and by t h e  not s i g n i f i c a n t l y r e l a t e d  abundance, but densiometer  accounted  f o r Hl%  abundance.  ( s . l . ) abundance, but i n c o m b i n a t i o n  these f a c t o r s  Crown c l o s u r e  moosehorn  potential  s i g n i f i c a n t and  abundance. Much o f t h e r e s i d u a l  to Alectoria  appeared t o  production.  in Alectoria  s i g n i f i c a n t and a c c o u n t e d  Taken s e p a r a t e l y , s l o p e  related  Community  statistically  of A l e c t o r i a  e x p l a i n a b l e by v e g e t a t i o n  11.  among s i t e s  ( s . l . ) abundance on  of the v a r i a t i o n  A regression  Alectoria  for lichen  of a l e c t o r i a  f o r 31%  statistically  success i n  abundance.  Community and t h e T r a n s i t i o n  annual s o l a r  1  v e g e t a t i o n communities d e s c r i b e d , the Douglas-  have t h e g r e a t e s t  9.  interpreters  were u n a b l e t o d i s t i n g u i s h  h i g h and low l i c h e n  fir/Salal  affected  abundance.  Photo i n t e r p r e t e r s  with  8.  ecology  f o r UQ%  of the v a r i a t i o n  measurements in Alectoria  by  Alectoria  o f crown  closure  abundance.  120  13.  Basal area  was not s i g n i f i c a n t l y r e l a t e d to a l e c t o r i a -  ( s . l . ) abundance.  14.  The r a t i o o f crown l e n g t h to t r e e height appeared to be  related to alectoria  ( s . l . ) abundance only on D o u g l a s - f i r / S a l a l  s i t e s and on T r a n s i t i o n s i t e s .  15.  Forest p r o d u c t i v i t y , as measured by the height of the  codeainant t r e e l a y e r , accounted f o r 64% of the v a r i a b i l i t y i n alectoria  16.  (s.l.)  abundance.  Q u a n t i t i e s of a l e c t o r i a  (s.l.) l i t t e r f a l l  on the three  study s i t e s ranged from 31.9 to 151.2 kg/ha/180 d i f f e r e n c e s among s i t e s were s t a t i s t i c a l l y  1-7.  Alectoria (s.l.) l i t t e r f a l l  days;  significant.  represented  10.5-16.1 % of  A l e c t o r i a biomass i n the t r e e canopy.  18.  A l e c t o r i a ( s . l . ) d e p o s i t i o n r a t e s i n c r e a s e d from e a r l y  winter  to l a t e winter  on a l l three  p l o t s , but comparison:with  other s t u d i e s suggested that t h i s p a t t e r n may not be g e n e r a l .  19.  No r e l a t i o n s h i p c o u l d be e s t a b l i s h e d between  r a t e s and p r e c i p i t a t o n o r l i t t e r f a l l by t o t a l i z i n g  20. and  litterfall  r a t e s and wind, as measured  anemometers.  D i f f e r e n c e s between A l e c t o r i a ( s . l . ) accumulation i n s i d e outside exclosures  were s t a t i s t i c a l l y  s i g n i f i c a n t on a l l  1.21  three study  sites,  indicating  consumption  of these l i c h e n s  by  herbivores.  21.  D i f f e r e n c e s between  inside  and  indicating  h e r b i v o r e s was  Based on  exclosures, a severe range  23.  and  24.  Alectoria  utilization  winter range  Deer use  16.7  consumption of these l i c h e n s  (s.l.) was  guantities inside  estimated  a r e a , 80.0  Areas s e l e c t e d  a t 25.7  by  and  by d e e r  as  outside  kg/ha/180 days i n  was  i n the  measured  g r e a t e r i n the  poor  winter range  winter range  tended  to  area.  by severe area.  be  h i g h i n l i c h e n a b u n d a n c e , a l t h o u g h t h e r e was  correspondence.  