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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 SUSAN KINGSBURY STEVENSON B.A., Swarthmore College, 1969 A THESIS SUBMITTED IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF THE FACULTY OF GRADUATE STUDIES (DEPARTMENT OF FORESTRY) We accept th is thesis as conforming to the required standard UNIVERSITY OF BRITISH COLUMBIA May, 1978 ~c) Susan K. Stevenson, 1978 by MASTER OF SCIENCE In presenting th i s thes is in pa r t i a l fu l f i lment of the requirements for an advanced degree at the Univers i ty of B r i t i s h Columbia, I agree that the L ibrary sha l l make it f ree ly ava i l ab le for reference and study. I fur ther agree that permission for extensive copying of th is thesis for scho lar ly purposes may be granted by the Head of my Department or by his representat ives. It is understood that copying or pub l i ca t ion of th is thesis fo r f inanc ia l gain sha l l not be allowed without my writ ten permission. Department of The Univers i ty of B r i t i s h Columbia 2075 Wesbrook Place Vancouver, Canada V6T 1W5 Date TJA^JL // /f7t? ABSTRACT Biomass of a r b o r e a l l i c h e n s used as winter food by b l a c k t a i l e d deer was s t u d i e d . Three methods were used to assess abundance of A l e c t o r i a sarraentosa and B r y o r i a spp. Lichen biomass was measured by sampling the l i c h e n s from f e l l e d t r e e s . A system of v i s u a l estimates of l i c h e n abundance was developed and r e l a t e d to biomass estimates obtained by sampling. A p r e d i c t i v e eguation {Y = 158.03 {A x CL), where Y i s l i c h e n biomass, A i s an estimate o f l i c h e n cover on a p o r t i o n of the t r e e crown, and CL i s crown l e n g t h ; n = 40; S y, x = 376,89; r 2 = 0,75) was used t o e x t r a p o l a t e l i c h e n biomass values from sampled t r e e s to unsampled t r e e s . The value of l a r g e - s c a l e c o l o u r i n f r a r e d a i r photography as a t o o l f o r i n v e n t o r y i n g l i c h e n abundance was assessed, using densitometry and photo i n t e r p r e t a t i o n . Some p o s i t i v e r e l a t i o n s h i p s with l i c h e n abundance were found using each method, but n e i t h e r densitometry nor photo i n t e r p r e t a t i o n was demonstrated t o have s t r o n g p o t e n t i a l f o r use i n i n v e n t o r y i n g abundance of a r b o r e a l l i c h e n s . Biomass of A l e c t o r i a (sensu l a t o ) on the 14 p l o t s s t u d i e d ranged from 21 to 1528 kg/ha. P h y s i c a l and v e g e t a t i v e c h a r a c t e r i s t i c s of the p l o t s were measured and r e l a t e d to l i c h e n abundance. Taken to g e t h e r , s l o p e , aspect, and e l e v a t i o n accounted f o r 82 percent of the v a r i a t i o n i n l i c h e n abundance. F o r e s t p r o d u c t i v i t y was n e g a t i v e l y r e l a t e d to l i c h e n abundance, l i t h i n t h e range of s i t e s s t u d i e d , A l e c t o r i a (s.1.) was most abundant on moderate to steep s o u t h - f a c i n g s l o p e s , at e l e v a t i o n s above 500 m, where tree growth was poor. i i i To assess a v a i l a b i l i t y of l i c h e n s and t h e i r u t i l i z a t i o n by deer, l i t t e r f a l l was measured i n s i d e and o u t s i d e e x c l o s u r e s on t h r e e s i t e s where l e v e l s of deer use i n winter were known. The r e l a t i o n s h i p s between l i t t e r d e p o s i t i o n r a t e s and weather were examined. Q u a n t i t i e s of A l e c t o r i a (s.1.) l i t t e r were 69.9 kg/ha/180 days i n a severe winter range area, 151.2 kg/ha/180 days i n a m i l d winter range area, and 31.9 kg/ha/180 days i n a poor winter range area. On a l l t h r e e s i t e s , s i g n i f i c a n t l y more (p<0.05) A l e c t o r i a l i t t e r was present i n s i d e than o u t s i d e e x c l o s u r e s ; g u a n t i t i e s of n o n - l i c h e n l i t t e r d i d not d i f f e r s i g n i f i c a n t l y . U t i l i z a t i o n of A l e c t o r i a was 37, 53, and 52 percent of a v a i l a b l e g u a n t i t i e s on the three s i t e s , r e s p e c t i v e l y . The r e l a t i o n s h i p between l i c h e n abundance and s e l e c t i o n of winter h a b i t a t by b l a c k t a i l e d deer was assessed, based on p e l l e t group counts, t r a c k counts, and the data of other i n v e s t i g a t o r s . Areas s e l e c t e d by deer as winter range tended to be moderate or high i n l i c h e n abundance. i v TABLE OF CONTENTS ABSTRACT i i L I S T OF TABLES .............................................. v i L I S T OF FIGURES . . . . v i i L I S T OF APPENDICES .......................................... i x ACKNOWLEDGEMENTS X I . INTRODUCTION 1 R a t i o n a l e ........................................... 1 O b j e c t i v e s 3 The S t u d y A r e a ...................................... 4 I I . EACKGBCUND 7 Dse Of A r b o r e a l L i c h e n s By U n g u l a t e s 7 N u t r i t i o n a l V a l u e Of A r b o r e a l F o r a g e L i c h e n s ........ 14 E c o l o g y Of A l e c t o r i o i d L i c h e n s ...................... 16 I I I . , QUANTIFICATION OF BIOMASS OF AEBOHEAL FORAGE LICHENS .. 22 Methods . ... 26 V i s u a l E s t i m a t e s Of L i c h e n Abundance 26 A i r P h o t o Q u a n t i f i c a t i o n Of L i c h e n Abundance ..... 30 B i c a a s s S a m p l i n g 36 C o m p u t a t i o n s And R e s u l t s ............................ 39 C o m p u t i n g B i o m a s s T o t a l s ......................... 39 R e s u l t s Of A i r P h o t o Q u a n t i f i c a t i o n 45 D i s c u s s i o n . ............. .......... .. ................ 50 E v a l u a t i o n Of Methods ............................ 50 B i o m a s s R e s u l t s Compared W i t h L i t e r a t u r e R e p o r t s . 55 I V . , RELATIONSHIPS BETWEEN LICHEN BIOMASS AND S I T E V CHAfiACTERISTICS . . ..... . . ..... . 58 Methods 60 Besults . . 63 Vegetation Communities ........................... 63 Measures Of Physical Environment ................. 68 Forest Measures 75 Discussion 81 V. AVAILABILITY OF ASBOREAL LICHENS AND UTILIZATION BY DEES 85 Methods ........................... ,. .. 87 L i t t e r f a l l Measurement ........................... 87 Habitat Selection ................................ 90 Besults ............................................. 92 Total L i t t e r f a l l Quantities ...................... 92 Lichen L i t t e r f a l l In Relation To Lichen Biomass ,. 94 L i t t e r Deposition In Relation 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 Plots ..................... 106 Lichen Abundance And Deer Use .................... 107 Discussion .......................... ... ............. 110 L i t t e r f a l l ..................................... .. 110 Deer Use And lichen Abundance 116 VI. , SUMMABY AND MANAGEMENT RECOMMENDATIONS ................. 118 Summary ...... ........ . .... 118 Eecommendations ..................... ................ 122 LITEEAIUEE CITED 124 v i LIST OF TABLES I. N u t r i t i o n a l value of arboreal forage lichens ....... 1 5 I I . Biomass of Ale c t o r i a on sampled trees .............. 4 1 I I I . Optical density values and red:green f i l t e r r a t i o s of trees with high and low lichen biomass .......... 4 6 IV. Spearman's rho values for predictions of lichen abundance on i n d i v i d u a l trees . . . . . . . . . . . . . . . . . . . . . . , 4 8 V. 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. L i t t e r f a l l quantities inside exclosures ............ 9 3 VIII. Alectoria l i t t e r f a l l in r e l a t i o n to Al e c t o r i a standing crop ..... 9 5 IX. L i t t e r f a l l quantities i n s i d e and outside e x c l o s u r e s . 1 0 4 X. Deer use and snow depths at Plots 3, 4, and 5, March 28-29, 1977 ........................................ 1 0 8 XI. Habitat selection and lichen biomass ............... 1 0 9 XII. Comparison of reported quantities of lic h e n l i t t e r f a l l ......................................... I l l « v i i LIST OF FIGURES 1. Location of the study area (Jones 1975) 5 2. , Measurement of crown width p a r a l l e l and perpendicular to is ix© sXcp^ • # • * • • * • * * • . « '• *(•••••«.,•.'•'*••• • '• •' • • *'-••'•.•••• 29 3. Division of a tree into layers for visu a l estimates of 1 xc i i ©n cii^un d c^ n c € • • • • • • • «* • • *: * • %-•>• • • • • • • * • ... 29 '*» . Mfictoria saraentosa on Douglas-fir; normal colour i f i l m (A) and colour infrared film with Jfratten 12 f i l t e r and two CC20 magenta f i l t e r s (B) 31 5. Colour i n f r a r e d a i r photographs of plot with low 1 xch-€ E JDXGIUCL ss * • *"• • * • • • • • * • : • • "• *./#:» *"*,"• • *••#>•»" #• • • •  • 34 6. Colour infrared a i r photographs of plot with high 1 i c li s n 13 x o sts ss »• * • « • • * • * * * * * * • • • • • •. • • • • •« * • • • * . 35 7. Regression used to calculate lichen biomass of visu a l 8. Lichen biomass per tree based on independent v i s u a l estimates by two observers ...:....... *..... >....... 44 9. Vegetation community and Alectoria biomass (A): vegetation community and percent Alectoria IB) . . . . . . . 69 10., Relationship between slope and percent A l e c t o r i a ..... 70 11. Relationship between aspect and percent Alectoria .... 71 12. Linear regression of percent Alectoria on poten t i a l annual radiation . . . . . . . . . . . . . . . . . . v .... . . . v i . . . . . . . . . . 72 13. Linear regression of percent Alectoria on elevation .. 73 14. Relationship between crown closure and percent Alectoria i densiometer (A); moosehorn (B); visua l estimates (C) ......... .... ........ . . * •. v * . . . « * . . . . . 76 v i i i 15. R e l a t i o n s h i p between b a s a l area and percent A l e c t o r l a . 78 16. 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 h e i g h t of codcffiinant t r e e l a y e r ................................ 79 17. R e l a t i o n s h i p between crown l e n g t h / t r e e height and pcxcsnt AC T~tor 13^ * • • • • • • • •» * » • • • • •"» • •*-#-• • • - ' 80 18. Layout of l i t t e r f a l l p l o t s 8 9 19. 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 l i t t e r i n e a r l y winter (E), mid-winter (ft), and l a t e winter ( L ) , 1S76-20. D e p o s i t i o n r a t e s of other {non-Alectorioid) l i c h e n l i t t e r i n e a r l y winter ( E ) , mid-winter (H), and l a t e winter (L) , 1976-1977 97 21. D e p o s i t i o n r a t e s o f a l l l i t t e r i n e a r l y winter ( E ) , mid-winter ( f 4 ) , and l a t e winter (L), 1976-1977. ...... 9 8 22. P r e c i p i t a t i o n at Port Hardy and Hoss Camp i n e a r l y winter (E) , mid-winter (M) , and l a t e winter (L), 1 S1 € *" 1 S *77 • • * « # • • * * • * • * • • ••«:*•:••.•*••••*••*• • • • • • • • • • m- * • 100 23. R e l a t i v e amounts of wind i n mature timber (A) and c l e a r o u t s (B) i n e a r l y winter (E), mid-winter (H) , and l a t e winter (L) , 1976-1977 . . . , V . , . . , . . . . . . . . 1 0 1 24. A l e c t o r i a u t i l i z a t i o n on P l o t s 3, 4, and 5, with 95% cc n £ icls nc€ 1 x 01 i t s • • * • • • • • « « • • * • * * • * • • • * • •» •• • • *. * # • • * * * • 105 25. D e p o s i t i o n r a t e s of forage l i t t e r ( l i c h e n s and green c o n i f e r f o l i a g e ) i n the study area, 1973-1974 (Rochelle 1 9 7 8 ) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 114 26. D e p o s i t i o n r a t e s o f three l i t t e r f a l l components a t Northwest Bay, 1975-1S76 (Kale unpublished d a t a ) . . . . . 115 i x LIST OF APPENDICES I. , C a l c u l a t i o n of A l e c t o r i a ( s . l . ) biomass t o t a l s f o r p l o t s 131 I I . D i f f e r e n t i a t e d t a b l e o f v e g e t a t i o n data 146 X ACKNOWLEDGEMENTS I r e c e i v e d p e r s o n a l f i n a n c i a l a s s i s t a n c e f o r graduate study from the Canadian W i l d l i f e S e r v i c e , Canadian F o r e s t Products L t d . , and the U n i v e r s i t y of B r i t i s h Columbia. T h i s study was funded by the B r i t i s h Columbia F i s h and W i l d l i f e Branch through a grant to Dr. F.L. B u n n e l l . Dr. D.S. Eastman, i / c W i l d l i f e Research, handled the funding arrangements; without h i s a s s i s t a n c e and encouragement, the study would not have been completed. Dr. D. Hebert, R e g i o n a l W i l d l i f e B i o l o g i s t , provided a d d i t i o n a l support. The study was conducted with the c o o p e r a t i o n and a s s i s t a n c e of Canadian F o r e s t Products, L t d . Many i n d i v i d u a l s w i t h i n the company aided me; I would l i k e to acknowledge i n p a r t i c u l a r the a s s i s t a n c e of Mr. Stan Chester, Mr. J u l i u s Kapitany, Mr. Greg Jones, and the s t a f f of the f i r e shop. Mr. Gordon Flowerdew f e l l e d the t r e e s f o r biomass sampling, and Mr. Henry Shear allowed me to use h i s cabin. Mr. And Mrs. T. C a r l y l e helped with the f i e l d w o r k i n many ways. The members of my Graduate Committee, Dr..H.B. S c h o f i e l d and Dr. P.A. Murtha provided d i r e c t i o n and c o n s t r u c t i v e c r i t i c i s m . I am e s p e c i a l l y g r a t e f u l to my s u p e r v i s o r , Dr. F.L. Bunnell, f o r h i s guidance, encouragement, and p a t i e n c e . Mr. Parker W i l l i a m s , of I n t e g r a t e d Resources Photography, Ltd. provided e x c e l l e n t a i r photographs. I wish to thank a l l those who worked long hours a s s i s t i n g me i n the f i e l d and with the p r e p a r a t i o n of the manuscript. The support of my husband, David, encouraged me through a l l staqes the study. 1 I. INTRODUCTION Rationale The use of clearcuts and young second growth as winter range by blacktailed deer 10docoileus hemionus cclumbianus Richardson) has been documented f o r many areas (Brown 1961, Gates 1968, M i l l e r 1968), but where snowfall i s high, mature timber appears important to deer in winter. Edwards (1956), for example, found that ungulate numbers in B r i t i s h Columbia declined during years of deep snow, but that adeguate cover could offset the e f f e c t s of snow. Cowan (19 56) recognized the importance of stands of coniferous trees in providing shelter from heavy snowfalls to bl a c k t a i l e d deer i n the northern part of i t s range. Gates (1968) noted that early serai stages of vegetation are of li m i t e d value to deer when food a v a i l a b i l i t y i s r e s t r i c t e d by deep snow. Jones (1975) reported that on northern Vancouver Island deer, i n winter, used mature timber habitats more in t e n s i v e l y than logged habitats, especially when snow was deep and soft. However, not a l l forest types were found to be of egual value, and further research on winter habitat was recommended. Much of the area in B r i t i s h Columbia that i s occupied by blacktailed deer i s prime timber-producing land which i s subject to extensive logging. It may be essential to the maintenance of blacktailed deer populations that appropriate areas of mature timber be i d e n t i f i e d and set aside as habitat f o r wintering 2 animals. T h i s s i t u a t i o n has c r e a t e d an urgent need f o r research i n t o the c h a r a c t e r i s t i c s of b l a c k t a i l e d deer winter range. Because food i s o f t e n c r i t i c a l to deer i n w i n t e r , and forage i s l i m i t e d under the canopy of a mature f o r e s t , winter range i n v e s t i g a t i o n s must c o n s i d e r forage a v a i l a b i l i t y . In the northern part of the range of b l a c k t a i l e d deer a r b o r e a l l i c h e n s are a major component of the winter d i e t (Cowan 1945; Gates 1968; Jones 1975; R o c h e l l e 1978). The l i c h e n s which are most h e a v i l y used as forage belong t o the genera Dsnea^ A l e c t o r i a ^ and B r y p r i a . They are l o n g , pendulous l i c h e n s o f t e n r e f e r r e d to as "beard l i c h e n " or " o l d man's beard". They grow on mature t r e e s and are most commonly a v a i l a b l e t o deer as l i t t e r f a l l on the f o r e s t f l o o r . The purpose of the present study was to i n v e s t i g a t e the p r o duction of a r b o r e a l forage l i c h e n s and t h e i r u t i l i z a t i o n by b l a c k t a i l e d deer. 3 O b j e c t i v e s Four o b j e c t i v e s were d e f i n e d f o r the p r o j e c t : 1. To develop and e v a l u a t e methods of q u a n t i f y i n g biomass of a r b o r e a l forage l i c h e n s . 2. To r e l a t e a r b o r e a l forage type, common f o r e s t i n v e n t o r y c h a r a c t e r i s t i c s . l i c h e n s to v e g e t a t i o n community measures, and other s i t e 3. To assess winter a v a i l a b i l i t y and u t i l i z a t i o n of forage l i c h e n s . 4. To r e l a t e abundance of a r b o r e a l forage l i c h e n s to s e l e c t i o n of winter 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 located in the Nimpkish Valley on north-central Vancouver Island (Fig. 1). The si t e of most of the fieldwork was the southeastern portion of the Nimpkish drainage, near the confluence of Croman Creek and the Davie River. Geologic history, physiography, and climate of the study area were described by Willms (1971). The area i s mountainous, with U-shaped g l a c i a l valleys at about 300 m a . s . l . r i s i n g to peaks of 1500 to 1800 m. The climate i s characterized by moderate temperatures but high precipitaton in winter. Snowfall i n the study area i s generally greater than the 10 2 cm mean annual t o t a l recorded at Boss Camp, located 14 km northwest of the study area at an elevation of 140 m a . s . l . (Hillms 1971). Vegetation i n the study area f a l l s within the Coastal Western Hemlock Zone, the Mountain Hemlock Zone, and the Alpine Tundra Zone (Krajina 1965). Most of the fieldwork was carried out within the Coastal Western Hemlock Zone. Clearcut logging began in the study area i n 1947 (Willms 1971) and, at present, the area i s a mosaic of regenerating stands aged 30 years or l e s s , and mature stands aged 300 years or more. A l l study plots were located in mature stands. Although many species of arboreal lichen are present i n the study area, the species most used as forage belong to the closely related genera A l e c t o r i a and Bryoria. Alggtoria sarmentosa (Ach.) Ach. i s the most abundant species of thi s group i n the study area. Bryoria glabra (Mot.) Brodo and D. Hawksw. i s the most abundant of the Brioriae.. Other species Figure .1. Location of the study area (Jones 1975). 6 o c c a s i o n a l l y found i n the study area are B.. f r i a b i l i s Brodo and D. Hawksw., B± oregana (Tuck.) Brodo and D. Hawksw., B. c a p i l l a r i s (Ach.) Brodo and D. Hawksw., B. t r i c h o d e s (Michx.) Brodo and D. Hawksw. subsp. t r i c h o d e s . and B. t r i c h o d e s subsp. americana (Hot.) Brodo and D. Hawksw. The genus Usnea, r e p o r t e d from other l o c a l i t i e s as important f o r forage, i s extremely r a r e i n the study area (only one known specimen). I t occurs more commonly at the north end of the Nimpkish V a l l e y where the c l i m a t e i s more oceanic. The f i e l d w o r k was c a r r i e d out during the p e r i o d May 1976 to September 1977. 7 I I . BACKGROUND Use of A r b o r e a l Lichens by Unsulates A number of c e r v i d s p e c i e s use a r b o r e a l l i c h e n s as forage. In g e n e r a l , consumption of a r b o r e a l l i c h e n s seems to be a s s o c i a t e d with high s n o w f a l l . Sources of a r b o r e a l l i c h e n f o r a g e i n c l u d e the lower branches of t r e e s , l i t t e r f a l l , n a t u r a l l y f a l l e n t r e e s , and t r e e s f e l l e d by l o g g e r s . B l a c k t a i l e d deer commonly use l i c h e n s a v a i l a b l e as l i t t e r f a l l . On southern Vancouver I s l a n d , Cowan (1945) found that an a r b o r e a l l i c h e n which he i d e n t i f i e d as Dsnea barbata •1 c o n s t i t u t e d 36% by volume of winter rumen contents of b l a c k t a i l e d deer. Lichens were consumed from broken branches brought down by snow and high winds, and from r e c e n t l y f e l l e d t r e e s . On the southeastern c o a s t of Vancouver I s l a n d Gates (1968) observed that a r b o r e a l l i c h e n s , made a v a i l a b l e by strong winds and by l o g g i n g , c o n s t i t u t e d 13%, by volume, of winter rumen samples and were the t h i r d most important winter forage item. Jones (unpublished d a t a ) 2 found 10% and 4% a r b o r e a l l i c h e n i n rumen samples of b l a c k t a i l e d deer c o l l e c t e d d u r i n g severe and 1 I v i s i t e d Cowan's study area near Goldstream Lake i n June, 1976 and noted t h a t A l e c t o r i a and B r ^ o r i a spp. were about as 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 consuming Dsnea.x i t seems l i k e l y t h a t a l l pendulous f r u t i c o s e l i c h e n s i n the rumen samples were i d e n t i f i e d as Usnea barbata. 2 Canadian F o r e s t Products L t d . , Woss, B r i t i s h Columbia. 8 mild w i n t e r s , r e s p e c t i v e l y , i n the Nimpkish V a l l e y on northern Vancouver I s l a n d . H i s f i e l d o b s e r v a t i o n s suggested t h a t l i c h e n s were more h e a v i l y used than these data i n d i c a t e . L i c h e n g u a n t i t i e s i n rumen samples may have been low because most rumens were c o l l e c t e d from deer i n logged areas (Jones 1975). In the same study area, B o c h e l l e (1978) found that A l e c t o r i o i d l i c h e n s were a major d i e t a r y item f o r deer c o l l e c t e d i n timbered stands during winter. They occurred at a volume of 35.5%, with 100% freguency. The l i c h e n s were second o n l y to s a l a l i f i a u l t h e r i a s h a l l o n i , which had a volume of 43.6%; a l l other s p e c i e s occurred i n volumes o f l e s s than 5%. A r b o r e a l l i c h e n s are eaten by other North American deer as w e l l , but they may not be a c r i t i c a l component o f the winter d i e t . Schroeder (1974) observed both mule deer (Odocqileus hemionus hemionus) and white t a i l e d deer (Odocoileus v i r g i n ^ anus) feed i n g on A l e c t o r i a sarmentosa from f a l l e n t r e e s i n the S e l k i r k Mountains of southeastern B r i t i s h Columbia. In South Dakota, Usnea spp. were of secondary importance as winter food items f o r mule deer and w h i t e t a i l e d deer; more l i c h e n s were eaten as snow depth i n c r e a s e d (Schneewis et a l . 1972). S c o t t e r (1964) r e p o r t e d t h a t d u r i n g p e r i o d s of deep snow, t r a p p e r s i n w e s t - c e n t r a l A l b e r t a cut down t r e e s so t h a t mule deer c o u l d f e e d on the a r b o r e a l l i c h e n s . Data presented by Willms e t a l . (1976) i n d i c a t e that consumption o f l i c h e n s by mule deer near Kamloops decreased from 6.9% of rumen contents i n e a r l y f a l l to 0.6% i n midwinter. T h i s change probably occurred because a r b o r e a l l i c h e n s were l e s s abundant on the animals* winter ranges than on f a l l ranges. In an oak woodland h a b i t a t i n northwestern 9 C a l i f o r n i a , the a r b o r e a l l i c h e n s Bamalina r e t i c u l a t a i i i l e n z i e s i i ) and Usnea spp. were found i n rumens of b l a c k t a i l e d deer throughout the year, though l a r g e s t volumes (up to 24%) occurred i n winter (Book et a l . 1972). The use of a r b o r e a l l i c h e n s on the lower branches c f t r e e s by c a r i b o u and r e i n d e e r (Rangifer spp.) has been documented by many authors. Nasimovitch (1955) repo r t e d t h a t , i n heavy s n o w f a l l areas i n the S o v i e t Union, w i l d r e i n d e e r i R a n j i f e r tarandus tarandus) fed more on a r b o r e a l l i c h e n s than on t e r r e s t r i a l l i c h e n s , whereas i n low s n o w f a l l areas, t e r r e s t r i a l l i c h e n s were the main winter food. In the mountains of the Kola peninsula and the southern U r a l s , r e i n d e e r l e f t a l p i n e areas when snow became dense and c r u s t e d to descend i n t o the f o r e s t , changing t h e i r d i e t from t e r r e s t r i a l t o a r b o r e a l l i c h e n s , Skuncke (1969) noted t h a t i n Sweden, a r b o r e a l l i c h e n s are o f t e n necessary t o r e i n d e e r , e s p e c i a l l y i n l a t e w i n t e r , and the Lapp herders sometimes f e l l t r e e s or break o f f l i c h e n - b e a r i n g branches f o r the animals. Sulkava and H e l i e (1975) found t h a t domesticated r e i n d e e r i n F i n n i s h c o n i f e r f o r e s t s dug i n the snow f o r ground l i c h e n s and grasses u n t i l January when snow depths reached 40-50 cm. During the r e s t of the winter they f e d almost e n t i r e l y on beard l i c h e n s ( A l e c t o r i a j u b a t a ^ - A f c . f r e i g n t j i ^ JL. A l J l g x a j , km, sarmentosaJL a v a i l a b l e from the lower branches of t r e e s and from f a l l e n branches brought down by storms or melting snow. F e l l e d t r e e s a t l o g g i n g s i t e s were used as sources of l i c h e n s by about o n e - t h i r d of the animals. 10 In North America, there has been some co n t r o v e r s y concerning the importance of l i c h e n s , i n c l u d i n g a r b o r e a l l i c h e n s , to c a r i b o u (Rangifer tarandus cariboul. of the boreal f o r e s t . Cringan (1957) r e p o r t e d t h a t l i c h e n s were more important as winter food than woody browse to c a r i b o u of the S l a t e I s l a n d s i n Lake s u p e r i o r . He argued t h a t a r b o r e a l l i c h e n s , produced mainly i n climax f o r e s t s , were p o t e n t i a l l y more important than t e r r e s t r i a l l i c h e n s because of the s u s t a i n e d supply a v a i l a b l e from f a l l e n t r e e s , independent of c a r i b o u p o p u l a t i o n l e v e l s . His c o n c l u s i o n that c a r i b o u r e g u i r e climax f o r e s t s t o maintain steady p o p u l a t i o n s was l a t e r guesticned by Euler et a l . (1976), who observed t h a t S l a t e I s l a n d c a r i b o u made e x t e n s i v e use of immature as w e l l as mature stands i n winter. S c o t t e r (1964) presented anecdotal evidence f o r the importance of a r b o r e a l l i c h e n s t o c a r i b o u , p a r t i c u l a r l y when snow was deep and c r u s t e d , but found o n l y small amounts of A l e c t o r i a and Usnea spp. i n rumens c o l l e c t e d i n December and January i n the northern b o r e a l f o r e s t ( S c o t t e r 1967). He s p e c u l a t e d t h a t l a r g e r g u a n t i t i e s might have been found had samples been taken l a t e r i n winter. A h t i and Hepburn (1967) summarized a v a i l a b l e i n f o r m a t i o n on the food h a b i t s of O n t a r i o c a r i b o u i n winter. Ground l i c h e n s seemed t o be p r e f e r r e d to t r e e l i c h e n s where both were a v a i l a b l e . In the southern part of the b o r e a l f o r e s t , where snow was deeper and t e r r e s t r i a l l i c h e n s l e s s abundant, a r b o r e a l l i c h e n s were thought t o be more important than i n the north. 1 1 Bergerud (1972) found t h a t a r b o r e a l l i c h e n s represented 54% of the contents of e i g h t rumens of Newfoundland c a r i b o u c o l l e c t e d i n March under severe snow c o n d i t i o n s . In g e n e r a l , the animals fed i n c r e a s i n g l y on a r b o r e a l l i c h e n s as the snow became deeper. Bergerud considered that a r b o r e a l l i c h e n s were used because they were a v a i l a b l e , but that they were not n e c e s s a r i l y p r e f e r r e d or r e q u i r e d . . : M i l l e r (1976) noted t h a t c a r i b o u i n northwestern Manitoba changed t h e i r d i e t from t e r r e s t r i a l l i c h e n s and g r a s s l i k e p l a n t s t o a r b o r e a l l i c h e n s and woody browse i n l a t e winter when a hard c r u s t formed on the snow. Consumption of e p i p h y t i c l i c h e n s i n winter i s not always a s s o c i a t e d with deep or c r u s t e d snow. In Stardom's (1975) study area i n s outheastern Manitoba, a r b o r e a l l i c h e n a v a i l a b i l i t y was g r e a t e s t i n open tamarack ( L a r i x l a r i c i n a ) and black spruce (Picea mariana) bogs. Ground l i c h e n a v a i l a b i l i t y was g r e a t e s t on jack pine jP i n u s banksiana) r i d g e s where the snow cover was t h i n n e r because o f i t s i n t e r c e p t i o n by t r e e s , and s o f t e r because there was no wind c r u s t . Caribou f e d mainly on a r b o r e a l l i c h e n s i n bogs u n t i l snow depths reached 60 cm and wind c r u s t s developed. The animals then moved from the bogs to r i d g e s , where they f e d mainly on t e r r e s t r i a l l i c h e n s . Apparently they were able to o b t a i n ground l i c h e n s by c r a t e r i n g d e s p i t e average snow depths which reached 65-70 cm on the r i d g e s d u r i n g a severe winter. A r b o r e a l l i c h e n s are the p r i n c i p a l winter food of mountain c a r i b o u i n B r i t i s h Columbia. In Wells Gray Park, c a r i b o u spent the e a r l y p a r t of the winter i n low e l e v a t i o n f o r e s t s where they fed f i r s t on ground v e g e t a t i o n and then, a f t e r snow became deep. 12 on a r b o r e a l l i c h e n s (Edwards and B i t c e y 1959) . Twenty-six of seventy (37%) f e e d i n g o b s e r v a t i o n s made during t h i s p e r i o d noted the consumption of a r b o r e a l l i c h e n (Edwards and B i t c e y 1960). By January the snow was f i r m enough to support the animals and they migrated to tim.ber.line f o r e s t s where they remained u n t i l A p r i l (Edwards and B i t c e y 1959). The consumption of a r b o r e a l l i c h e n s represented 11 of 12 (92%) f e e d i n g o b s e r v a t i o n s between January and A p r i l (Edwards and R i t c e y 1960} * Freddy (1974) found that i n the S e l k i r k Range i n southeastern B r i t i s h Columbia, mountain c a r i b o u used the spruce-f i r J Pice a enqelmannii - Abies l a s i pear pal f o r e s t zone throughout the w i n t e r , but used the lower p a r t o f the zone more from October to February and the upper part more from March to May. Twenty-three of t h i r t y - t h r e e (70%) f e e d i n g o b s e r v a t i o n s during the p e r i o d October-May c o n s i s t e d of a r b o r e a l l i c h e n s . Caribou fed mainly on l i c h e n s from the lower branches of t r e e s , though f a l l e n t r e e s were a l s o used; deep snowpacks at high e l e v a t i o n s i n c r e a s e d amounts of l i c h e n a v a i l a b l e to c a r i b o u . Freddy c o n s i d e r e d a r b o r e a l l i c h e n s an extremely v a l u a b l e food f o r the S e l k i r k c a r i b o u and concluded t h a t mature s p r u c e - f i r f o r e s t s were probably e s s e n t i a l t o c a r i b o u s u r v i v a l . A r b o r e a l l i c h e n s are a minor component of the d i e t of moose (Alces a l c e s ) i n some areas. Cowan et a l . (1950) i m p l i e d that Usnea barbata was used by moose i n the Quesnel r e g i o n o f B r i t i s h Columbia. Eastman (1978) noted t h a t i n l a t e winter. L o b a r i a pulmonaria c o n s t i t u t e d 10 percent of the rumen c o n t e n t s of moose i n a c o n i f e r o u s f o r e s t near P r i n c e George, B r i t i s h Columbia. Des i Meules (1965) found no evidence of moose fe e d i n g on a r b o r e a l 13 lichens i n Laurentide Park, Quebec, although lichens were abundant in some areas studied. Importance of arboreal lichens to Eocky Mountain elk l£ervus canadensis nelsoni) i s apparently low. Kufeld (1973) summarized the r e s u l t s of 48 food habits studies and mentioned only one (DeNio 1938) i n which lichen consumption was reported. Hash (1973) found that arboreal lichens made up 2,435 of the winter diet and occurred in 35% of 57 elk rumens co l l e c t e d in northern Idaho. Murie (1951) described arboreal lichens as "very palatable" to Eoosevelt elk (Cervus canadensis roosevelti) on the Olympic Peninsula and noted that they were consumed both from f a l l e n branches and from trunks of standing trees. Packee (1975) rated use of Usnea spp. by Eoosevelt elk as medium i n winter and spring, but added that the importance of lichens i n the diet of elk i s poorly understood, Arboreal lichens are important i n the winter d i e t s of some ungulates i n the Soviet Union. Nasimovitch (1955) reported that they are a p r i n c i p a l winter food of musk deer (Moschus moschiferus) throughout i t s range, and of moose i n parts of i t s range. Arboreal lichens are important secondary foods i n parts of the range of red deer iCervus elaphus) , sika deer (C. nippon) , roe deer (Capreolus capreolus) . b i s o n (Bison bonasusl , and chamois (Rupicapra rupicapra) -. 14 j M i i i i i 2 i a l Value of Arboreal Forage Lichens Available data on the chemical composition of arboreal lichens of the genera Alectoria Isensu lato) and Usnea are summarized i n Table I.,In general, crude protein lev e l s are lower than the 6-7% generally recommended as minimum maintenance level s for deer (French et a l . 1956; Dietz 1972). Protein l e v e l s in the study area are the lowest that are reported, although they might have been higher had Rochelle sampled Brvoria spp. as well as Alectoria sarmentosa. The Ca-P r a t i o reported by Bergerud (1972) i s favourable. The d i g e s t i b i l i t y values reported by Bochelle (1978) are extremely high - 75-85% compared to 20-50% for other major forage species during winter. Furthermore, Alectoria sarmentosa appears to have a synergistic e f f e c t on net d i g e s t i b i l i t y when combined with ether forage plants (J.A. Hochelle 1 unpublished data). Seme writers have suggested that Alectoria spp. are a good energy source (Scotter 1965, 1972), but preliminary r e s u l t s of analysis of v o l a t i l e f a t t y acids in Alectoria sarmentosa indicate that the lichens may be r e l a t i v e l y low i n available energy (Bochelle unpublished data). The n u t r i t i o n a l value of Alectoria (s.1.) i s discussed more f u l l y by Bochelle (1978). 1 Weyerhaeuser Co. Ltd., Centralia, Washington. TABLE I. Nutr i t iona l value of arboreal forage l ichens. In V itro Dry Matter D i g e s t i b i l i t y [%) Crude Protein (%) Fat (*)• Fi ber (*) Ash (%) NFE (%) Ca P (*) (*) Rochelle 1978 A. sarmentosa 75 - 85 1 <2 Sulkava & Helle 1975 Alectoria spp. 4.2 Bergerud 1972 "A. jubata"2 5.78 0.56 7.47 1.45 84.74 0.11 0.07 Packee 1975 A. sarmentosa Usnea subfloridana 3.06 9.25-9.38 Schroeder 1974 "A. rjubata"3 4.6 2.34 4.39 1.73 86.75 A. sarmentosa 2.55 7.05 1.42 0.57 88.42 Scotter 1965 "A. jubata"h 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 B lackta i led deer; winter Mainly B. trichodes subsp. americana (Brodo & Hawksworth 1977: 45). Bryoria spp. B. lanestris and B. simplicior (Brodo & Hawksworth 1977: 45). 