winter  w i n t e r range  1976-1977, a s  track counts,  areas than  and  kg/ha/180 d a y s i n a m i l d  kg/ha/180 days i n a poor  mild winter range  to-one  that  d u r i n g the w i n t e r o f  group c o u n t s  moderate o r  lichens  minimal.  a r e a , and  pellet  of non-Alectorioid  o u t s i d e e x c l o s u r e s were n o t s t a t i s t i c a l l y  significant,  22.  accumulation  no  one-  122  Becommendations  Besults Alectoria sinter that  of t h i s  study support  (s.1.)litterfall  diet  of b l a c k t a i l e d  i s an i m p o r t a n t deer,  winter  c o n s i d e r e d when mature t i m b e r  component o f  deer,  lichen  areas  described  i n Chapter  IV  abundance and  site  abundance  Preliminary steep  stratification  of s i t e s  slopes, r e l a t i v e l y  as  c o u l d be a c c o m p l i s h e d  or  photographs.  Methods f o r r a p i d  out i n t h i s  field  of l i c h e n  adapted.  Estimates of percent cover of A l e c t o r i a  sites,  i n Chapter  would be  methods d e s c r i b e d h e r e  similar  to obtain indices  could  w i t h i n a study b i o m a s s . The  a r e a , but  of  i n the  a  lichen Alectoria stems/ha  would p e r m i t c o m p a r i s o n  c o u l d n o t be r e l a t e d  r e g r e s s i o n used  be  ( s . l . ) on  t o t h e measfure " p e r c e n t  Such i n d i c e s  as  for inventory  IV. Measurement o f crown l e n g t h s and  unnecessary.  20-  (s.l.)  biomass,  b u t some o f t h e  used  moderate  forest  assessment o f A l e c t o r i a  study, i s t o o time-consuming  sample o f t r e e s c o u l d be  such  w i t h t o p o g r a p h i c maps and  purposes  abundance on  be  1965).  (Krajina  with south a s p e c t ,  abundance a r e n e e d e d . Q u a n t i f i c a t i o n carried  must  characteristics  h i g h e l e v a t i o n s , and,low  productivity 40-chain  they  provide g u i d e l i n e s f o r i d e n t i f y i n g  w i t h i n t h e C o a s t a l W e s t e r n Hemlock Zone  lichen  areas  where l i c h e n abundance i s h i g h .  B e l a t i o n s h i p s between l i c h e n  sites  the  stands are set aside  managers a r e t o c o n s i d e r a r b o r e a l l i c h e n s ,  able to i d e n t i f y  used  that  range.  If  areas  findings  even i n a m i l d w i n t e r . I n  a r e t o be managed f o r b l a c k t a i l e d  s h o u l d be  to  earlier  to  of  actual  present study  to  "  123  predict  l i c h e n b i o m a s s on  extrapolated  to other  unsampled  trees should  areas without  not  testing.  S e v e r a l recommendations f o r f u r t h e r r e s e a r c h emerge f r o m  the  guantifying  lichen  particularly usefulness could  be  present  abundance d i s c u s s e d  of o b l i q u e a e r i a l  methods  methods c f  i n Chapter I I I -  method - r e g u i r e t e s t i n g .  photographs f o r l i c h e n  of several aspects  l i c h e n s would  managers. S t u d i e s o f implications managed a s  for size  winter  provide  dispersal and  range.  