16 Ecolo<jI of A l e c t o r i o i d L ichens E c o l o g i c a l s t u d i e s must be based on a s a t i s f a c t o r y taxonomy. U n t i l r e c e n t l y , the l a c k of a workable and widely accepted c l a s s i f i c a t i o n of A l e c t o r i o i d l i c h e n s was an o b s t a c l e to e c o l o g i c a l r e s e a r c h . The p u b l i c a t i o n of Brodo and Hawksworth*s (1977) monograph d i s p e l l e d much co n f u s i o n and l a i d the groundwork f o r improved e c o l o g i c a l s t u d i e s . These authors d i v i d e d the former genus A l e c t o r i a i n t o f o u r genera w i t h i n North America - A l e c t o r i a , B r y o r i a , Pseudephebe, and S u l c a r i a . As S u l c a r i a does not occur i n B r i t i s h Columbia, and Pseudephebe i s a t e r r e s t r i a l genus, a l l the A l e c t o r i o i d l i c h e n s d i s c u s s e d i n t h i s t h e s i s belong to the genera AjLectoria and gryoria,, These genera are e a s i l y d i s t i n g u i s h e d i n the f i e l d by c o l o u r -A l e c t o r i a spp. are gray-green t o yellow-green, whereas B r y o r i a spp. are brownish. In some cases, i d e n t i f i c a t i o n to s p e c i e s i s d i f f i c u l t or i m p o s s i b l e i n the f i e l d . Ecology at the s p e c i e s l e v e l i s not considered i n the d i s c u s s i o n that f o l l o w s . A l e c t o r i a and B r y o r i a are e s s e n t i a l l y c o l d c l i m a t e genera, and most s p e c i e s are e i t h e r n o r t h e r n or a s s o c i a t e d with mountains (Brodo and Hawksworth 1977). Some of the more important e c o l o g i c a l f a c t o r s determining the abundance of A l e c t o r i o i d l i c h e n s w i t h i n t h e i r geographical range are s o l a r energy, water, n u t r i e n t s , s u b s t r a t e c h a r a c t e r i s t i c s , d i s p e r s a l o p p o r t u n i t i e s , and time. F a c t o r s t h a t c o n t r i b u t e to the removal of l i c h e n s such as breakage through the a c t i o n o f wind and r a i n , and consumption by animals are a l s o important. 17 The importance of l i g h t and humidity i n determining the abundance of A l e c t o r i a (s.l.) has been pointed out by other workers (Schroeder 1974) and may be inferred from the d i s t r i b u t i o n of these lichens i n trees. In general, l i g h t i n t e n s i t y increases and r e l a t i v e humidity decreases with increasing distance from the ground i n a forest stand (Onura et a l , 1955; Geiger 1966:298). On the basis of physiological studies of several species of epiphytic lichens and mosses at various illuminations and humidities, Hosokawa et a l . (1964) concluded that 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 at the lower l i m i t s by l i g h t and at the upper l i m i t s by water. In the humid west coast forest 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 positions where forests are denser, A l e c t o r i a i s found nigh i n the crown. On xeric s i t e s where stands are more open, Alectoria extends further down the tree and Bryoria i s present near the top of the tree. Both genera are sparse in the lower parts of trees except at very open s i t e s . This d i s t r i b u t i o n pattern, documented by the work of Szczawinski (1953) and Pike et a l . (1975) strongly suggests that i n the coastal f o r e s t , l i g h t i s the dominant climatic factor 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 Bryoria species of t h i s forest type are more photophilous than the Alectoria species. In the i n t e r i o r of B r i t i s h Columbia, with i t s more continental climate, humidity i s more frequently a l i m i t i n g factor than i n the moist, coastal forest. Edwards et a l . (1960) noted that in some dry forests of S e l l s Gray Park, Al e c t o r i a (s.l.) was present i n the lower parts of the crowns, but absent 18 i n the upper p a r t s . I n parts of the S e l k i r k Range of southeastern B r i t i s h Columbia, occurrence of A l e c t o r i a and B r y o r i a appears t o be l i m i t e d by humidity. Thus, e x c e s s i v e t h i n n i n g can make f o r e s t s too dry f o r continued l i c h e n growth (Schroeder 1974). Many authors have noted, i n c o a s t a l areas as well as i n t e r i o r areas, t h a t abundance of A l e c t o r i a ( s . l . ) i n c r e a s e s with e l e v a t i o n (Ahti 1962, B a r i c h e l l o 1975, Rochelle 1978). Although other v a r i a b l e s a l s o change with e l e v a t i o n , i n c r e a s e d l i c h e n abundance a t higher e l e v a t i o n s i s probably due i n p a r t to i n c r e a s e d humidity, a s s o c i a t e d with h i g h e r p r e c i p i t a t i o n , lower summer temperatures, and t h e r e f o r e lower evaporation r a t e s . The n u t r i e n t supply of l i c h e n s i s d i s s o l v e d i n water t h a t comes i n cont a c t 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 s s e l e c t i v e l y accumulate a wide range of organic and mineral n u t r i e n t s (Hale 1974). A l e c t o r i o i d l i c h e n s have green algae as phyccbionts, and are t h e r e f o r e dependent on e x t e r n a l n i t r o g e n sources such as n i t r a t e s and o r g a n i c n i t r o g e n , which are leached from the t r e e canopy by r a i n f a l l . In the study a r e a , n u t r i e n t -poor f o r e s t s i t e s with good l i g h t p e n e t r a t i o n o f t e n have much higher l i c h e n abundance than r i c h e r s i t e s with a denser t r e e canopy. I t t h e r e f o r e seems u n l i k e l y t h a t n u t r i t i o n i s a major f a c t o r l i m i t i n g growth of A l e c t o r i o i d l i c h e n s , except p o s s i b l y very high i n the crowns of t r e e s 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 has not been enriched by l e a c h i n g . A i r p o l l u t i o n i s thought t o be an important f a c t o r i n the present d i s t r i b u t i o n of s e v e r a l s p e c i e s of B r y o r i a i n Europe, and 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 and i n d u s t r i a l 19 areas i s limited (Brodo and Hawksworth 1977). Sheridan et a l . (1976) found that biomass of Alectoria (Bryoria) fremontii was exremely reduced below control s i t e values as much as 16 km from a pulp m i l l i n Montana. The information currently available indicates that species of A l e c t o r i a and Bryoria d i f f e r in t h e i r s e n s i t i v i t y to a i r p o l l u t i o n , but i t i s not yet possible to describe their r e l a t i v e s e n s i t i v i t i e s (Brodo and Hawksworth 1977) . The Best common substrates of corticolous A l e c t o r i a (s.l.) are coniferous trees and trees with s i m i l a r bark c h a r a c t e r i s t i c s such as birch (Brodo and Hawksworth 1977). It i s not clear whether these lichens actually prefer a c i d i c bark due to some physiological requirements, or whether they require the climate and general environment of coniferous forests and simply occupy the most available substrate. According to Brodo and Hawksworth, the l a t t e r p o s s i b i l i t y i s more l i k e l y , as a wide variety of trees within each forest type are used as substrates. Although su b s t r a t e - s p e c i f i c i t y of A l e c t o r i o i d lichens i s r e l a t i v e l y lew compared to seme corticolous lichens, especially species which are i n closer contact with the substrate (Brodo 1973), some important differences i n abundance of Ale c t o r i a is.1.) cn conifer species 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 trees and their s u i t a b i l i t y as habitats for epiphytes have been discussed by Szczawinski (1953) and Ahti (1962). Moisture capacity, mineral content, pH, physical texture, 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 potentially important i n determining s u i t a b i l i t y of a tree as a substrate (Brodo 1973). 20 The establishment of a lichen species i n an area depends on the a v a i l a b i l i t y of viable propagules, either sexual or vegetative. Spore-producing structures are rather rare among Alectoria (s.l.) and most species depend mainly on vegetative methods of propagation (Brodo and Hawksworth 1977) . Vegetative reproduction by windblown thallus fragments i s of major importance i n short-range di s p e r s a l , but there i s some evidence that species of Al e c t o r i a and Bryoria that produce soredia -clumps of a l g a l c e l l s surrounded by fungal hyphae - have an advantage i n long-range dispersal over species that do not (Stevenson unpublished data). Fragmentation and i s o l a t i o n of old forest stands by human settlement, agriculture, and logging may severely r e s t r i c t the d i s t r i b u t i o n of some lichen species that cannot disperse e f f e c t i v e l y over long distances. Rose (1976) found that some arboreal lichens i n Great B r i t a i n , including several species of Alectoria ( s . l . ) . are presently limited to ancient, undisturbed forests and are apparently unable to colonize suitable habitats at a distance. 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 of some species i n North America. Studies of e f f e c t i v e dispersal distances of lichens are needed, as they have implications f o r forest management, p a r t i c u l a r l y with respect to siz e and spacing of clearcuts. The abundance of A l e c t o r i a (s.l.) in a suitable f o r e s t s i t e depends on how much time has been available for establishment and growth of the lichens. Growth rates of lichens are generally slew compared to other plant groups. 5There are many technical d i f f i c u l t i e s involved i n measuring growth rates of fruticose 21 arboreal lichens, and consequently l i t t l e useful information i s available on growth rates of a l e c t o r i o i d lichens. The r e l a t i v e l y great abundance of a l e c t o r i a (s.l.) i n mature forest stands as opposed to immature stands has been noted by many authors (Edwards et a l . 1960, Ahti 1962, Ahti and Hepburn 1967). There i s a need for research on the dynamics of lichen development i n the course of forest succession, and associated changes i n lichen bicmass. 22 I I I . QUANTIFICATION OF BIOMASS OF ARBOREAL FORAGE LICHENS In order to study l i c h e n p r o d u c t i o n and r e l a t e i t to other ecosystem f a c t o r s , i t was necessary to have methods f o r measuring l i c h e n abundance. The f i r s t o b j e c t i v e was t o develop and e v a l u a t e techniques of q u a n t i f y i n g biomass o f a r b o r e a l forage l i c h e n s . Three methods o f a s s e s s i n g l i c h e n biomass were used. L i c h e n bicasass was measured by sampling the l i c h e n s from f e l l e d t r e e s . A system of v i s u a l estimates of l i c h e n abundance was developed, r e l a t e d to biomass e s t i m a t e s obtained by i n t e n s i v e sampling, and extended to other areas as an i n v e n t o r y t o o l . The value of l a r g e - s c a l e a i r photography as a p o t e n t i a l i n v e n t o r y t o o l was t e s t e d . In each case, q u a n t i f i c a t i o n was l i m i t e d to a r b o r e a l l i c h e n s of the genera A l e c t o r i a and B r y o r i a * Many t e c h n i c a l and s t a t i s t i c a l problems are a s s o c i a t e d with attempts to q u a n t i f y the biomass of epiphytes on an a r e a l b a s i s . Access to epiphytes growing i n the crown of a l a r g e t r e e i s d i f f i c u l t ; c l i m b i n g the t r e e r e q u i r e s e l a b o r a t e equipment (e.g. Pike et a l . 1972, 1977), and f e l l i n g the t r e e d e s t r o y s and s c a t t e r s some of the epiphytes. I n t e n s i v e biomass sampling can produce a good estimate of epiphyte q u a n t i t i e s i n a s m a l l area, but i s too time-consuminq to be used f o r i n v e n t o r y purposes. Rapid estimates of epiphyte abundance s u i t a b l e f o r i n v e n t o r y purposes are o f t e n i m p r e c i s e and are d i f f i c u l t t o r e l a t e to a c t u a l epiphyte biomass. 23 Some investigators (Scotter 1962, Andre et a l . 1975, Hein and Speer 1975) have determined lichen biomass by removing and weighing the entire l i c h e n load from one or more trees within a forest type. These authors studied f a i r l y small trees (25 m or l e s s ) ; the method would be i n e f f i c i e n t i f trees were large and lichen guantiti.es great. Various subsampling schemes have been used to study larger trees. Edwards et a l . (1960) counted the branches in each 3 m height i n t e r v a l , selected two representative branches for sampling, and calculated biomass within each height i n t e r v a l by mu l t i p l i c a t i o n . Forman (1975) selected a representative one-third of the crown of large trees for sampling and multiplied the res u l t s by three. Pike et a l . (1972, 1977) developed a very intensive method of c a l c u l a t i n g t o t a l epiphyte biomass on a large (65 m) standing Douglas-fir (Pseudctsuqa menziesii) tree. Each branch was assigned an "importance value" on the basis of detailed observations of i t s size and epiphyte biomass, which were made from a climbing position on the trunk of the tree. Importance values were used as the basis f o r a random proportionate scheme of sampling branches for detailed study. These values were l a t e r used to extrapolate epiphyte biomass from sample branches to the entire tree by means of a l i n e a r regression of biomass on epiphyte importance value. Most researchers (Edwards et a l . 1960, Scotter 1962, Andre et a l . 1975) have extended th e i r biomass t o t a l s for i n d i v i d u a l trees to an areal basis by multiplying by trees per unit area. Unless the forest i s quite uniform or the sample size very large, t h i s method could be expected to lead to errors. Forman 24 (1975) found 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 epiphyte g u a n t i t i e s and t r e e diameter, and used r e g r e s s i o n s of epiphyte q u a n t i t y on DBH to c a l c u l a t e t o t a l s f o r unsampled t r e e s . Q u a n t i f i c a t i o n o f forage l i c h e n s on an i n v e n t o r y b a s i s has not o f t e n been r e p o r t e d , other than simple r a t i n g s of l i c h e n abundance cn a low-raedium-high s c a l e ( B a r i c h e l l o 1975, Jones 1975). A n t i (1962) used estimates of percent l i c h e n cover, r a t i n g s of abundance of twigs on a 1-5 s c a l e , and numbers of stems/acre to c a l c u l a t e a "range index" of l i c h e n abundance on the lower 3 meters of t r e e s i n H e l l s Gray Park. T h i s method gave r e s u l t s which c l o s e l y p a r a l l e l e d the ge n e r a l impressions of l i c h e n abundance recorded by A h t i i n h i s f i e l d notes. D. R u s s e l l 1 and S. R u s s e l l (pers. comm.) developed a system of es t i m a t i n g l i c h e n abundance on the lower p a r t s o f t r e e s i n i n t e r i o r B r i t i s h Columbia f o r e s t s which i n v o l v e d v i s u a l l y d i v i d i n g the t r e e i n t o s e c t i o n s , counting branches i n each s e c t i o n , and e s t i m a t i n g percent cover on r e p r e s e n t a t i v e branches. N e i t h e r cf these systems i s s u i t a b l e f o r e s t i m a t i n g l i c h e n abundance on t h e e n t i r e crown of t r e e s i n the coast f o r e s t . A l i t e r a t u r e search r e v e a l e d no previous attempts to inve n t o r y a r b o r e a l l i c h e n abundance by means of remote sensing, although t e r r e s t r i a l l i c h e n communities have been mapped frcm a i r photographs ( B u t t r i c k 1978). Despite the absence of previous r e s e a r c h , i t seemed reasonable to expect that d i f f e r e n c e s i n the appearance of the f o r e s t canopy due to v a r i a t i o n s i n l i c h e n 1 Yukon Game Branch, 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 be d e t e c t a b l e i n l a r g e - s c a l e a i r photographs. Since the i n i t i a t i o n of the present study, Harestad (1978) used normal c o l o u r o b l i q u e a i r photographs to assess l i c h e n abuncance i n the study area. He used a dot g r i d technique to q u a n t i f y l i c h e n abundance on 23 s i t e s , i n c l u d i n g 9 p l o t s on which I q u a n t i f i e d l i c h e n abundance using ground measurements. When data were transformed to l o g | 0 # a r e g r e s s i o n of a i r photo p r e d i c t i o n s on l i c h e n bit-mass as measured on the ground i n t h i s study was 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 < 0.06) and had an r z of 0.41 (Harestad 1978) . 26 Methods Visual estimates of lichen abundance Quantification of lichen abundance by v i s u a l estimates was conducted on 14 plots. Procedures for l a y i n g out plots and selecting trees for lichen estimates varied s l i g h t l y depending on whether or not the plot was used for testing photo-prediction of lichen abundance. Nine non-photo plots were chosen to represent areas for which information on winter deer use or lichen l i t t e r f a l l was available (Jones 1975, Rochelle 1978, Harestad 1978). These plots were sguare and were i n i t i a l l y 0.1 ha i n area, but after completing three plots, I increased plot size to 0.2 ha to improve representation of the f o r e s t type. A l l trees within the plot which were 18 cm or more in diameter were inventoried. Trees were temporarily l a b e l l e d with numbered p l a s t i c cards, and the following information was recorded for each tree: - DBH (diameter at 1.5 m above the point of germination); - species; - dominance class (dominant, codominant, intermediate or overtopped); - l i v i n g or dead (dead trees 10 m or more i n height were further c l a s s i f i e d as dead potential (i. e . merchantable) or dead useless (B.C. Forest Service 1973); and 27 - l e a n ( t h i s was noted i f t r e e s leaned more than approximately 10 degrees from v e r t i c a l ) . i Frequencies of i n d i v i d u a l species-dominance c l a s s e s were then compiled and 15 t r e e s s e l e c t e d f o r l i c h e n e s t i m a t e s using a random p r o p o r t i o n a t e scheme (Cochran 1963:89), such t h a t the chance of any p a r t i c u l a r t r e e being i n c l u d e d i n the sample was p r o p o r t i o n a t e to the number of t r e e s i n i t s species-dominance c l a s s . Dead t r e e s , l e a n i n g t r e e s , and overtopped t r e e s were omitted from the s e l e c t i o n process. In the case of the f i v e a i r photo p l o t s , p l o t boundaries were l a i d out w i t h i n the area of s t e r e o o v e r l a p cn the photographs, and l a t e r l o c a t e d on the ground. A i r photo p l o t s were v a r i a b l e i n s i z e and shape because of topographic i r r e g u l a r i t i e s . In the l a b o r a t o r y , a l l l i v i n g t r e e s having bases i n s i d e the p l o t boundaries on the a i r photograph were i d e n t i f i e d to s p e c i e s and assigned a dominance c l a s s . F i f t e e n t r e e s were s e l e c t e d f o r l i c h e n estimates u s i n g the random p r o p o r t i o n a t e sampling scheme. A f t e r p l o t boundaries were l a i d out on the ground, a t r e e i n v e n t o r y was c a r r i e d out and s p e c i e s i d e n t i f i c a t i o n s made from the photograph were c o r r e c t e d where necessary. Standard f o r e s t mensuration techniques were used on a l l p l o t s to o b t a i n t r e e heiqht (TH), crown le n g t h (CL), and crown width (Ca^ ) of t r e e s which had been s e l e c t e d f o r l i c h e n estimates. Crown l e n g t h was d e f i n e d as the v e r t i c a l d i s t a n c e , i n meters, from the highest l i v i n g p a r t o f the crown t o the lowest l i v i n g part of the crown. Crown width was d e f i n e d as the average diameter, i n meters, of the crown at the widest p a r t , and was 28 determined by averaging measurements taken p e r p e n d i c u l a r to the slope and p a r a l l e l to the sl o p e ( f i g . 2). Lich e n e s t i m a t e s were made from a viewing p o s i t i o n from which as much as p o s s i b l e o f the crown could be seen. Sometimes, i t was necessary to s h i f t to a second viewing p o i n t to see a p o r t i o n of the t r e e . The l i v e crown was v i s u a l l y d i v i d e d i n t o four l a y e r s ( F i g . 3), and the f o l l o w i n g e s t i m a t e s were made f o r each l a y e r with the a i d of b i n o c u l a r s ; - crown width (CW» ) : The crown width of the t r e e , which had alre a d y been measured, was used as a r e f e r e n c e p o i n t f o r e s t i m a t i n g the width of each l a y e r , i n meters ( i = - % t r e e cover ( T C j ) : For each l a y e r , percent t r e e cover was estimated within an imaginary r e c t a n g l e having dimensions CWj x CL/4 ( F i g . 3 ) . Tree cover i n c l u d e d trunk, branches, twigs, f o l i a g e , and epiphytes — anything which was part of the t r e e or which grew on i t . - % A l e c t o r i a ( s . l . ) (A|): The percent of t r e e cover c o n s i s t i n g of A l e c t o r i a and B r y o r i a was estimated f o r each l a y e r . To assess i n t e r o b s e r v e r r e l i a b i l i t y , a f i e l d a s s i s t a n t and I made independent v i s u a l estimates of l i c h e n abundance on the t r e e s i n P l o t 1. A r e g r e s s i o n o f one s e t of o b s e r v a t i o n s on the other was c a l c u l a t e d , and Student's t - t e s t (Dixon and Massey 1969:114) used t o determine whether the i n t e r c e p t d i f f e r e d s i g n i f i c a n t l y from zero, or the s l o p e d i f f e r e d s i g n i f i c a n t l y from one. 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 three estimates described here were made on a l l trees studied. During the second summer, only Alectoria was estimated. The reason i s given in the discussion of computations and re s u l t s . Air photo quantification of lichen abundance Eefore f l y i n g the area, I photographed lichens using various combinations of films and f i l t e r s . Contrast between conifer foliage and lichens was best with colour-infrared f i l m , a Wratten 12 f i l t e r , and two CC20 Magenta f i l t e r s (Fig. 4) . Ae r i a l photography was done by Integrated Resources Photography, Ltd. of Vancouver on June 5 and 6, 1976, using Kodak Aercchrcme Infrared 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 wing-tips of a Cessna 180 were exposed simultaneously to produce stereo images. A l l areas were flown at a height of approximately 152 m above the tops of the trees to obtain imagery at the scale 1:2000; for some f l i g h t lines 1;4000 and 1;8000 photographs were also obtained. The use of a i r photographs to detect lichen abundance was tested by two methods: densitometry and photo inte r p r e t a t i o n . Densitometry i s a technigue that may be used to quantify dye-layer density i n a colour photograph. Colour-infrared positive transparencies consist of three developed dye-layers and a stable base. The densities cf these dye-layers are inversely related to the spectral reflectance pattern of the subject photographed, although other factors influence dye-layer density as well (Murtha 1972). In colour-infrared f i l m , spectral A Figure 4. Alectoria savmentosa on Douglas-f i r : normal colour f i l m (A) and colour in f rared 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 r e f l e c t a n c e i n the n e a r - i n f r a r e d region of the spectrum a f f e c t s the cyan dye-forming l a y e r . A densitometer i s used with c o l o u r e d f i l t e r s to measure the d e n s i t y of each dye l a y e r s e p a r a t e l y i n 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 i f e r needles r e f l e c t s t r o n g l y i n the i n f r a r e d r e g i o n of the spectrum ( S t e i n e r and Gutermann 1966, Gates 1970, Hurtha 1972). Since the dyes used i n c o l o u r - i n f r a r e d f i l m are s u b t r a c t i v e , v i s u a l l y green c o n i f e r needles appear red to magenta when the f i l m i s developed ( F i g . 4 ) . In c o n t r a s t , A l e c t o r i a sarmentosa appears very l i g h t . Data on 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 A l e c t o r i a sarmentosa are not a v a i l a b l e , but high r e f l e c t a n c e at a l l wavelengths has been repo r t e d f o r other l i g h t - c o l o u r e d l i c h e n s ( S t e i n e r and Gutermann 1966, Gates 1970). Such a 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 would r e s u l t i n a l i g h t image on c o l o u r - i n f r a r e d f i l m . A Macbeth TR-524 Transmission R e f l e c t i o n Densitometer was used with Wratten f i l t e r s 92 ( r e d ) , 93 (green), and 94 (blue) t o measure dye-layer d e n s i t i e s of the t r e e s i n the photo p l o t s f o r which l i c h e n biomass had been c a l c u l a t e d . The sample t r e e s were d i v i d e d i n t o c l a s s e s of high and,low l i c h e n biomass as determined from ground p l o t s , separated by the mean. S t u d e n t s t t e s t (Dixon and Massey 1969:114) was used t o compare the mean de n s i t y of each d y e - l a y e r , and the red:green f i l t e r r a t i o , of the two c l a s s e s f o r a l l t r e e s together and f o r sepa r a t e s p e c i e s . The a b i l i t y of photo i n t e r p r e t e r s to p r e d i c t l i c h e n abundance on i n d i v i d u a l t r e e s and on s i t e s was t e s t e d . Ten s u b j e c t s with v a r y i n g degrees o f experience with photo i n t e r p r e t a t i o n and with l i c h e n e c c l o g y were used. 33 The interpreters were shown normal colour and colour-infrared photographs, taken from the ground, of Alectoria -covered Bouglas-fir trees (Fig. 4). They were then shown two stereo pairs, cne representing a plot with low l i c h e n biomass (Fig. 5) and one representing a plot with high l i c h e n biomass (Fig. 6), i n which a t o t a l of 34 trees were l a b e l l e d according to lichen abundance. The trees l a b e l l e d i n the photographs were ones on which biomass sampling or v i s u a l estimates had been carried out. They were rated on a 6-point scale T (trace) through 5 (abundant), corresponding broadly to kilograms of lichen on the trees. Interpreters were then shown three additional stereo pairs of ground plots, each with 15 trees c i r c l e d , and asked to rate lichen abundance on those trees, using the l a b e l l e d sets for reference. C i r c l e d trees were ones for which visual estimates had been made. F i n a l l y , 9 of the 10 interpreters were asked to rate general l i c h e n abundance on eleven stereo pairs of ground plots as high,, medium, or low, using four labelled photo pairs for reference. In addition to sampling other photo interpreters, I made predictions of lichen abundance on trees i n photo plots selected for v i s u a l estimates before locking at the trees i n the f i e l d . I did not make predictions for s i t e s because I had f i e l d knowledge of t h e i r lichen abundance. To determine whether interpreter ratings of lichen abundance on i n d i v i d u a l trees were correlated with calculated lichen biomass, Spearman's rank correlaton c o e f f i c i e n t was used (Conover 1971:245). Spearman's rho or " r " was calculated for the ratings of each interpreter on each of the three plots and used Figure 5. Colour infrared a i r photographs of p lot with low l ichen biomass. 35 36 to t e s t f o r p o s i t i v e c o r r e l a t i o n of r a t i n g s with biomass. Ihe i n t e r p r e t e r s 1 rho sco r e s were used as data f o r the Mann-8hitney t e s t (Concver 1971:224) to determine w h e t h e r , a b i l i t y t o judge 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 photo i n t e r p r e t a t i o n experience or by f i e l d knowledge of A l e c t o r i a ( s . l . ) e cology. Subjects who had previous experience with i n t e r p r e t a t i o n of black and white, 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 photographs were c o n s i d e r e d t o have a high l e v e l of photo i n t e r p r e t a t i o n experience; s u b j e c t s who had experience with only normal c o l o u r and black and white a i r photographs, or l e s s , were considered i n e x p e r i e n c e d i n t e r p r e t e r s . Subjects who had been with me i n the study area d u r i n g the f i e l d work were co n s i d e r e d to have knowledge of A l e c t o r i a ( s . l . ) ecology; o t h e r s were cot. For s t a t i s t i c a l a n a l y s i s , i n t e r p r e t e r s ' judgements of l i c h e n abundance on s i t e s as low, medium, or high were g u a n t i f i e d as 1, 2, or 3. The mean r a t i n g f o r each s i t e was c a l c u l a t e d and Spearman's rank c o r r e l a t i o n c o e f f i c i e n t was used to determine whether mean r a t i n g s f o r s i t e s were c o r r e l a t e d with c a l c u l a t e d l i c h e n biomass t c t a l s f o r s i t e s . Biomass sampling Biomass sampling was c a r r i e d out durin g the f i r s t summer of fi e l d w o r k i n two photo p l o t s , chosen t o repr e s e n t areas of high and low l i c h e n abundance. In each photo p l o t , f i v e t r e e s were s e l e c t e d t o be f e l l e d f o r biomass sampling. The t r e e s were chosen from those f o r which v i s u a l estimates had been made, and were s e l e c t e d t o represent the major l i c h e n - b e a r i n g , s p e c i e s -dominance c l a s s e s . 37 The four compass d i r e c t i o n s were spray-painted on the holes of the ten 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 three D o u g l a s - f i r broke up badly when they f e l l , but the cedars and the hemlock were reasonably i n t a c t . D i f f e r e n t sampling methods were used i n the two cases. The branch was the basic sampling u n i t f o r i n t a c t t r e e s . 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 rees were sampled; l i c h e n abundance on the lower s i d e s of the t r e e s was assumed eguivalent to that on the upper s i d e s . A l l branches greater than 0.5 meters long p r o j e c t i n g from the upper 180 degrees of the trunk were numbered and the 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 occurred; - branch length (BL); - density f a c t o r (DF) (a s u b j e c t i v e estimate of branch surface area, independent of branch length, rated on a 1 to 5 scale) ; - estimated percent A l e c t o r i a ( s . l . ) cover of branch surface area (%A); and - average length of A l e c t o r i a ( s . l . ) clumps (LA). These data were used i n the f i e l d to c a l c u l a t e a Lichen Estimate (LE) f o r each branch, using the formula: LE = BL x DF x 3SA/10 0 x LA (1) The l i c h e n Estimates were then used as the basis f o r a randcm proportionate sampling 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 sampling, such that the p r o b a b i l i t y of any given branch being s e l e c t e d was proportionate to i t s Lichen Estimate. This sampling scheme, l i k e 38 that proposed by Pike et a l . (1972, 1977), ensured that branches with high lichen biomass were sampled intensively. Lichen biomass was sampled on the trunks of four of these trees. A l l Alectoria and Bryoria was collected from 10 x 30 cm guadrats at 3 m i n t e r v a l s along the sides of the trunk previously l a b e l l e d as north-facing and south-facing. Some adjustment in the aspect of the sample l i n e was necessary when the north or south side was facing d i r e c t l y down. This particular sampling scheme was part of a separate study of 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 trees. Lichen l i t t e r from the immediate area of the f a l l e n trees was collected separately f o r each layer of the crown. The three Douglas-fir were so badly broken up that branches could not be used as sampling units. Two of these trees were sampled by running eight one-meter-wide transects across the area occupied by f a l l e n trees, perpendicular to the trunk. A l l Alect o r i o i d lichens were collected from within these transects, i regardless of whether they occurred on branches or on the ground. Lichens co l l e c t e d from the trunk were bagged separately. The third Douglas-fir had a very small crown and a l l the lichens were collected from the area surrounding t h i s tree. The sample trees were aged by counting annual rings at stump height, approximately 0.6 m. Lichens were hand-picked, bagged and labe l l e d in the f i e l d , and dried i n a hose dryer for storage. In the laboratory, they were cleaned of debris, even-dried at 60-65°C for 24 hours, and weighed with a Mettler P1210 e l e c t r i c scale to the nearest 0.01 g. 3 9 Computations and Results Computing biomass t o t a l s Hand-picking the A l e c t o r i a and Bryoria from the ten sample trees represented two seeks of work for a crew of four. Time reguired for cleaning, drying, and weighing was 54 hours. Lichen weights for the sample branches were then extrapolated to arrive at biomass t o t a l s for trees and for s i t e s . Descriptive data for branches (BL, DF, %&, LA) were used to estimate lichen weights for branches that had not been sampled. A regression of lic h e n weights for the sample branches of a l l trees on Lichen Estimates (Equation 1) gave an r 2 value of C.61. To obtain the best possible l i c h e n weight estimates for branches that were not sampled, several d i f f e r e n t regression equations were evaluated for each tree, and the eguation yi e l d i n g the hiqhest r 2 value was used to calculate lichen bicmass for unsampled branches on that tree. In most cases, the best reqression equation was of the form y = b 0 + ( b j X , ) • ( b i X z ) • (b^Xj) * lhHxH) (2) where x, throuqh x 4 represent BL, DF, %h, LA, but i n one case y = b 0 + fa, (x, x 2x 3x v) (3) accounted for more of the va r i a t i o n i n weights. In general, r 2 values increased with the order in which trees were described (namely, 0.34, 0.30, 0.59, 0.53, 0.74, 0.65, 0.69), suggesting that my a b i l i t y to make consistent estimates improved with practice. 40 Total lichen biomass for each layer of each tree was obtained by summing l i c h e n weights for branches, multiplying by two to account f o r the lower, unsampled side of the tree, and adding the weight of the l i t t e r associated with that layer. Layer weights were summed to obtain t o t a l s for trees. Trunk t o t a l s were calculated and added to the tree t o t a l s . For trees with high lichen loads, lichen biomass on the trunk was consistently 8% of the t o t a l lichen biomass, but for trees with lower lichen loads, percent l i c h e n biomass of the trunk was higher and more variable (Table I I ) . To estimate trunk biomass for trees from which no trunk samples had been taken, 8% of t o t a l lichen biomass was added fo r trees having more than 2C00 g of l i c h e n , and 171 f o r trees having less that 2000 g. A l l biomass t o t a l s and percentages occurring on the trunks of sampled trees are summarized i n Table I I . Ages of the sampled trees, included in Table I I , showed l i t t l e v a riation. Seven of the ten trees were between 350 and 400 years old. To obtain lichen bio mass t o t a l s for each s i t e , i t was necessary to extrapolate to trees that had not been sampled, using visual estimates of lichen abundance. Several regression models were evaluated to determine which had the greatest predictive capacity. Lichen weight for layers of sampled trees was the dependent variable and crown length (CL) or crown length for layers (CL/4), crown width for layers (CH^; where i = 1,...,4), percent tree cover (TCj) , and percent A l e c t o r i a (s.l.) (A^) were the independent variables. A single eguation was used for a l l trees, regardless of species or dominance cl a s s because 41 TABLE II. Biomass of Alectoria on sampled trees Plot Tree Species Dominance Class Age Alectoria Biomass (g) % of Biomass Occurring on Trunk 1 112 WH D 3701 5693 8% 1 13 WH C 3501 7036 a 1 22 WH I 350 1851 a 1 119 WRC C 260 436 15% 1 101 DF C 395 1507 14% 2 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 Symbols used in Table: WH - western hemlock WRC - western red cedar DF - Douglas-fir AF - amabilis f i r D - dominant C - codominant I - intermediate a - not sampled Center of trunk rotten; age estimated. 42 the sample s i z e was too small to warrant separate equations. The r e g r e s s i o n y = 81.617 + 148.58 (A£ x CL) (4) (n = 40; S Y . X = 376.57) had an r 2 value of 0.75. B i o l o g i c a l l y , i t i s reasonable to expect a m u l t i p l i c a t i v e r e l a t i o n s h i p between a measure c f percent cover and a measure cf a v a i l a b l e s u b s t r a t e . To avoid the problem of p o s i t i v e l i c h e n biomass p r e d i c t i o n s f o r t r e e s with no il§£ifiria ( s . l . ) cover, the r e g r e s s i o n was f o r c e d through z e r o , and the eguation Y = 158. 