of the  inventory  Wildlife  slow-growing t r e e s provide  of  useful to  d i s t a n c e s o f l i c h e n s would  spacing  availability  biology  information  of c l e a r c u t s i n areas managers need  growth r a t e s o f l i c h e n s i n d e v e l o p i n g  changes i n f o r a g e  have  to  be  information  f o r e s t s to p r e d i c t  during succession.  The  hypothesis  better substrates for lichens t  than  rapidly-growing  research  The  assessed.  Alectorioid  that  on  Ideas f o r improving  a p p l i c a t i o n o f t h e 3P  Clarification  about  study.  be  i s needed on  w e a t h e r , and  the  trees should  be  tested. Finally,  r e l a t i o n s h i p s among l i t t e r f a l l  standing crop  of l i c h e n s i n the  tree  further rates, canopy.  124  LITERATURE CITED  Abee, A. and D. Lavender. 1S72. Nutrient c y c l i n g i n t h r o u g h f a l l and l i t t e r f a l l , i n 450-year-old D o u g l a s - f i r stands. Pp. 133-143 i n J.F. F r a n k l i n , L.J. Dempster, and R.H. .flaring, eds. Research on c o n i f e r o u s f o r e s t ecosystems: f i r s t year progress i n the c o n i f e r o u s f o r e s t biome, US/IBP. A h t i , T, 1962. E c o l o g i c a l i n v e s t i g a t i o n s on l i c h e n s i n H e l l s Gray P r o v i n c i a l Park, with s p e c i a l r e f e r e n c e t o t h e i r importance to mountain c a r i b o u . Dept. of Botany, Univ. of H e l s i n k i , 69 pp. A h t i , T. and R.L. 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Von  Schrenk, H. 1898. On the mode of d i s s e m i n a t i o n of Usnea barbata,. Trans. Acad. S c i . S t . L o u i s 8: .189-198..  Bard, B.H. 1964, L i v e crown r a t i o and stand d e n s i t y i n young, even-aged red oak stands. F o r . S c i , 10(1): 56-65. Hein, R.H. and J.E. Speer. 1S75. L i c h e n biomass i n Acadian and b o r e a l f o r e s t s of Cape Breton I s l a n d , Nova S c o t i a . Bryol. 78(3): 328-333. B i l l m s , B.D. 1971. The i n f l u e n c e of f o r e s t edge, e l e v a t i o n , aspect, s i t e index, and roads on deer use o f logged and mature f o r e s t , northern Vancouver I s l a n d , B r i t i s h Columbia. M.Sc. T h e s i s , Univ. of B r i t i s h Columbia, Vancouver. 184 PP.  B i l l m s , B.D., A. McLean, and R. B i t c e y . 1976. Feeding h a b i t s of mule deer on f a l l winter and s p r i n g ranges near Kamloops, B.C. Can. J . Anim. S c i . 56 (3): 5 31-542.  131  APPENDIX Calculation  of A l e c t o r i a Totals  Symbols used  in  I (sensu l a t o )  Biomass  for Plots  appendix;  AF  - amabilis f i r (Abies amabilis)  DF  - douglas-fir  GF  - grand  MH  - mountain  WH  - western  WP  - white  WRC  - western  YC  - yellow cypress  (Pseudotsuga  menziesii)  f i r JAbies grandis) hemlock hemlock  pine  (Tsuga (Tsuga  jginus  red cedar  C  -  codominant  D  -  dominant  I  - intermediate  mertensianaV heterophylla)  monticola) (Thuja plicata).-  (Cfaamaecyparis n o o t k a t e n s i s )  Footnotes 1/ B a s e d  on  visual  2/  on  biomass  Based  e s t i m a t e s e x c e p t as  indicated.  sampling.  