03 (A i x CL) (5) (n = 40; S Y > X = 376.89; r 2 = 0.75) was used to c a l c u l a t e l i c h e n biomass of t r e e s f o r which v i s u a l estimates had been made. Eguation 5 i s i l l u s t r a t e d i n Figure 7. Because the estimates of CW and TC d i d not s i g n i f i c a n t l y improve the r 2 of the r e g r e s s i o n s t e s t e d , they were omitted during the second summer cf f i e l d w o r k . B e s u l t s of the t e s t of i n t e r o b s e r v e r r e l i a b i l i t y are shown i n Figure 8. The r 2 o f the r e g r e s s i o n was 0.85. The i n t e r c e p t d i d not d i f f e r s i g n i f i c a n t l y from 0 (t = 1.07; t 0 o s- = 2.10), and the s lope d i d not d i f f e r s i g n i f i c a n t l y from 1 (t = 1.22; t 0 0 5 = 2.10), i n d i c a t i n g t h a t n e i t h e r set of o b s e r v a t i o n s was s y s t e m a t i c a l l y high or low i n r e l a t i o n to the other. Using c a l c u l a t e d l i c h e n weights f o r 15 t r e e s on each p l o t , l i c h e n biomass f o r s i t e s was computed. Mean l i c h e n biomass f o r each species-dominance c l a s s was used t o c a l c u l a t e t o t a l l i c h e n biomass f o r t h a t c l a s s on each p l o t . Using p l o t areas computed from f i e l d measurements, t o t a l A l e c t o r i a ( s . l . ) biomass (kg/ha) 4 0 0 0-n 3 0 0 0 H rt 2 0 0 0 -— cn 1 0 0 0 -l 1 — i 1 i i r 4 6 8 1© 12 14 P E R C E N T A L E C T O R I A X C R O W N L E N G T H Figure 7. Regression used to ca lcu late l ichen biomass of visual estimate trees (y =• 158.03 x; n = 4.0; S y . x = 376.89;, r 2 - 0.75). 44 Figure 8. Lichen biomass per tree based on independent visual estimates by two observers (y = 0.379 + 0.89 x; n = 19; S y . x ' = 733.51; r 2 = 0.85). 45 was c a l c u l a t e d f o r each s i t e . R e s u l t s a re presented i n Table VI, and s u p p o r t i n g c a l c u l a t i o n s are given i n Appendix I , Res u l t s of a i r photo g u a n t i f i c a t i o n Table I I I summarizes mean o p t i c a l d e n s i t y v a l u e s and red:green f i l t e r r a t i o s of the photo images of high and low l i c h e n biomass t r e e s . Separate comparisons were not made f o r western red cedar because only one t r e e had high l i c h e n biomass. For a l l t r e e s taken together and f o r D o u g l a s - f i r , comparison of o p t i c a l d e n s i t y values 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 with high l i c h e n biomass with those f o r t r e e s with low l i c h e n biomass r e v e a l e d no s i g n i f i c a n t d i f f e r e n c e s . In the case of western hemlock, d i f f e r e n c e s i n the red:green f i l t e r r a t i o between high and low l i c h e n t r e e s approached s t a t i s t i c a l s i g n i f i c a n c e (p<0.07). The d i r e c t i o n o f the d i f f e r e n c e s i n the r a t i o suggests t h a t t r e e s with high l i c h e n biomass had r e l a t i v e l y high r e f l e c t a n c e i n the red r e g i o n of the spectrum, compared to t r e e s with low l i c h e n biomass. T h i s i s c o n s i s t e n t with published r e p o r t s on 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 s of other l i g h t - c o l o u r e d l i c h e n s and of c o n i f e r f o l i a g e ( S t e i n e r and Gutermann 1966, Gates 1970). To f u r t h e r assess t h e value o f the red:green f i l t e r r a t i o as a p r e d i c t o r of l i c h e n abundance I regre s s e d red:green f i l t e r r a t i o s f o r western hemlock t r e e s cn l i c h e n biomass values of those t r e e s . R e s u l t s were d i s a p p o i n t i n g ; although the r e g r e s s i o n was h i g h l y s i g n i f i c a n t (t = 3.0497; tQ Q 5 = 2.021), i t accounted f o r c n l y 18% of the v a r i a t i o n i n the l i c h e n biomass. 46 TABLE I I I. Optical density values and red:green f i l t e r rat ios of trees with high and low l ichen biomass (x ± S.D.) Red Fi 1 ter Student 1s t Green F i1ter Student 1s t Blue Fi1ter A l l Trees High l ichen n = 34 3? = 3917 ± 1268 g Low l ichen n = 45 x = 910 ± 726 g 0.529 ± 0.144 0.537 ± 0.185 0.208 1 0.976 ±0.290 0.936 + 0.325 0.563 1 1.507 ±0.369 1.455 ±0.356 Student's t 0.642 1 Red:green rat io 0.559 ± 0.127 0.602± 0.163 Student's t 1.2881 Western hemlock High l ichen n = 28 x = 3996 ± 1260 g Low l ichen n = 15 x = 1084 ± 764 g 0.526 ± 0.129 0.502± 0.128 0.574 2 0.999 ± 0.231 1.089 ± 0.252 1.1902 1.527 ±0.310 0.532 ± 0.092 0.498 2 1.9062 1.576 ± 0.301 0.470 ± 0.118 Douglas-fir High l ichen n - 5 x = 3736 ± 1424 Low l ichen n = 10 x = 1379 ± 589.5 0.560 ±0.238 0.518 ±0.088 0.507 3 0.954 ±0.512 0.895 ± 0.192 0.330 3 1.508 ±0.626 0.615 + 0.116 0.302 3 0.182 3 1.444 ±0.203 0.601 ± 0.153 1 t 0 5 = 2.000 2 t 0 5 = 2.021 3 t 0 5 = 2.160 47 B e s u l t s of i n t e r p r e t e r s ' r a t i n g s of l i c h e n abundance on i n d i v i d u a l t r e e s are presented i n Table IV, Spearman's rho t e s t compares the i n t e r n a l ranking of p r e d i c t e d values with the i n t e r n a l ranking of the a c t u a l biomass v a l u e s . Thus, the i n t e r p r e t e r ' s a b i l i t y t o d i s t i n g u i s h t r e e s of high and low l i c h e n biomass w i t h i n a p l o t i s t e s t e d , but not h i s a b i l i t y to a s s i g n a b solute amounts of l i c h e n t o i n d i v i d u a l t r e e s . In a l l but 30 of the 33 cases (91S), l i c h e n p r e d i c t i o n s were p o s i t i v e l y c o r r e l a t e d with c a l c u l a t e d l i c h e n biomass, but i n o n l y 13 cases {39%) was the c o r r e l a t i o n s t a t i s t i c a l l y s i g n i f i c a n t at the 0.05 confidence l e v e l . The r e s u l t s c f the Hann-Whitney t e s t , used t o e v a l u a t e whether previous experience with photo i n t e r p r e t a t i o n or with l i c h e n ecology a f f e c t e d a b i l i t y t o judge l i c h e n abundance, were p o s i t i v e i n both cases. The e f f e c t of each f a c t o r was s t a t i s t i c a l l y s i g n i f i c a n t (photo i n t e r p r e t a t i o n : T = 155, T * o o 5 = 89; l i c h e n ecology : T = 218, T 0 0 5 = 83) . Of the two f a c t o r s , p r e v i o u s experience with l i c h e n ecology appeared to have the stronger i n f l u e n c e . The importance of a p r i o r i e xperience i s r e f l e c t e d i n the scores o f I n t e r p r e t e r s 3 and 7 - the two i n d i v i d u a l s who were most f a m i l i a r with the ecology of the study area. Success i n r a t i n g l i c h e n abundance on i n d i v i d u a l t r e e s i n a p l o t i s of l i t t l e p r a c t i c a l value unless one can a l s o r a t e the general l e v e l cf l i c h e n abundance on t h e p l o t , R e s u l t s of Spearman's rho t e s t on i n t e r p r e t e r s ' p r e d i c t i o n s of l i c h e n abundance on p l o t s were ne g a t i v e ; t h e r e was a weak negative c o r r e l a t i o n between p r e d i c t e d l e v e l of l i c h e n abundance and 48 TABLE IV. Spearman's rho values for predictions of l ichen abundance on indiv idual trees Interpreter 1 2 3 4 5 6 7 8 9 10 11 Photo Interpretation Experience L L L L L L H H H H H Knowledge of l ichen Ecology Plot 9 .20 .16 1 .59 .53 .45 1 .16 .53 . l l 1 .25 .34 .25 Plot 10 .38 .27 .55 .04 .70 .31 .85 -.03 .66 .39 .57 Plot 11 .20 -.07 .62 .44 .59 .33 .66 .59 .42 -.30 .71 Spearman's rho = 0.44; n = 15. Underlined values are s t a t i s t i c a l l y s i gn i f i cant . (p< 0.05) 1 n = 14; Spearman's rho = 0.46. 49 a c t u a l l e v e l of l i c h e n abundance {xho = -0.35). Apparently, some photo i n t e r p r e t e r s can compare l i c h e n abundance on t r e e s w i t h i n a p l o t with reasonable success, but they a r e unable t o d i s t i n g u i s h between areas with high l i c h e n abundance and areas with low l i c h e n abundance. P o s s i b l e e x p l a n a t i o n s f o r these r e s u l t s are d i s c u s s e d i n the f o l l o w i n g s e c t i o n . 50 D i s c u s s i o n Evaluation of methods The methods of determining lichen biomass that were developed i n t h i s study represent a compromise between the need to quantify biomass on a variety of s i t e s , and the desire f o r reasonable confidence i n the numbers obtained. The study was not designed in such a way that i t i s possible to put confidence 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 biomass t o t a l s can be assessed. The use of f e l l e d trees for epiphyte sampling has been c r i t i c i z e d by workers carrying out detailed research on epiphyte d i s t r i b u t i o n within a single tree (Pike et a l . 1972), but I do not believe i t caused any serious problems i n the present study. Despite some breakage of tranches and scattering of epiphytes, l i t t l e d i f f i c u l t y was experienced i n determining which of the f a l l e n lichen had come from the sample tree. Some error probably was caused by twigs knocked off adjacent trees of the same species by the f a l l i n g sample tree. This potential source of error i s a factor that should be taken into consideration when direction of f e l l i n g i s decided. Seme of the regressions r e l a t i n g Lichen Estimates to lichen biomass on branches did not have as high r 2 values as expected; those of the f i r s t two trees sampled were only 0.34 and 0.30. Fortunately, the r a t i o of sampled branches to unsampled branches was high (1:2 cr better), and the branches sampled tended to be 51 those with high l i c h e n abundance. T h e r e f o r e , the biomass c a l c u l a t i o n s f o r t r e e s were based to a c o n s i d e r a b l e extent on a c t u a l samples. Despite poor r 2 values i n a few cases, i t seems b e t t e r to e x t r a p o l a t e from r e g r e s s i o n s than t o assume t h a t unsampled branches are the same as sampled ones, as some workers have done. I b e l i e v e t h a t g u a n t i f i c a t i o n of l i c h e n biomass on branches could be improved by using a method analagous to the 3P ( p r o b a b i l i t y p r o p o r t i o n a t e to p r e d i c t i o n ) sampling scheme used by f o r e s t e r s to determine timber volume (Grosenbaugh 1967; Cochran 1963:251). T h i s method r e q u i r e s a p r e c i s e (but not n e c e s s a r i l y accurate) p r e d i c t i o n f o r each u n i t i n the p o p u l a t i o n , and measurement without e r r o r of a s m a l l sample from the p o p u l a t i o n . Using t h i s approach, the r e s e a r c h e r would p r e d i c t how many l i c h e n clumps of an a r b i t r a r y (but c o n s i s t e n t ) s i z e were present on each branch. Sample branches would be s e l e c t e d with a p r o b a b i l i t y p r o p o r t i o n a t e t o the p r e d i c t i o n of l i c h e n biomass. L i c h e n biomass on the sample branches would be weighed, and the r e s u l t s used t o determine a c o r r e c t i o n r a t i o of p r e d i c t e d biomass to measured biomass. An advantage to t h i s method i s t h a t i t i s easy to put confidence l i m i t s around the f i n a l biomass e s t i m a t e s , and i f the p r e d i c t i o n s are c o n s i s t e n t , the confidence l i m i t s are r e l a t i v e l y narrow. In the present study, 15 t r e e s on each p l o t were s e l e c t e d f o r v i s u a l estimates of l i c h e n abundance ac c o r d i n g to a random p r o p o r t i o n a t e sampling scheme. Trees were s e l e c t e d i n p r o p o r t i o n to t h e i r r e p r e s e n t a t i o n i n each species-dominance c l a s s . The scheme ensured that the species-dominance c l a s s e s having the 52 l a r g e s t numbers were most h e a v i l y sampled. But s i n c e as many as 9 species-dcminance c l a s s e s might be present w i t h i n a p l o t , a c l a s s was sometimes represented by only one or two t r e e s . V a r i a b i l i t y of l i c h e n abundance w i t h i n a c l a s s was sometimes great. Except i n f o r e s t s where species-dominance c l a s s e s are r e l a t i v e l y homogenous i n t h e i r l i c h e n abundance, estimates of l i c h e n biomass i n a stand probably c o u l d be improved by s t r a t i f y i n g t r e e s d i r e c t l y a c c o r d i n g to l i c h e n abundance, e.g. on a 1-5 scales Lichen abundance c l a s s e s would then be used as the s t r a t a f o r t r e e s e l e c t i o n , and as the b a s i s f o r e x t r a p o l a t i o n of l i c h e n biomass to the stand. The system of v i s u a l e stimates of l i c h e n abundance seems to be workable. The r 2 value of the r e g r e s s i o n equation of l i c h e n weights on v i s u a l e stimates was 0.75 f o r a l l species-dominance c l a s s e s . I t would have been d e s i r a b l e t o f e l l a second s e t of t r e e s to t e s t the p r e d i c t i v e value of the r e g r e s s i o n , but t h i s was impossible i n the present study because of l i m i t a t i o n s of time and money. In t e r - o b s e r v e r r e l i a b i l i t y seems to be w i t h i n an a c c e p t a b l e range, although i t should be c a r e f u l l y monitored i f more than one r e s e a r c h e r makes v i s u a l e stimates. The system has s e v e r a l l i m i t a t i o n s which should be noted. F i r s t , i t depends on the a b i l i t y of the r e s e a r c h e r to f i n d one or two p o s i t i o n s from which the e n t i r e crown of the t r e e , or at l e a s t most of i t , can be viewed. In very t a l l or very dense f o r e s t s , t h i s may not be p o s s i b l e . The present study was c a r r i e d out mainly on s i d e h i l l s i t e s ; I b e l i e v e t h a t some of the v a l l e y bottcm s i t e s w i t h i n the study area would have been d i f f i c u l t t o assess. 53 Second, the system was designed to discriminate among trees which vary greatly i n t h e i r lichen loads. Estimates of a l e c t o r i a (s.l.) cover for layers of trees encountered i n the study area ranged from 0 to 80%. In i t s present form, the system would not be appropriate f o r making f i n e r d i s t i n c t i o n s as, for example, discriminating among trees i n young stands i n which Ale c t o r i a -cover never exceeded 5%. Third, i t must not be assumed that the regressions used in the present study to extrapolate from sampled trees to unsampled trees are general i n the i r a p p l i c a b i l i t y . I f lichen biomass i s to be assessed i n other areas, further biomass sampling should be undertaken i n those forest types to determine relationships between v i s u a l estimates and lichen biomass. Results of the densitometry indicate some po t e n t i a l for detection of lichen abundance using red:green f i l t e r r a t i o s . Use of r a t i o s rather than absolute density values eliminates much of the variation i n optic a l density readings caused by d i f f e r e n t i a l l i g h t i n g cf trees. I t i s l i k e l y that a larger sample siz e would show that red:green r a t i o s of trees with high lichen biomass d i f f e r s i g n i f i c a n t l y from those of trees with low lich e n biomass, at least for some species. However, the present r 2 value of the regression ( r 2 = 0.18) suggests that the predictive value of the r a t i o i s low. Photo interpretation r e s u l t s were disappointing. The i n a b i l i t y of interpreters to distinguish areas of high lichen abundance from areas of low lichen abundance suggests that the potential usefulness of a i r photography as a lichen inventory tool i s low. Several factors may explain these poor r e s u l t s . 54 F i r s t , the presence of B r y o r i a mixed with A l e c t o r i a high i n the crowns of t r e e s may have d e t r a c t e d from an i n t e r p r e t e r ' s a n i l i t y to d e t e c t abundance o f A l e c t o r i o i d l i c h e n s i n g e n e r a l . B r y o r i a spp. are dark i n c o l o u r and may have caused underestimates of l i c h e n abundance where they o c c u r r e d . However, I b e l i e v e t h a t two other f a c t o r s were more important., S e v e r a l i n t e r p r e t e r s remarked that they were confused by d i f f e r e n c e s i n photo s c a l e . I had o r i g i n a l l y planned f o r t h r e e photo s c a l e s : 1:2000, 1:4000, and 1:8000. But t h e d i f f i c u l t i e s of f l y i n g at a constant height above ground l e v e l over mountainous t e r r a i n were such t h a t the photographs taken at the two lower f l y i n g h e ights were v a r i a b l e i n s c a l e . , F o r t h i s reason, the photographs assessed by i n t e r p r e t e r s a l s o v a r i e d i n s c a l e . A r b o r e a l l i c h e n s tended t o be more n o t i c e a b l e i n the l a r g e s c a l e photographs, r e g a r d l e s s of a c t u a l abundance, and t h i s may account f o r i n t e r p r e t e r s ' i n a b i l i t y to d i s t i n g u i s h among s i t e s . ; The t h i r d f a c t o r t h a t may have c o n t r i b u t e d t o the poor r e s u l t s i s the growth h a b i t of the l i c h e n s . A l e c t p r i a sarmentosa and most a r b o r e a l s p e c i e s of B r y o r i a have a pendent growth h a b i t ; they tend t o form elongated clumps which hang from branches and twigs. They may be v i s i b l e from d i r e c t l y above, but the degree t o which they are v i s i b l e from above may not be a good r e p r e s e n t a t i o n of t h e i r a c t u a l biomass. One i n t e r p r e t e r remarked t h a t l i c h e n abundance was e a s i e r to assess on t r e e s near the edge of the photograph because they were seen o b l i g u e l y . In the study area, I have observed t h a t areas of high and low l i c h e n abundance can be d i s t i n g u i s h e d with b i n o c u l a r s 55 from a viewpoint a c r o s s a v a l l e y . These o b s e r v a t i o n s suggest t h a t o b l i q u e a e r i a l photography might be u s e f u l i n i n v e n t o r y i n g l i c h e n abundance, at l e a s t at the reconnaissance stage. Biomass r e s u l t s compared with l i t e r a t u r e r e p o r t s The l i c h e n biomass t o t a l s obtained i n t h i s study are compared with r e s u l t s o b t a i n e d i n other north temperate and b o r e a l ecosystems i n Table V. The f i g u r e s are not s t r i c t l y comparable, as some are based on a l l l i c h e n s and others i n c l u d e only A l e c t o r i o i d l i c h e n s , but they provide an i d e a of the range i n magnitude of l i c h e n p r o d u c t i o n that occurs. The lowest biomass t o t a l i n t h i s study (21 kg/ha) i s s m a l l e r than any other t o t a l s r e p o r t e d , but the highest (1528 kg/ha) i s g r e a t e r than most other r e p o r t e d values. I t i s only about h a l f as great as the ttaximum reported value - 3289 kg/ha, r e p o r t e d f o r an Englemann s p r u c e - s u b a l p i n e f i r f o r e s t i n the i n t e r i o r of B r i t i s h Columbia, TABLE V. Comparison of reported biomass tota ls of epiphytic lichens Source Area Forest Type Epiphytes Oven-dried weight (kg/ha) Edwards et a l . 1960 Wells Gray Park, B.C. Scotter 1962 Rudnova, Tonkonogov, & Dorokhova 1964 (c ited in Rodin & Bazi lev ich 1967) B.iazrov 1969 W.D. Boehm 1972 unpublished data Pike et a l . 1972 Black Lake, northern Saskatchewan Archangel Province, U.S.S.R. U.S.S.R. Ross Lake, Washington Mi xed Pinus Picea Abies Picea Abies Picea Pinus conifers contorta engelmannii-lasiocarpa engelmannii-lasiocarpa mariana banksiana Alectoria sarmentosa and Bryoria spp. Oregon Pinus excelsa Picea abies-mixed deciduous Pinus contorta Pseudotsuga menziesii Tsuga heterophylla Pseudotsuga menziesii A l l l i chens 1 Bryopogon implexus3 A l l lichens Alectoria sarmentosa Bryoria spp. A l l lichens 1* 837 282 754 3289 rl 198 2 2051 2 700 600 282 47-233 527 564 Ln ON TABLE V. (Continued) Oven-dried Source Area Forest Type Epiphytes weiqht (kg/ha) Schroeder 1974 Selk irk Mountains, B.C. & Washington Picea engelmannii-Abies lasiocarpa 430 Picea engelmannii-Abies lasiocarpa A l l macrolichens 5 103 Larix occidentalis 429 Andre et a l . 1975 France Abies alba A l l l ichens 1040 Wein & Speer 1975 Cape Breton Is land, Nova Scotia Abies balsamea-Picea mariana A l l l ichens 47-280 Turner & Singer 1976 Western Washington Abies amabilis-Tsuga mertensiana A l l l ichens 1900 This study Northern Vancouver Island Tsuga heterophylla-Pseudotsuga menziesii-Thuja plicata Alectoria sarmentosa and Bryoria spp. 21-1528 1 Pr imari ly Bryoria spp ., Evernia mesomorpha. Usnea hirta. 2 A i r -d r ied weight. 3 Probably Bryoria implexa. 4 Only epiphytes on overstory Douglas-f ir included. 5 Pr imari ly Alectoria sarmentosa and Bryoria spp. 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 abundance 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 v e g e t a t i v e c h a r a c t e r i s t i c s of s i t e s . The r a t i o n a l e f o r t h i s p o r t i o n of the study was that the i n f o r m a t i o n gathered would improve understanding of the ecology of A l e c t o r i o i d l i c h e n s , and would a s s i s t i n i d e n t i f y i n g f o r e s t areas with abundant l i c h e n . I expected t h a t high l i c h e n abundance would be c o r r e l a t e d with measures which are themselves a s s o c i a t e d with high l e v e l s of s u n l i g h t reaching the t r e e canopy. High l i c h e n abundance should be a s s o c i a t e d with south aspects and steep s l o p e s because these f a c t o r s i n c r e a s e the p o t e n t i a l amount of s o l a r r a d i a t i o n i n c i d e n t on a s u r f a c e at a given l a t i t u d e . L i c h e n abundance should be c o r r e l a t e d n e g a t i v e l y with crown c l o s u r e (except at low l e v e l s of crown c l o s u r e , where s u b s t r a t e becomes l i m i t i n g ) , because crown c l o s u r e l i m i t s the amount of l i g h t p e n e t r a t i n g the t r e e canopy. I a l s o expected that high l i c h e n abundance would be c o r r e l a t e d p o s i t i v e l y with the r a t i o of l i v e crown to t r e e height, a measure which has been proposed as an i n d i c a t o r of crown c o m p e t i t i o n . The r a t i o tends t o be high i n open-grown stands and low i n dense stands, where the lower branches r e c e i v e inadequate l i g h t and die (Hard 1964, Chiam 1967). I a l s o expected that l i c h e n abundance would i n c r e a s e with i n c r e a s i n g e l e v a t i o n . T h i s trend had a l r e a d y been noted i n the study area (Bochelle 1978) and an adjacent watershed ( B a r i c h e l l o 1975), and i s probably a s s o c i a t e d with i n c r e a s i n g humidity. 59 although crown closure also tends to be more favourable for 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 collected at the fourteen plots on which lichen abundance was estimated. A complete f l o r i s t i c l i s t of vascular plants and major bryophyte species of the forest f l o o r was made at each plot. A percent cover value was estimated f o r each species, and each species except those i n the tree layer was assiened a d i s t r i b u t i o n class. Distribution classes were:1 1. rare in d i v i d u a l 2. few scattered indi v i d u a l s 3. single patch 4. several scattered individuals 5. a few small patches 6. several well-spaced patches 7. continuous cover of well-spaced indiv i d u a l s 8. continuous dense cover with a few openings 9. continuous dense cover uninterrupted Vegetation was c l a s s i f i e d using the method of tabular comparison developed by the Braun-Blanquet School of Phytosccioloqy (Mueller-Dcmbois and Ellenberg 1974: 177-193);. Topographic measurements taken were elevation, measured with an altimeter; slope, measured with a clinometer; and aspect, measured with a compass. For computer analysis, aspect 1 Developed by the Vegetation Functional Subcommittee of the B r i t i s h Columbia Land Resources Committee, Environment and Land Use' Committee, Ministry of Environment, B r i t i s h Columbia. Presently the Resource Analysis Branch, Ministry of Environment, B r i t i s h Columbia. 61 readings were converted to departure i n degrees from due south. Measurements of slope and aspect together with s o l a r r a d i a t i o n t a b l e s (Buffo et a l . 1972) were used to determine annual p o t e n t i a l s o l a r r a d i a t i o n i n c i d e n t on each s i t e except P l o t 12. The t a b l e s are not a p p l i c a b l e to P l o t 12 because i t i s l o c a t e d i n a narrrow v a l l e y i n which day l e n g t h i s r e s t r i c t e d by topography (Buffo e t a l . 1972). Crown c l o s u r e was measured u s i n g the "mocsehorn" technigue (Garrison 1949) and a s p h e r i c a l densiometer (Lemmon 1956).Mean crown c l o s u r e values were c a l c u l a t e d from 16 readings l o c a t e d s y s t e m a t i c a l l y w i t h i n each p l o t . Crown c l o s u r e of the main t r e e canopy was a l s o estimated v i s u a l l y . Tree i n v e n t o r y data (see Chapter 3) were used to c a l c u l a t e b a s a l area. A l l l i v i n g t r e e s over 18 cm i n diameter were i n c l u d e d , as were a l l dead p o t e n t i a l (merchantable) t r e e s 18 cm or more i n diameter and 3 m or more i n height. Down t r e e s were in c l u d e d i f they met the above c r i t e r i a ( B r i t i s h Columbia F o r e s t S e r v i c e 1973). The height of the codcminant t r e e l a y e r was estimated, based on the measured heights of the v i s u a l estimate t r e e s . The mean r a t i o of crown l e n g t h to t r e e height (CL/TH) was c a l c u l a t e d , based on measurements of v i s u a l e s timate t r e e s (see Chapter 3). The v a r i a b l e s d e s c r i b e d above were r e l a t e d to a v a r i a b l e r e p r e s e n t i n g l i c h e n abundance: mean percent a l e c t o r i a ( s . l . ) per l a y e r on Tsuga s p e c i e s . 1 T h i s v a r i a b l e was chosen as a 1 Hereafter r e f e r r e d to as "percent A l e c t o r i a " 62 measure of the q u a l i t y of a s i t e f o r l i c h e n growth, r e g a r d l e s s of the s i z e or number of t r e e s on the s i t e . Hemlocks were used r a t h e r than a l l t r e e s p e c i e s because l i c h e n biomass on Tsuga s p e c i e s was almost i n v a r i a b l y g r e a t e r than on other s p e c i e s , suggesting t h a t on other s p e c i e s s u b s t r a t e may be more important than s i t e as a l i m i t i n g f a c t o r . B e s u l t s of the v e g e t a t i o n a n a l y s i s were a l s o r e l a t e d t o t o t a l A l e c t o r i a ( s . l . ) biomass. 1 The height of the codcminant t r e e l a y e r was r e l a t e d t o e l e v a t i o n as w e l l as t o percent Ale c t o r i a , . R e l a t i o n s h i p s amonq v a r i a b l e s were t e s t e d u s i n g the l i n e a r r e q r e s s i o n model of the computer proqram MIDAS o f the S t a t i s t i c a l Research Laboratory, U n i v e r s i t y of Michigan. 1 H e r e a f t e r r e f e r r e d to as " A l e c t o r i a biomass" 63 Besults Table VI summarizes the data used in the analysis of lichen abundance in r e l a t i o n to s i t e c h a r a c t e r i s t i c s . Vegetation communities The vegetation c l a s s i f i c a t i o n resulted i n the delineation of three communities (Appendix I I ) . Because my data base was i n s u f f i c i e n t for a comprehensive c l a s s i f i c a t i o n of the vegetation i n the study area, the following comments are based on f i e l d impressions and published 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 well as on data co l l e c t e d at plots. Plots 1, 3, and 10 formed a group which was l a b e l l e d the Douglas-fir/Salal Community (DF/S). The c h a r a c t e r i s t i c combination of species for t h i s community i s Pseudotsuqa menzijasii^ Gaultheria shallon, Hemitomes congestum, Stokesie11a oreganaA V i o l ^ sempervirens„ and Boschniakia hookeri. The community develops on xeric s i t e s , most frequently on south-facing slopes, ridges, and knolls. In the study area, i t may occur from the valley bottom to about 700 m, although at high elevations i t i s limited to very xeric s i t e s . The Douglas-fir/Salal Community i s similar to the Hyloccmio (splendentis) - Eurhynchio (oregani) - Gaultherio (shallonis) -Pseudotsugetum menziesii (the Gaultheria shalIon association) of Kojima and Krajina (1975) and to the cjrthic Gaultheria forest type of O r l o c i (1961). Kojima and Krajina (1975) described t h e i r Gaulthjgria s h a l l on association as occurring on x e r i c s i t e s with extremely well-drained and shallow s o i l s , and a r e l a t i v e l y high 64 TABLE VI. Site character i s t ics of plots 7 . Potential Crown Closure {%) Crown ; Height R , Plot A^ectorza P e r c e n t Elevation . . Slope Annual Length/ of Co- j ? " * 1 Vegetation Number 7, 0 n?u S S, Alectoria (m) aspect ( D e g r e e s ) Radiation Densio- Moose- ... , , Tree Dominant L 2 / , , Community < k9/ h a) (cal/cm 2/year) meter horn V i s u a l Height Tree's (m) ( m / h a ) 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 UNCLASSIFIED 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 65 slope gradient. Crown closure was r e l a t i v e l y low (x = 65%) and l i g h t i n tensity under the tree canopy the highest (x = 3239 lux) reported in their study. Forest productivity of the Gaultheria shallcn association i s low; the s i t e index for Douglas-fir was reported as 33 m/100 years and for western hemlock, 26 m/1-,00 years (Kojima and Krajina 1975). Plots 5, 8, 14, and 151 formed a group which was l a b e l l e d the Amabilis Fir/Oval-leaved Huckleberry Community (AF/OH).The c h a r a c t e r i s t i c combination of species for t h i s community includes Abies amabilis, Ccrnus canadensisRubu-s pedatus^ vaccinium ovalifolium, Menziesia ferruqjnea, L i s t e r a caurina, Clintenia u n i f l o r a , Streptopus roseus, Plagiothecium undulatorn, and Tsucja mertensiana. The community develops on mesic s i t e s , most frequently on gentle or moderate north-, east-, or west-facing slopes above 500 m i n elevation. The Amabilis Fir/Oval-leaved Huckleberry Community i s comparable to the Rhytidiadelpho (lorei) - Plagiothecio (undulati) - Rubo (pedati) - Vaccipio (alaskaensis) - Abieto (amabilis) - Tsuqetum heterophyllae (the Vaccinium alaskaense association) of Kojima and Krajina (1975), and the clintonicsum variant of the Abieto - Tsuqetum heterophyllae of O r l o c i (1961). The Vaccinium alaskaense association (Kojima and Krajina 1975) occurs on mesic-subhygric s i t e s i n the wetter subzone of the Coastal Western Hemlock Zone (Krajina 1965). It i s c h a r a c t e r i s t i c of gentle slopes and of terraces near the bottom of valleys, and i t s d i s t r i b u t i o n appears to be related to the 1 No lichen estimates were performed on Plot 15. 66 d i s t r i b u t i o n of snow-pack. Crown c l o s u r e was high (x = 80%) i n the Vaccinium alaskaense a s s o c i a t i o n , and l i g h t i n t e n s i t y under the canopy was r a t h e r low (x = 1943 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 f o r western hemlock was 40 m/100 years and f o r a m a b i l i s f i r , 37 m/100 years (Kojima and K r a j i n a 1975). P l o t s 6, 9, 11, and 13 formed a t h i r d group which i n c l u d e d s p e c i e s c h a r a c t e r i s t i c o f both the p o u g l a s - f i r / S a l a l Community and the A m a b i l i s F i r / O v a l - l e a v e d Huckleberry Community. This group, termed the T r a n s i t i o n Community (THANS), occurs on s i t e s t h a t are i n t e r m e d i a t e between t y p i c a l D o u g l a s - f i r / S a l a l s i t e s and Amabilis F i r / O v a l - l e a v e d Huckleberry s i t e s . P l o t 6 i s on a s u b x e r i c , s o u t h e a s t - f a c i n g s l o p e at an e l e v a t i o n (669 m a . s . l . ) that i s too high f o r the pure D o u g l a s - f i r / S a l a l Community except on very x e r i c s i t e s (e.g. P l o t 1). P l o t 11 i s on a x e r i c s i t e , but at an e l e v a t i o n (792 m a . s . l . ) o u t s i d e the range of the D o u g l a s - f i r / S a l a l Community . P l o t s 9 and 13 are on w e s t - f a c i n g s l o p e s t h a t are steep and w e l l - d r a i n e d . Four p l o t s c o u l d not be c l a s s i f i e d u s i n g the Braun-Blanguet approach, probably because the sample s i z e was too s m a l l . Of these f o u r , two p l o t s (2 and 7) are s t r u c t u r a l l y and f l o r i s t i c a l l y s i m i l a r t o the hyloccmiosum s p l e n d e n t i s of the Hyocomio ( s p l e n d e n t i s ) - Eurhynchio (oregani) - Mahonio (nervosae) - Pseudotsugo - Tsugetum h e t e r o p h y l l a e (the Hyloconiium splendens v a r i a n t of the moss a s s o c i a t i o n ) d e s c r i b e d by Kojima and K r a j i n a (1975). T h i s v a r i a n t was d e s c r i b e d as o c c u r r i n g i n the d r i e r subzone of the C o a s t a l Western Hemlock Zone {Krajina 1965), on s e l l - d r a i n e d mesic s i t e s where s l o p e s 67 are moderate and s o i l s r e l a t i v e l y deep. Crown closure was high (x = 85%) and l i g h t intensity under the forest canopy low (x = 1750 lux). Productivity was intermediate; the s i t e index was 40 m/100 years for Douglas-fir and 37 m/100 years for western hemlock. The Hyloccmium splendens variant of the moss association has Pseudotsuga menziesii f Tsuga heterophylla, and Thuja p l i c a t a i n the tree canopy, low cover i n the shrub and herb layers, and a well-developed mess layer including HyjLocomium splendens, Rhytidiadelphus l o r e u s A and Stokesiella 2Iia2Bli Because Plots 2 and 7 occurred on mesic, moderately sloped, low elevation s i t e s and were s t r u c t u r a l l y and f l o r i s t i c a l l y s i m i l a r to the Hyloccmium s p l e n d e n s s i t e s described by Kojima and Krajina (1S75), they were given the tentative label of "Moss Community" {BOSS) . The two remaining plots were not c l a s s i f i e d . I had expected that Plot 4 would belong with the Transition Community, but the results of the analysis dc not support c l a s s i f y i n g i t with that group. The plot has few species c h a r a c t e r i s t i c of the Douglas-f i r / S a l a l , Aoabilis Fir/oval-leaved Huckleberry, or Transition Communities, but d i f f e r s from Moss s i t e s i n i t s high cover (40%) of Vaccinium alaskaense . Plot 12 d i f f e r s from a l l other plots i n i t s vegetation and i t s topographic position. I t has very low cover and species d i v e r s i t y i n a l l layers beneath the tree canopy. The plot i s located on a steep south-facing slope near the bottom of a narrow east-west valley, where i n s o l a t i o n i s re s t r i c t e d by topography. More data would be needed to c l a s s i f y Plots 4 and 12. 68 The relationship between vegetation community and lichen abundance i s shown i n Figure 9. The vegetation communities which appear to have the greatest potential for lichen production are the Douglas-fir/Salal Community and the Transition Community, regardless of whether Alectoria biomass (Fig, 9A) or percent Alectoria (Fig. 9E) i s measured, within these two communities there i s a strong tendency f o r lichen abundance to increase with elevation. Plot U, the u n c l a s s i f i e d plot which I expected to group with the Transition Community, i s also r e l a t i v e l y high i n lichen abundance by both measures. The Amabilis Fir/Oval-leaved Community i s low in- Alectoria biomass, but i n two of the three plots percent A l e c t o r i a i s moderate rather than low, These results suggest that some s i t e s of t h i s type are moderate in s u i t a b i l i t y for lichen production, but that amount of available substrate i s lew. The Moss Community i s low in l i c h e n production by both measures. Although only two s i t e s were sampled, f i e l d impressions support the conclusion 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 physical environment Scattergrams of percent Alectoria on slope, aspect, potential annual radiation, and elevation on percent Alectoria are given in Figures 10 to 13. Variation i n slope (Fig. 10} accounts for very l i t t l e of the variation i n percent Alectoria;• the regression has an r 2 value cf only 0.18 and 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 . 