3/ S p e c i e s / d c m i n a n c e species/dominance  class class  not sampled;  based  on  most  similar  132  PLOT 1  Species/ Dominance Class  Lichen Tree Biomass/ Number Tree (g)  x Lichen Biomass/ C l a s s (g)  Number of Trees in Class  T o t a l Lichen Biomass/ C l a s s (kg)  WH-D  112  56 92.59  56 92.59  5  28. 46  WH-C  2 13 27 38 45 79 81  4956.33 70 36.22 4933.80 3614.26 4716.03 2950.93 4628.63  46 90.89  51  239.24  WH-I  22 31  1850.93 2666.77  2258.85  48  108. 42  WHC-C  69 70  1772. 96 2596.78  2184.87  28  61. 18  WBC-I  17 119  95.96 435.97  26 5.97  22  5.86  DF-D  84 101b  3686.52 6063.12  4874.82  6  29.25  DF-C  34 98 101a  502.9 3 3106.25 1506.85  1705.34  9  15.35  AF-I  1705.34  1  DF-I  1705.34  1  T o t a l l i c h e n biomass i n p l o t (kg)  491. 16  1/plot area (ha)  x 3. 11  T o t a l l i c h e n biomass  (kg/ha)  1528.04  133  PLOT 2  Species/ Dominance Class  Lichen Tree Biomass/ Number Tree(g)  x Lichen Biomass/ C l a s s (g)  Number of Trees i n Class  T o t a l Lichen Biomass/ C l a s s (kg)  WBC-I  5  4785.90  4785.90  3  14. 36  WH-C  13 57  620.89 511.04  56 5.97  11  6.23  HH-I  12 26 41  90. 14 43.62 82.87  72.21  26  1.88  SfiC-C  14 31 33 42 48 49 59  947.26 93.78 202.09 555.21 226.80 1034.89 191.18  464.46  22  10.22  DF-D  50 55 61  1809.21 1521.12 3233.26  2187.86  5  10.94  DF-C  54 63  570.65 1628.34  1095.50  3  3. 30  7  0.51  8 8C-C  T o t a l l i c h e n biomass i n p l o t (kg) 1/plot area  x 3.69  (ha)  T o t a l l i c h e n biomass  47. 44  (kg/ha)  175. 25  134  PLOT  Species/ Dominance Class  3  Lichen Tree Biomass/ Number Tree(g)  x Lichen Biomass/ C l a s s (g)  Number of Trees in Class  T o t a l Lichen Biomass/ C l a s s (kg)  WH-C  15  1068.6 9  106B.69  3  3.21  WH-I  36  610.16  610.16  1  .61  WBC-C  27  181.94  181.94  2  .36  WRC-I  5 17  134.97 170.84  152.91  4  .61  Df-D  18  3474. 54  3474.54  2  6.95  DF-C  2 7 8 10 13 28 32 35  2498.64 1507.6 4 2304.07 3631.95 4631. 19 2498.64 3887.96 4587.79  3193.49  16  51.10  DF-I  9  3679.66  3679.66  1  3.68  T o t a l l i c h e n biomass i n p l o t (kg) 1/plot area (ha) T o t a l l i c h e n biomass  66.52 x  (kg/ha)  10.00 665.20  135  PLOT 4  Species/ Dominance C l ass  Lichen Biomass/ Tree Number T r e e (g)  x Lichen Biomass/ Class(g)  Number of T r e e s in Class  Total Lichei Biomass/ C l a s s (kg)  HH-C  17 18 37 53 56 57 62  29 56.89 3771.89 3200.09 987.89 2655.80 3778.69 2607.16  2851.20  24  68.43  HH-I  3 6 47 63  174.73 181.94 1705. 13 593.70  663.88  12  7.9 7  WRC-C  5  459.65  459.65  3  1.38  HRC-I  49  508.09  50 8.09  3  1.52  DI-C  34 50  3737.75 1237.88  2487.82  6  14.93  (kg)  94. 23  Total  lichen  1/plot area Total  lichen  biomass  i n plot  x 10.0 0  (ha.) biomass  (kg/ha)  942.27  136  PLOT 5  Species/ Dominance Class  Lichen Tree Biomass/ Number Tree{g)  x Lichen Biomass/ Class(g)  Number of Trees in Class  T o t a l Lichen Biomass/ C l a s s (kg) 8. 