1500H cn 1000" CO < 50(H ® n PLOT 10 3 1 DF/S 13 9 6 11 TRANS 5 14 8 AF/OH 7 2 MOSS 3 0 H 20H 10H i PLOT 10 3 1 DF/S 13 9 6 11 TRANS 5 14 8 AF/OH 7 2 MOSS 12 4 U I I 12 4 U Figure 9. Vegetation community and Aleetoria biomass (kg/ha) (A); vegetation community and percent Alectoria (B). Within communities, plots are arranged in order of increasing e levat ion. U = unc la s s i f i ed . 40-< 3 0 H rr O i— o UJ < 20-F-z LU O DC LU 0- 10-v O 10 A V 20 30 S L O P E degrees I DF/S community | TRANS community AF/OH community | MOSS community unc las s i f ied I _ T _ 40 I 50 Figure 10. Relationship between slope and percent Alectoria (y = 7.6012 + 0.33442; n = 14; S V . Y = 9.75; r 2 = 0.18). 71 40-1 30H o o LU 20H LU O CC LU V 10-_Q_ 30 60 90 120 150 ASPECT (degrees from south) "3o~ 180 • DF/S community A TRANS community v AF/OH community B MOSS community o unc las s i f ied Figure 11. Relationship between aspect and percent Alectoria (y = 23.376 - 0.120 x; n = 14; S y . x = 9.31; r 2 = 0.25) 72 ( 40 -I 16 18 P O T E N T I A L A N N U A L R A D I A T I O N 10,000 x c a l / c m 2 / y r r 20 22 © DF/S community A TRANS community v AF/OH community a MOSS community o unclassified Figure 12. Linear regression of percent Alectoria on potential annual radiation (y = -19.081 + 0.00197 x; n. = 13; S y . x = 8.56; r 2 = 0.37). 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 unc las s i f ied Figure 13. Linear regression of percent Alectoria on elevation (y = -15.465 + 0.0527 x; n = 14; S y . x = 8.14; r 2 = 0.47). 74 The r 2 value of percent A l e c t o r i a on aspect, measured i n degrees frcm due south, i s 0.25, s l i g h t l y higher than that, c f 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 s i g n i f i c a n t {Fig. 11). The t r i a n g u l a r shape c f the scattergram suggests t h a t s o u t h e r l y aspect i s a necessary but not s u f f i c i e n t c o n d i t i o n f o r high l i c h e n abundance. P o t e n t i a l f o r high percent alectoria appears to i n c r e a s e as aspect approaches due south, but the p o t e n t i a l i s not always r e a l i z e d . D i f f e r e n c e s i n v e g e t a t i o n community do not appear to e x p l a i n the low l e v e l s of l i c h e n abundance on some p l o t s , although l i c h e n g u a n t i t i e s cn both Moss s i t e s are lower than expected on the b a s i s of aspect.. The scattergram i l l u s t r a t e s the occurrence o f D o u g l a s - f i r / S a l a l s i t e s cn south aspects, 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 s on north aspects — the aberrant s i t e has a slope of only 5 degrees — and T r a n s i t i o n s i t e s on i n t e r m e d i a t e a s p e c t s . P o t e n t i a l annual s o l a r r a d i a t i o n ( F i g . 12) e x p l a i n s 37% of the v a r i a t i o n i n percent A l e c t o r i a . The r e g r e s s i o n i n d i c a t e s a s t a t i s t i c a l l y s i g n i f i c a n t dependence of percent A l e c t o r i a on s o l a r r a d i a t i o n ( t = 2. 522, t 0 O 5 =2.201). Like F i g u r e 11, t h i s scattergram has a t r i a n g u l a r shape which suggests t h a t s o l a r r a d i a t i o n s e t s an upper l i m i t on l i c h e n p r o d u c t i o n . Most D o u g l a s - F i r / S a l a l and T r a n s i t i o n s i t e s have amounts of A l e c t o r i a higher than that p r e d i c t e d by the r e g r e s s i o n . Percent A l e c t c r i a on Amabilis F i r / O v a l - l e a v e d Huckleberry s i t e s i s c l o s e to p r e d i c t e d v a l u e s , whereas t h a t cn Moss s i t e s i s lower. The s i n g l e environmental f a c t o r which accounts f o r the g r e a t e s t v a r i a t i o n i n percent A l e c t o r i a i s e l e v a t i o n ( F i g . , 13), which has an r 2 value of 0.47. The dependence of percent 75 A l e c t o r i a cn e l e v a t i o n i s h i g h l y s i g n i f i c a n t (t = 3.267, t a o y = 3.055). Much of the v a r i a t i o n around the r e g r e s s i o n l i n e i s e x p l a i n a b l e by d i f f e r e n c e s i n v e g e t a t i o n community. Douglas-f i r / S a l a l s i t e s are c o n s i s t e n t l y higher i n percent A l e c t o r i a than p r e d i c t e d by the r e g r e s s i o n ; Amabilis F i r / O v a l - l e a v e d Huckleberry s i t e s are c o n s i s t e n t l y lower. Amounts of A l e c t o r i a on T r a n s i t i o n s i t e s are c l o s e to p r e d i c t e d v a l u e s . Environmental v a r i a b l e s i n combination e x p l a i n more of the v a r i a t i o n i n percent 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 m u l t i p l e r e g r e s s i o n y = -38.639 • 0.043 (x ( ) + 0.00016658 mz) (6) (n = 13; S Y . / = 5. 99) where y i s percent A l e c t o r i a , x, i s e l e v a t i o n i n meters, and xz i s p o t e n t i a l s o l a r r a d i a t i o n i n c al/cm 2/year, has an r 2 c f 0.72. The r e g r e s s i o n y = -14.043 + 0.0455 (x () - 0.0903 (x 2) * 0.3715 (x 3 ) (7) (n = 13; S y. x = 5.06) where y i s percent 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 2 i s departure from due south i n degrees, and x 3 i s s l o p e i n degrees, has an r 2 of 0.82. F o r e s t measures Scattergrams and l i n e a r r e g r e s s i o n s of crown c l o s u r e measured by densiometer, moosehorn, and v i s u a l e s t i m a t e on percent A l e c t o r i a are given i n F i g u r e 14, Only the r e g r e s s i o n of percent A l e c t o r i a on densiometer readings i s s t a t i s t i c a l l y s i g n i f i c a n t {t = 3.3174, t 0 0 5 = 3.055). Crown c l o s u r e as measured by the densiometer accounts f o r 48% of the v a r i a t i o n i n 40-i < DC O 3 0 I-o LU _1 <20 - | h -z LU o LX10' LU CL 70 "80" 90 100 CROWN C L O S U R E Densiometer ® « C L 40-1 301 20H 10H 1 40-1 40 © • ® v A 50 60 70 80 CROWN C L O S U R E Moosehorn — i 90 30-20-1<H A O © v © -9-50 60 70 80 CROWN C L O S U R E Visual - i 90 DF/S community A TRANS v AF/OH a MOSS o Unclass i f ied Figure 14. Relationship between crown closure and percent Alectoria: densiometer (y = 140.7 - 1.473 x; n = 14; S y . x = 8.08 r 2 = 0.48) (A); moosehorn (y = 42.859 - 0.375 x; n = 14; S y . x = 9.95 r 2 = 0.21) (B); v isual estimates (y = 27.382 - 0.19005 x; n = 14; S y . x = 11.01 r 2 = 0.03) (C) 77 percent a l e c t o r i a . The r e g r e s s i o n of b a s a l area on percent A l e c t o r i a 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 {Fig. 15), The height of the codcniinant t r e e l a y e r ( F i g , 16) i s a good p r e d i c t o r of percent A l e c t o r i a ( r 2 = 0.64). Tree height i s a l s o s i g n i f i c a n t l y c o r r e l a t e d with e l e v a t i o n (r = -.6180, r 0 0 5 = 0,5324). The r e g r e s s i o n of crown l e n g t h / t r e e height i n d i c a t e s l i t t l e r e l a t i o n s h i p ( r 2 = 0.18), except 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 s i t e s and T r a n s i t i o n s i t e s ( F i g . 17). I suspected t h a t the ccor r e l a t i o n s h i p might be due to the d i f f e r e n t combination of shade-t o l e r a n t and s h a d e - i n t o l e r a n t t r e e s on each s i t e , so I repeated the r e g r e s s i o n u s i n g crown l e n g t h / t r e e height r a t i o s of western hemlock and D o u g l a s - f i r s e p a r a t e l y . In each ca s e , the r 2 of the r e g r e s s i o n was f u r t h e r reduced. < 3 0 -o r— O UJ _ l < 2 0 -\-LU O DC LU ° - 1 0 -8 0 9 0 1 0 0 BASAL AREA m 2 /ha 1 1 0 1 2 0 . © DF/S community A TRANS community v AF/OH community B MOSS community o unc lass i f ied Figure 15. Relationship between basal area and percent Alectoria (y = 36.907 - 0.235 x; ri = 14; S y . x = 10.71) r2 = 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 unc las s i f ied 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— z LU O DC LU O 10 H _ — , [—^ — r .65 .70 .75 C R O W N L E N G T H / T R E E H E I G H T o DF/S community A TRANS community v AF/OH community a MOSS community o unc las s i f ied .80 Figure 17. Relationship between crown length/tree height and percent Alectoria (y = -49.68 + 0.93 x; n = 14; S y . x = 10.14; r 2 = 0.18). 81 D i s c u s s i o n T e s t s of r e l a t i o n s h i p s between physical.environment v a r i a b l e s and percent a l e c t o r i a support the hypothesis that s o l a r r a d i a t i o n and e l e v a t i o n are major determinants of the q u a l i t y o f s i t e s i n the study area f o r l i c h e n p r o d u c t i o n . The r e g r e s s i o n of percent a l e c t o r i a on each of these v a r i a b l e s i s s t a t i s t i c a l l y s i g n i f i c a n t . The f a c t t h a t separate r e g r e s s i o n s of s l o p e and aspect are not s i g n i f i c a n t i s not s u r p r i s i n g , as the e f f e c t of each v a r i a b l e on s o l a r r a d i a t i o n depends on the other. The high r 2 (0.82) of slope and aspect considered with e l e v a t i o n i n d i c a t e s the importance of these v a r i a b l e s . The f a c t t h a t the r e g r e s s i o n of e l e v a t i o n , s l o p e , and aspect (Equation 7) accounts f o r more of the v a r i a b i l i t y i n percent A l e c t o r i a than t h a t of e l e v a t i o n and s o l a r r a d i a t i o n {Eguation 6) suggests t h a t s l o p e and aspect operate i n a more complex manner than simply determining p o t e n t i a l amounts o f i n c i d e n t s o l a r r a d i a t i o n on a s i t e . For i n s t a n c e , slope i s probably important i n determining how much l i c h e n s u b s t r a t e w i t h i n the t r e e canopy at a s i t e i s exposed to s u n l i g h t , as well as how much s u n l i g h t reaches the s i t e . S o l a r r a d i a t i o n appears to s e t an upper l i m i t on p o t e n t i a l f o r l i c h e n p r o d u c t i v i t y . At lower e l e v a t i o n s , l i c h e n abundance tends t o be lower than p r e d i c t e d by s o l a r r a d i a t i o n , probably because of decreased moisture, v e g e t a t i o n community i n t e g r a t e s the e f f e c t s of s l o p e , a s p e c t , e l e v a t i o n , and other environmental v a r i a b l e s on a s i t e - s p e c i f i c l e v e l . The s i t e r s p e c i f i c i t y of v e g e t a t i o n may enhance i t s p r e d i c t i v e value. For example, P l o t 82 10 i s g e o g r a p h i c a l l y c l o s e t o P l o t 2 and has only s l i g h t l y acre p o t e n t i a l annual r a d i a t i o n , but has much high e r l i c h e n abundance. P l o t 10 i s l o c a t e d on a convex s l o p e and has thus developed i n t o a D o u g l a s - f i r / S a l a l s i t e ; P l o t 2, a Boss s i t e , i s l o c a t e d i n an adjacent draw and probably r e c e i v e s l e s s s o l a r r a d i a t i o n than expected on the b a s i s of i t s gross p o s i t i o n . In ge n e r a l , D o u g l a s - f i r / S a l a l s i t e s and T r a n s i t i o n s i t e s appear to have the best p o t e n t i a l f o r l i c h e n p r o d u c t i v i t y , e s p e c i a l l y at higher e l e v a t i o n s . T h i s i s probably because they develop where i n c i d e n t s o l a r r a d i a t i o n i s high. Conversely, Moss s i t e s develop where s o l a r r a d i a t i o n i s r e l a t i v e l y low, and probably never have high l i c h e n abundance. l i k e v e g e t a t i o n community, f o r e s t measures i n t e g r a t e the e f f e c t s of many environmental v a r i a b l e s . Tests of r e l a t i o n s h i p s between f o r e s t measures and percent A l e c t o r i a p r o d u c e d mixed r e s u l t s . The r e l a t i o n s h i p s between crown c l o s u r e measures and percent A l e c t o r i a are p u z z l i n g . A f t e r c a r r y i n g out the f i e l d w o r k , I expected t h a t v i s u a l estimates of crown c l o s u r e would be more c l o s e l y r e l a t e d t o l i c h e n abundance than e i t h e r densicmeter or moosehorn readings because cover of understory t r e e s was i n c l u d e d i n the instrument readings. The poor r e l a t i o n s h i p between v i s u a l estimates of crown c l o s u r e and l i c h e n abundance may have been due to the d i f f i c u l t y of e s t i m a t i n g cover of the main canopy without c o n s i d e r i n g understory t r e e s . The apparent r e l a t i o n s h i p between crown c l o s u r e as measured by the densiometer and percent A l e c t o r i a i s d i f f i c u l t to e x p l a i n and should be i n t e r p r e t e d c a u t i o u s l y , c o n s i d e r i n g the poor r e s u l t s o f the other crown c l o s u r e 83 measures. The poor r e l a t i o n s h i p cf the r a t i o of crown length to t r e e h e i g h t with percent A l e c t o r i a was s u r p r i s i n g . I expected the r a t i o to be a good i n d i c a t o r of the amount of s u n l i g h t that penetrates the t r e e canopy, i n t e g r a t i n g the e f f e c t s of i n c i d e n t s o l a r r a d a t i o n and canopy s t r u c t u r e . C o n s i d e r i n g t r e e s p e c i e s s e p a r a t e l y , I expected the r e l a t i o n s h i p t o hold b e t t e r f o r D o u g l a s - f i r than f o r western hemlock, because D o u g l a s - f i r i s s h a d e - i n t o l e r a n t i n the C o a s t a l Western Hemlock Zone ( K r a j i n a 1965). As the sample s i z e f o r D o u g l a s - f i r was s m a l l ( u s u a l l y 2-4 t r e e s cn each of 10 p l o t s ) , i t i s p o s s i b l e t h a t a r e l a t i o n s h i p e x i s t s and I was unable to d e t e c t i t . A l t e r n a t i v e l y , l i g h t c o n d i t i o n s i n the lower p a r t of the canopy as measured by branch s u r v i v a l may be a poor i n d i c a t o r c f l i g h t c o n d i t i o n s higher i n the canopy, as r e f l e c t e d i n l i c h e n abundance. Cf the two f o r e s t measures suggested by B o c h e l l e • s (1978) data, b a s a l area i s not s i g n i f i c a n t l y r e l a t e d to l i c h e n abundance, but height of the codominant t r e e l a y e r accounts f o r 64% of the v a r i a b i l i t y 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 probably r e s u l t s from the f a c t t h at t r e e growth, l i k e v e g e t a t i o n community, i n t e g r a t e s the e f f e c t s of many p h y s i c a l v a r i a b l e s . The low s i t e index of the G a u l t h e r i a s h a l l o n a s s o c i a t i o n -s i f f i i l a r to the D o u g l a s - f i r S a l a l Community of the present study - was noted by Kojima and K r a j i n a (1975). Steep, s o u t h - f a c i n g s l o p e s , which are g e n e r a l l y x e r i c or s u b x e r i c , provide adverse moisture and n u t r i e n t c o n d i t i o n s f o r t r e e growth i n the C o a s t a l western Hemlock zone ( K r a j i n a 1969), but good c o n d i t i o n s f o r l i c h e n growth. I n c r e a s i n g e l e v a t i o n i s s i g n i f i c a n t l y a s s o c i a t e d 84 with decreasing t r e e height and i n c r e a s i n g l i c h e n abundance. I t i s a l s c p o s s i b l e that slow-growing t r e e s provide b e t t e r s u b s t r a t e s f o r l i c h e n s than r a p i d l y growing t r e e s , but t h i s hypothesis has not been e v a l u a t e d . Use of f o r e s t p r o d u c t i v i t y as an i n d i c a t o r of l i c h e n abundance should be done c a u t i o u s l y , and should not be extended to v e g e t a t i o n communities not re p r e s e n t e d i n t h i s study. Bog s i t e s , f o r example, which occur i n the study area have poor t r e e growth but would not n e c e s s a r i l y have high l i c h e n abundance. S e v e r a l f a c t o r s have been noted which are p o t e n t i a l l y u s e f u l i n i d e n t i f y i n g f o r e s t s i t e s with high l i c h e n abundance i n the study area and s i m i l a r areas. Although the sample s i z e d i d not permit t e s t i n g f o r s i g n i f i c a n t d i f f e r e n c e s i n l i c h e n abundance among v e g e t a t i o n communities, a probable ranking c f v e g e t a t i o n communities i n order of t h e i r i n c r e a s i n g p o t e n t i a l f o r l i c h e n production i s suggested by F i g u r e 9; the Moss community, the A m a b i l i s F i r / O v a l - l e a v e d Huckleberry Community, the T r a n s i t i o n Community, and the D o u g l a s - f i r / S a l a l Community . Vegetation community i s a u s e f u l i n d i c a t o r of l i c h e n abundance when considered i n c o n j u n c t i o n with e l e v a t i o n . Topographic f e a t u r e s a s s o c i a t e d with high i n s o l a t i o n may i n d i c a t e high l i c h e n abundance: moderate to steep s l o p e s , south a s p e c t , and absence of topographic f e a t u r e s t h a t block s u n l i g h t . Low f o r e s t p r o d u c t i v i t y may i n d i c a t e good s i t e s f o r l i c h e n growth when i t i s a s s o c i a t e d with x e r i c or s u b x e r i c c o n d i t i o n s . 85 V. AVAILABILITY OF ABBOBEAL LICHENS AND UTILIZATION BY DEES The t h i r d and f o u r t h o b j e c t i v e s of the study are t r e a t e d together as they are c l o s e l y r e l a t e d . The t h i r d o b j e c t i v e was to assess winter a v a i l a b i l i t y and u t i l i z a t i o n of forage l i c h e n s by measuring l i t t e r f a l l i n s i d e and o u t s i d e e x c l o s u r e s . I a l s o wished to examine the timing of 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 t o wind and p r e c i p i t a t i o n . The f o u r t h o b j e c t i v e was to r e l a t e abundance of forage l i c h e n s to s e l e c t i o n of winter h a b i t a t by deer. H a b i t a t s e l e c t i o n was r e l a t e d t o l i c h e n abundance i n two ways; by monitoring deer use of the l i t t e r f a l l p l o t s , and by l o c a t i n g " v i s u a l e s t i m a t e " p l o t s i n areas f o r which i n f o r m a t i o n on winter deer use was a v a i l a b l e {Jones 1975, Harestad 1978). B o c h e l l e (1978) s t u d i e d composition and monthly r a t e s of l i t t e r f a l l i n the study area d u r i n g the winter o f 1973-1974. He found that l i t t e r f a l l r epresented a r e l a t i v e l y important source of winter food f o r deer: amounts of p o t e n t i a l forage ( l i c h e n s and green c o n i f e r f o l i a g e ) i n l i t t e r f a l l approached or exceeded amounts of rooted f o r a g e . Q u a n t i t i e s of ftlectprla ( s . l . ) spp. deposited d u r i n g a 180-day winter period ranged from 2.04 kg/ha on a low e l e v a t i o n s i t e t o 115.35 kg/ha on a m i d - e l e v a t i o n s i t e . L i t t l e seasonal p a t t e r n was apparent i n the monthly data, and no r e l a t i o n s h i p between s n o w f a l l and l i t t e r f a l l c o u l d be determined ( B o c h e l l e 1978). Chickenwire fences were c o n s t r u c t e d around some of the l i t t e r c o l l e c t i o n p l o t s on two s i t e s . Despite the s a a l l sample s i z e and the f a c t t h a t fences were not e n t i r e l y e f f e c t i v e i n e x c l u d i n g deer, q u a n t i f i e s of A l e c t o r i a t s . l . ) were grea t e r i n s i d e than o u t s i d e the fenced areas, and the d i f f e r e n c e 86 was 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 < 0.05) on one s i t e . 