16  HH-D  8 14  3495.76 4665.28  4 080.52  2  HH-C  6 9 11 18 29 30  2520.13 2825.71 3562.59 1176.86 3594.93 2052.33  2622.04  7  18.35  HH-I  15 24  7.73 1383.40  695.57  3  2.09  WBC-C  17  309.30  30 9.30  1  .31  Af-C  1 4 5  354.63 146.73 803.75  435.04  3  1.31  Af-I  20  65.08  65.08  1  .07  T o t a l l i c h e n biomass i n p l o t (kg)  x 10.0 0  1/plot area (ha) T o t a l l i c h e n biomass  30.29  (kg/ha)  302.90  137  PLOT 6  Species/ Dominance Glass  Lichen Tree Bioniass/ Number Tree(g)  x Lichen Biomass/ C l a s s <g)  Number of Trees i n Class  T o t a l Lichen Biomass/ C l a s s (kg)  WH-D  30  6005.92  6005.92  2  12.01  WH-C  56 92 108 133  3233. 23 2236.49 902.11 24 91.86  2215.92  41  90.85  WH-I  5 45 83 127  58. 43 1071.46 1812.34 3677.34  1654.89  30  49.65  WBC-C  58  825.37  825.37  10  8.25  WBC-I  104  1049.47  1049.47  12  12.59  DF-D  54  3349.41  3349.41  4  13.40  DF-C  15  1380.61  1380.61  5  6.90  DF-I  31  51.78  51.78  2  0.10  MH-C  113  1230.67  1230.67  1  1.23  T o t a l l i c h e n biomass i n p l o t (kg)  194.98  1/plot area (ha)  x 5.00  T o t a l l i c h e n biomass  (kg/ha)  974.90  138  PLOT 7  Species/ Dominance Class  Lichen Tree Biomass/ Number Tree(g)  x Lichen Biomass/ C l a s s (g)  Number of Trees i n Class  T o t a l Lichen Biomass/ C l a s s (kg)  WH-D  27  1298.13  129 8.13  1  1.30  WH-C  10 1ft 22 52 61  1275.99 1419.66 2 109.20 2667.64 1393.39  1773.18  11  19.50  WH-I  19 26 57 58  40. 68 165.66 73.31 11.69  72.84  11  0.80  WBC-D  39  1 12.01  1  0.11  WBC-C  33  99.03  4  0.40  WBC-I  50  13. 68  1  0.01  DF-D  51  1776.65  3  5.33  DF-C  30  236.07  2  0.47  Total 1/plot Total  l i c h e n biomass i n p l o t (kg) area  (ha)  l i c h e n biomass (kg/ha)  27.92 x 5.00 139.60  139  PLOT 8  Species/ Dominance Class  Lichen Tree Biomass/ Number Tree eg)  x Lichen Biomass/ C l a s s (g)  Number of Trees i n Class  T o t a l Lichen Biomass/ C l a s s (kg)  WH-D  82  50 58.76  505 6.76  2  10.12  WH-C  28 43  1983.05 1917.81  1450.43  8  11 .60  BH-I  49  1491.03  1491.03  3  4. 47  AF-C  3 87  919. 08 17,53  468.31  7  3.28  AF-I  13 58 74  24.24 48. 81 18.92  30.66  10  0.31  MH-C  6 50  3909.76 1072.38  2491.07  6  14.95  YC-D  51  5200.49  520 0.49  1  5.20  YC-C  39 84  1086.08 1468.49  1277.29  6  7.66  YC-I  54  152.76  152.76  1  0. 15  T o t a l l i c h e n biomass i n p l o t  (kg)  57.74  1/plot area (ha)  x 5.00  T o t a l l i c h e n biomass (kg/ha)  288.71  140  PLOT 9  Species/ Dominance Class  Lichen Tree Biomass/ Number Tree(g)  x Lichen Biomass/ C l a s s (g)  Number of Trees i n Class  Total Lichen Biomass/ C l a s s (kg)  WH-D  20  7238.69  7238.69  6  43.4 3  WH-C  47 52 78 98 103 106  1953.80 5503.25 3727.46 2263. 12 1232.15 3316.15  3204.68  50  160.23  WH-I  2 112  3801.73 529. 73  216 5.73  30  64. 97  WBC-C  16 92  266.36 764.74  515.55  13  6.70  WBC-I  17 41 13 29 38  304.77 154. 18 4 59.28 13 27.93 2040.51  229.48  38  8.72  1275.91  20  25.52  DF-C  DF-D  (1275.91)  3  3.83  DF-I  1275.913  2  2.55  T o t a l l i c h e n biotrass i n p l o t (kg)  315.95  1/plot  x 2.53  area (ha)  T o t a l l i c h e n biomass  (kg/ha)  800.46  141  PLOT 10  Species/ Dominance Class  Lichen Tree Biomass/ Number Tree (g)  x Lichen Biomass/ C l a s s (g)  Number of Trees i n Class  Total Lichen Biomass/ G l a s s (kg)  BH-B  35  3672.86  3672.86  4  14.69  WH-C  9 12 13 15 17 33  1880*01 3567.05 2326.51 3885.22 2534.49 1996.86  269 8. 36  17  45. 8 7  WH-I  30 43 47  871. 77 773.24 545.81  730.27  9  6.57  WBC-C  26  225.37  225.37  3  0.68  HRC-I  42  106.98  106.98  8  DF-C  1 10 20  2321.14 1858. 19 1739.49  1972.94  9  17.76  GF-C  6  146.9 9  146.99  1  0.15  2  3.95  DF-D  (1972.94)  DF-I  1972.943  Total  1  l i c h e n biomass i n p l o t (kg)  1.97  92.50 x 5.6 3  1/plot area (ha) T o t a l l i c h e n biomass  0.86  (kg/ha)  521.20  142  PLOT 11  Species/ Dominance Class  Lichen Tree Biomass/ Number Tree(g)  x Lichen Biomass/ C l a s s (g)  Number of Trees i n Class  T o t a l Lichen Biomass/ C l a s s (kg)  WH-D  9  3102.87  3102.87  2  6.21  HH-C  8 11 37 53 71  3283.93 2785.41 3112.35 2857.04 3157.43  3039.23  32  97.26  WH-I  63 77  4372.67 1934.93  3153.80  27  85. 15  WEC-C  31 54  2143.67 837.64  1490.66  9  13.42  WRC-I  60  1223.09  1223.09  13  15.90  DF-D  20  20 58. 81  2056.81  3  6.18  DF-C  36 45  3959.60 748.82  2354.21  11  25.90  DF-I  33  1627.08  1627.08  5  Total  l i c h e n biomass i n p l o t  1/plot area  (ha)  T o t a l l i c h e n biomass  (kg)  8. 14 258.16 x 5.87  (kg/ha)  1516.18  143  PLOT 12  Species/ Dominance Glass  Lichen Tree Biomass/ Number Tree(g)  x Lichen Biomass/ Class (gj  Number o f Trees i n Class  T o t a l Lichen Biomass/ C l a s s (Jig)  2033.86  2  4. 07  3,94 57.02 63. 90  41.62  9  0.0 4  17 20 74  0 8. 10 0  2.70  9  0.02  WBC-D  5  8. 47  8.47.  1  0.01  WBC-C  58 65  0 0  0  6  0  AF-C  72  5.23  3  0.02  Af-I  11 45 50 53  0 2.05 0 0  17  0.01  WH-D  1  WH-C  66 79 83  WH-I  2033.86  Total 1/plot Total  l i c h e n biomass i n p l o t (kg) area (ha) l i c h e n biomass (kg/ha)  4. 17 x 5.00  20.84  144  PLOT 13  Species/ Dominance Class  Lichen Tree Biomass/ Number Tree (g)  x Lichen Biomass/ C l a s s (g)  Number of Trees i n Class  T o t a l Lichen Biomass/ C l a s s (kg)  WH-D  30 55  3027. 43 2231.71  2629.57  4  10.52  WH-C  8 50 58  1246. 19 116.95 57. 83  473.66  7  3.32  TJH-I  32  353.81  353.81  4  1.42  WE C-C  4 10  0 23.96  11.98  5  0.06  WRC-I  6 7  23.00 74. 16  45.58  9  0.4 4  DF-D  19 42 48  560.29 485.35 884.54  64 3.39  7  4.50  DF-C  26 27  90. 97 434.87  262.92  4  1.05  T o t a l l i c h e n biomass i n p l o t (kg)  x 5.0 0  1/plot area (ha) T o t a l l i c h e n biomass  21. 30  (kg/ha)  106.55  145  PLOT 14  Species/ Dominance Class  Lichen Tree Biomass/ Number Tree (g)  x Lichen Biomass/ C l a s s (g)  Number of Trees i n Class  T o t a l Lichen Biomass/ C l a s s (kg) 22. 8 7  HH-C  9 14 33 35 56  353.08 20 94.49 233.85 6409.88 1304.62  2079.18  11  WH-I  51 59 59  12.87 143. 11 143.87  99.62  8  0. 80  REC-C  1 52  324.43 98.55  211.49  8  1.69  HBC-I  58  0  0  2  0  AF-C  8 69  799.30 1745.41  1272.36  6  7. 63  AE-I  43  9. 87  9.87  4  0. 04  YC-C  38  2163.82  1  2. 16  T o t a l l i c h e n biomass i n p l o t (kg)  x 5.00  1/plot area (ha) T o t a l l i c h e n biomass  35.20  (kg/ha)  175. 95  146  APPENDIX I I  Differentiated (Hueller-Dombois  Table  of Vegetation  and E l l e n b e r g  Data  1  1974: 177-193)  The f i r s t number i n e a c h box r e p r e s e n t s an e s t i m a t e o f p e r c e n t c o v e r . The s e c o n d number r e p r e s e n t s a d i s t r i b u t i o n c l a s s . Where o n l y one number i s p r e s e n t , i t r e p r e s e n t s p e r c e n t c o v e r . 1  Vegetation Community  TRANS  AF/OH  P l o t number Elevation (m) Aspect Slope (degrees)  14 8 15 5 603 707 695 573 N35W S05W N15W N75E 24 5 16 23  9 6 669 646 S60E N82W 32 30  Abies amabilis Covnus canadensis Rubus pedatus Vaccinium ovalifolium Menziesia fevruginea Listera cauvina Clintonia uniflora Streptopus roseus Plagiothecium: undulgtum Tsuga mevtensiana Chamaecyparis nootkatensis Sphagnum sp. Khizomnium sp. Coptis asplenifolium Gymnocarpium dryopteris  25 15 25 2/4 ' 1/5 1/4 '2/6 .1/5 1/4 10/5 5/5 15/6 3/4 3/4 1/2 1/2 1/4 1/5 1/5 1/5 1/2 5/5 1/3 7 2 2 5 1/5 2/5 1/5 2/5 2/6 1/5 1/5 2/6  10 3/6 1/5 1/2 1/2 1/1  1 2/7 1/1 1/6 25/6 1/1 1/2 1/1 1/4  2  Tsuga heterophylla Listera covdata Hylocomium splendens Rhytidiadelphus loreus Thuja plicata Vaccinium parvifolium  1 1/3 1/5 40/8 1/1  1/3 2 1  7 70/7  Pseudotsuga menziesii Gaulthevia shallon Eemitomes congestum Stokesiella ovegana Viola sempevvirens Boschniakia hookevi  13 11 792 426 S60E N87W 14 25  1/5  20 1/5 1/5 1/3  20 5/6 1/2  55 1/4 70/8 10/6 2 3/6  UNCLASSIFIED  1 10 3 469 686 543 S10W S15E S30E 20 32 54  12 4 2 7 487 709 282 442 S22W S60E N80W S40E 32 22 20 15  1 1/5  25 1/3  1 1/6  1 1/3  8 20/6  5 1/5 1/2  1/2 1/1 1/1 1/5  35 1/3 3/5 1/5  1/1 60 45 35 35 1/2 1/2 1/2 1/4 20/6 15/6 5/5 15/6 50/8 15/6 50/8 35/8 15 5 2 5/4 1/4 3/4  DF/S  40 50 60 1/4 2/7 1/2 30/6 20/8 40/8 10/5 20/8 20/6 15 5 15 10/4 5/4 3/5  35 7 55 90/8 40/8 80/8 1/2 1/1 1/1 1/1 •"•1/1 1/2 1/3 1/2 1/1 1/4 45 9 30 1/4 1/4 :i/i 9/6 30/8 50/8 1/4 20/7 5/7 .5 15 7 2/4 15/7 12/7  20 3/3 3/4  80 35 25 70 1/4 1/6 1/2 3/7 10/6 50/8 70/8 30/8 10/6 10/7 10/7 30/8 15 5 4 35 2/4 15/7 4/7 3/2  Appendix  II  Continued  Vegetation Community Vaeciniwn alaskaense Chimaphila menziesii Rhytidiopsis robusta C o r a l l o r h i z a mevtensiana Goodyera o b l o n g i f o l i a Hypopitys monotvopa Bleohnum spioant Aehlys triphylla Pinus monticola Moneses u n i f l o r a Linnaea borealis T i a r e l l a tvifoliata Laotuaa muralis Polystichum lonehitzs Berberis nervosa Polystichum. munitum Athyrium filix-femina T i a r e l l a trifoliata var. laainiata Chimaphila umbellata Dryopteris austriaca Diaranum sp. Abies grandis Rosa gymnooarpa Sambucus raeemosa AIlotropa virgata Maianthemum dilatatum Bromus oarinatus Lysiohitum americanum Epilobium angustifolium Veratrum v i r i d e Viola g l a b e l l a Plagiomnium sp. Monotropa uniflorg.  AF/OH  TRANS  50/6 75/8 40/8 25/8 1/2 1/2 1/2 2/5 30/8 20/6 35/8 1/5 1/5 1/2 1/5 1/5 1/5 1/2 1/5 1/3 1/5 5/6 1 1/2 1/4 1/1 2/4  1/6 1/2  2/5 35/7 40/8 1/2 1/2 1/2 25/6 40/6 5/6 1/5 1/5 1/5 1/2 1/2 1/2 1/2 1/2 1  DF/S 2/5 1/4 2/5 1/5 1/2  1/1 10/6 1/1 1/5 15/7 1/5 1/5 1/1  1/1 2/4 40/7 2/7 1/4 1/2 1/2 1/2 30/7 2/5 1/2 1/21/21/2 1/21/11/41/2 1/2 1/2 1/4 1/3 1/2 1/3 1/5 1 1 1 1/3 1/2 1/4 1/3  1/2 1/3  1/3 1  1/51/51/4 1/5 1/11/2 1/1 1/5 1/3 1/1 2/5 1/1 1/3  1/2 5/7 3/2 1/2 1/2 3/2  3/3 3/2 1/5 1/2  1/6 1/1  UNCLASSIFIED  1/3 1/3  1/6 1/1 1/5  1/5  1 1/5 1/1 1/3 1/1 1/5 1/5 1/1 1/5 1/5 1/5 1/5  

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