87 Methods L i t t e r f a l l measurement l i t t e r f a l l was measured d u r i n g the winter o f 1S76-1977 on p l o t s 3, 4, and 5. Because v i s u a l estimates of l i c h e n abundance were a l s o performed a t these p l o t s , i t was p o s s i b l e to r e l a t e l i t t e r f a l l to s t a n d i n g crop, four a d d i t i o n a l estimate p l o t s (2, 6, 7, and 11) were l o c a t e d a t s i t e s where B o c h e l l e (1978) measured l i t t e r f a l l d u r i n g the winter of 1973-1974. The t h r e e l i t t e r f a l l p l o t s were chosen to r e p r e s e n t an area used by deer under severe winter c o n d i t i o n s , an area used under mild winter c o n d i t i o n s , and an area not used as winter range. P l o t 5 was l o c a t e d i n an area used by a r a d i o s - c o l l a r e d , a d u l t doe during the f a l l o f 1975 (Harestad 1978), and near an area i d e n t i f i e d by G.H. Jones (unpublished data) as poor winter range. I t i s 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 l o c a t e d at 573 m a . s . l . on an ENE-facing s l o p e . When snow f e l l i n l a t e f a l l , the doe moved approximately 2.8 km to the g e n e r a l area of P l o t 4, P l o t 4 i s the s i t e which I expected, on the b a s i s cn i t s v e g e t a t i o n and topographic p o s i t i o n , t o be a T r a n s i t i o n s i t e . I t i s l o c a t e d at 709 m a . s . l . on an E S l - f a c i n g s l o p e . The doe remained i n t h i s area during much of the winter, but moved about 0.6 km t o the area of P l o t 3 at times of heavy s n o w f a l l . P l o t 3 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. I t i s a S E - f a c i n g D o u g l a s - f i r / S a l a l s i t e at 543 m a . s . l . 88 at each l i t t e r f a l l p l o t a deer e x c l o s u r e was c o n s t r u c t e d o f 4 and one-half i n c h (11.4. mm) nylon s e i n e net. The e x c l o s u r e s were 3 to 3.5 meters high and approximately 18 x 21 meters long. Twenty l i t t e r f a l l t r a p s 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 , Th€ ;one meter, sguare l i t t e r f a l l t r a p s were c o n s t r u c t e d of nylon screen s t a p l e d t o a frame of 1x2*s and r e i n f o r c e d at the c o r n e r s with 1/4 i n c h (6.3 mm} plywood gussets. L i t t e r f a l l t r a p s were placed h o r i z o n t a l l y by di g g i n g i n t o the sl o p e on the u p h i l l s i d e and b u i l d i n g up the downhill s i d e with s o i l and p i e c e s of wood, a g e n e r a l i z e d p l o t plan i s given i n Fig u r e 18. P l o t layout was s l i g h t l y modified a t each s i t e because of microtopographic f e a t u r e s and p o s i t i o n s of t r e e s . L i t t e r f a l l p l o t s were e s t a b l i s h e d i n November 1976. To determine the temporal p a t t e r n of l i t t e r d e p o s i t i o n , a d d i t i o n a l l i t t e r f a l l t r a p s were p l a c e d over 13 of the o r i g i n a l t r a p s cn each s i t e i n January and February 1977. The t r a p s t o be covered were randomly s e l e c t e d from the 20 i n s i d e the e x c l o s u r e . In the f i r s t week of Hay 1977, the l i t t e r was c o l l e c t e d from the l i t t e r f a l l t r a p s , bagged and l a b e l l e d , and d r i e d i n a hose dryer f o r storage. L a t e r , the l i t t e r was s o r t e d i n t o three c a t e g o r i e s : - a l e c t o r i a x i n c l u d i n g a l e c t o r i a sarmentosa and E r y o r i a spp. - other l i c h e n s ; and - n o n - l i c h e n l i t t e r f a l l . y Anemometer O O. O O O ' O O O O .0 o o o o o o o o o o o o o o o o o o o o a • • • • • a a • • • • • • • a a' • • • Exclosure • • • • • a ri • a • • • • n a n • • • • o o o o o o o o o o o o 0 o o o- o o o o Q Pellet group plot • Litterfall trap 10 meters Figure 18". Layout of l i tterfal l plots, CO V D 90 The l i t t e r was oven-dried at 60-65°C for 24 hours and weighed with a Mettler P201Q e l e c t r i c scale to the nearest 0.1 g. The Kruskal-Wallis test (Conover 1971: 256) was used to compare l i t t e r f a l l guantities on the three plots. The Mann-Bhitney 0 test (Siegel 1956; 116) was used to compare l i t t e r f a l l quantities inside and outside exclosures. To measure wind conditions, t o t a l i z i n g anemometers were set up approximately 2.5 meters above the ground i n the timber near the exclosures and, where possible, i n adjacent clearcuts. Anemometers were read during January, February, March, and May v i s i t s to the plots. Snow depth was measured during winter v i s i t s i f snow was present. Measurements were taken at 10 systematically located points, and the mean of these readings was used as a measure of snow depth for the plot., P r e c i p i t a t i o n data were obtained from weather stations at Woss Camp and at Port Hardy, approximately 85 km north of the study area on the east coast of Vancouver Island. 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 1976-1977 was monitored by p e l l e t group counts and, when possible, by track counts. At each s i t e f i f t y 9 m* c i r c u l a r p e l l e t group subplots (comparable to 100 f t 2 subplots) were counted and cleared in November and again i n early May. P e l l e t group data were tested for goodness of f i t with several freguency d i s t r i b u t i o n s , and normalized by transformation (Sokal and Eohlf 91 1969: 380). Analysis of variance (Sokal and Rohlf 1969: 204) and Duncan's New Multiple Range Test (Li 1964: 270) were performed on the transformed data. Ihen snow was present during v i s i t s to plots, track counts were made by walking two intersecting 100-meter transects, one p a r a l l e l to the contours and one perpendicular to the contours. A l l deer tracks crossed by the transects were counted. In addition to Plots 3-5, locations of v i s u a l estimate plots 8-10 and 12-14 were chosen on the basis of known deer use during previous winters. Plot 8, an Amabilis Fir/Oval-leaved Huckleberry s i t e , was used by a radio-collared adult buck during mild winter conditions (Harestad 1S78). Plot 9, a Transition s i t e , was used by the same animal when winter weather was severe. Plots 13 and 14, Transition and Amabilis Fir/Oval-leaved Huckleberry s i t e s respectively, represented adjacent habitats which the buck used l i t t l e or not at a l l during winter, although he apparently t r a v e l l e d through them. Plot 10 i s a Douglas-f i r / S a l a l area which a radio-collared adult doe moved into at the time of an autumn snowfall. I t i s located i n an area i d e n t i f i e d by Jones (unpublished data) as good winter range which receives high deer use. Plot 12, which was u n c l a s s i f i e d , was used as winter range by a two-year-old radio-collared buck. 9 2 Jesuits Total l i t t e r f a l l guantities Quantities of the various l i t t e r components deposited inside exclosures on the three s i t e s are given i n Table VII. Because l i t t e r c o l l e c t i o n periods varied s l i g h t l y among s i t e s , r esults were adjusted for a 180-day period. Host of the d i s t r i b u t i o n s of the l i t t e r f a l l samples were strongly skewed rig h t (e.g. Q= 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 guantile method (Conover 1971:110). The medians differed considerably from the means, as i s expected i n skewed d i s t r i b u t i o n s . Alectoria (s.l.) guantities ranged from 31.9 to 151.2 kg/ha on the three s i t e s ; r esults of the Kruskal-Hallis test (Conover 1971: 256) indicated highly s i g n i f i c a n t differences in Alectoria levels among s i t e s (T = 47.59; T o o s = 5.99) . Non-lichen l i t t e r f a l l ranged from 867.5 to 1005.0 kg/ha, and t o t a l l i t t e r f a l l from 978.4 to 1071.1 kg/ha. Differences among s i t e s i n ncn-lichen l i t t e r f a l l and t o t a l l i t t e r f a l l were also s i g n i f i c a n t but at a much lower l e v e l (non-lichen: T = 10.33; t o t a l : T = 6.96; T 0 O 5 = 5,99). Quantities of non-Alectorioid lichens ranged from 34.2 to 45.5 kg/faa; differences among plots were not s i g n i f i c a n t (T = 0.71; T 0 0 5 = 5.99). 93 TABLE VII. L i t t e r f a l l quantit ies inside exclosures L i t t e r f a l l Components Aleotovia Other Lichen Non-Lichen Total P lot 3 (DF/S) Mean for 180-day period (kg/ha) 69.9 41.0 867.5 978.4 Median for 180-day period (kg/ha) 62.9 28.5 662.9 786.9 95% confidence interva l around median 60.0-81.9 22.6-40.3 556.7-759.4 629.5-908.9 Plot 4 (Unclass if ied) Mean for 180-day period (kg/ha) 151.2 46.5 868.0 1065.7 Median for 180-day period (kg/ha) 130.8 31.5 737.7 890.2 95% confidence interva l around median 116.1-166.2 22.6-55.1 537.1-784.9 694.4-1012.1 Plot 5 (AF/OH) Mean for 180-day period (kg/ha) 31.9 34.2 1005.0 1071.1 Median for 180-day period (kg/ha) 25.9 26.9 1019.4 1086.5 95% confidence interval around median 22.5-40.5 20.9-43.8 762.8-1197.3 800.9-1254.6 94 Lichen l i t t e r f a l l i n r e l a t i o n to lichen biomass Table VIII presents lichen l i t t e r f a l l and l i c h e n biomass data for seven s i t e s . The 1973-1974 data were co l l e c t e d by Bochelle (1978), Data from the two studies must be compared cautiously, as differences i n sampling methods and i n weather between years probably influenced r e s u l t s , Bochelle*s plots were subject to grazing (with the exception of ten l i t t e r f a l l traps on Plot 7) whereas the 1976-1977 data are based on exclosed plots. Despite these differences, ranges of Alectoria l i t t e r f a l l values are comparable. Alectoria l i t t e r f a l l quantities appear to be related to Alectoria biomass i n the canopy, as indicated by the r e l a t i v e l y narrow range of r a t i o s of l i t t e r f a l l to standing crop. Percentages of l i t t e r f a l l based on Bochelle as (1978) data range from 7,6 to 9,4%, but f a l l e n lichens were subject to removal by ungulates. L i t t e r f a l l percentages based on the present study are higher: 10.5 - 16.1%. L i t t e r deposition i n r e l a t i o n to time and weather Daily deposition of Alectoria l i t t e r , other lichen l i t t e r , and t o t a l l i t t e r are given i n Figures 19-21. The early winter period includes November to mid-January; mid-winter, mid-January to l a t e February; and l a t e winter, late February to the f i r s t week of May. On a l l three plots, deposition rates of both l i t t e r components and of t o t a l l i t t e r increased consistently during the winter. Deposition rates of A l e c t o r i a increased from early to late winter by factors of 12.7, 9.5, and 25,6 on the three plots, whereas deposition rates of t o t a l l i t t e r f a l l increased 95 TABLE VIII. Alectoria l i t t e r f a l l in re la t ion to Alectoria standing crop Plot Alectoria standi ng crop (kg/ha) Alectoria l i t t e r f a l l 1976-77 (kg/ha) Alectoria l i t t e r f a l l 1973-74 (kg/ha) 1 L i t t e r f a l l / Standing Crop (%) 2 175 14.2 8.1 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 2000-05 JZ cn i 1 _ i 1500-_ j < LL DC LU h- 1000-h-_1 < DC 500-o h-o LU _J < P L O T [ f l M 3 L J5L M L 4 M 5 mean median and 95% confidence l im i t s Figure 19. Deposition rates of Alectoria l i t t e r in ear ly winter (E), mid-winter (M), and late winter (L) , 1976-1977. 97 < LL DC LU Z LU X o cc LU X o 1500-1 ^ I O O O H CO •a cc sz cn 500H PLOT 0 M .3 M 4 M L 5 mean median and 95% confidence l im i t s Figure 20;. Deposition rates of other (non-Alectorioid) l ichen l i t t e r in ear ly winter (E), mid-winter (M), and late winter (L), 1976-1977. 98 ! tOOOOi 8000 CO CO D) 6 0 0 0 H < ^ 4C00-LU \z _ J _ j 2000H _l < 0 0 E M L E M L E M L P L O T 3 4 5 mean median and 95% confidence l im i t s Figure 21. Deposition rates of a l l l i t t e r in early winter (E), mid-winter (M), and late winter (L), 1976-1977. 99 only 2.2, 3.7, and 2.7 times. D e p o s i t i o n r a t e s o f other l i c h e n s f o l l o w e d an i n t e r m e d i a t e r a t e of i n c r e a s e , i n c r e a s i n g hy f a c t o r s of 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 at Boss Camp and Port Hardy during e a r l y , mid- and l a t e winter of 1976-1977 are given i n Figu 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 winter and l e a s t during mid-winter. No r e l a t i o n s h i p between p r e c i p i t a t i o n and l i t t e r f a l l i s apparent. The winter of 1976-1977 was unusually mild. The onl y major s n o w f a l l occurred i n l a t e March 1977, d u r i n g the l a t e winter p e r i o d . I t i s p o s s i b l e t h a t the i n c r e a s e d l i t t e r f a l l during t h i s p e r i o d was r e l a t e d t o t h a t storm, but th a t p o s s i b i l i t y cannot be evaluated with a v a i l a b l e data. Figure 23 summarizes r e l a t i v e amounts of wind i n the timber by the e x c l o s u r e s and i n adjacent c l e a r c u t s d u r i n g the three p e r i o d s . There i s no c l e a r c u t adjacent to P l o t 4. U n f o r t u n a t e l y , the anemometer l o c a t e d i n the c l e a r c u t by P l o t 5 d i d not f u n c t i o n p r o p e r l y , as the cups d i d not s p i n f r e e l y , the recorded readings probably underestimated true amounts of wind., Hind c o n d i t i o n s near ground l e v e l i n c l e a r c u t s should approximate wind c o n d i t i o n s at the top of the f o r e s t canopy, although more tu r b u l e n c e i s expected i n the canopy (Geiger 1966: 116, 274, 312). I had a n t i c i p a t e d t h a t anemometer re a d i n g s from the timber 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 cor r e s p o n d i n g c l e a r c u t r e a d i n g s , so t h a t timber readings could be co n s i d e r e d i n d i c e s to wind c o n d i t i o n s at the top of the canopy. Without r e l i a b l e r e a dings from the c l e a r c u t at P l o t 5, the assumption cannot be j u s t i f i e d . 100 o 10 — E O E UJ DC 0_ 5H E M L Port Hardy E M Woss Figure 22. P rec ip i ta t ion at Port Hardy and Woss Camp in ear ly winter (E), mid-winter (M), and la te winter (L ) , 1976-1977. 101 I 40-DC LU LU o LU zz. < c DC LU £ P L O T CO CO 20H M L 3 M 4 M 5 80-LU \— LU o w CO < ~-CO D 1 4 0 O DC < LU _J O PLOT E M L 3 E M L 5 Figure 23. Relative amounts of wind in mature timber (A) and clearcuts (B) in ear ly winter (E), mid-winter (M), and late winter ( L ) , 1 9 7 6 - 1 9 7 7 . 102 Anemometer rates on a l l s i t e s tended to be lower i n early winter than i n mid- or late winter. In the clearcut by Plot 3, anemometer rates increased as winter progressed, but changes i n l i t t e r f a l l rates were much greater than changes i n wind rates as measured by anemometers. The data do not preclude a relationship between l i t t e r f a l l rates and wind as measured by anemometers, but 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 outside the exclosure on Plot 5 were inadvertantly exposed to moisture during storage, and began to decompose. P a r t i a l decomposition affected the Alec t o r i a sarmentosa and Bryoria spp. more than the other lichens, and made i t impossible to separate them adequately from the rest of the sample. The Mann-Whitney U test (Siegel 1956:116) was used to determine whether weights of the p a r t i a l l y 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 unaffected samples., Results indicated that differences i n weights approached s i g n i f i c a n c e only i n the Alectoria category <U = 16.5; U 0 o S = 14.0).* Therefore, the p a r t i a l l y decomposed samples of A l e c t o r i a . but not of other l i t t e r components were eliminated from comparisons of l i t t e r f a l l inside and outside the exclosure. 1 In the Mann-Whitney U t e s t , the n u l l hypothesis i s rejected i f the calculated value for U i s smaller than the tabulated value. 103 Quantities of l i t t e r f a l l inside and outside exclosures are compared in Table IX. Results of the Mann-Whitney U test (Siegel 1956: 116) indicate that quantities of alectoria were s i g n i f i c a n t l y greater inside than outside exclosures on a l l three s i t e s , despite the reduced sample siz e at Plot 5. Quantities of non-Alectorioid lichens insi d e and outside exclosures did not d i f f e r s i g n i f i c a n t l y at any s i t e s , and i n one case were greater outside than inside. Amounts of non-lichen l i t t e r f a l l were also similar inside and outside exclosures and, i n cne case, guantities outside were greater than quantities inside. The s i g n i f i c a n t difference between t o t a l l i t t e r f a l l inside and outside the exclosure at Plot 4 was due partly to the larqe difference i n A l e c t o r i a quantities. A comparison of Alectoria with the other l i t t e r f a l l components suggests that differences i n Alecto ria guantities inside and outside exclosures were due to removal by herbivores, and not to fortuitous s p a t i a l patterns of l i t t e r deposition. Besults also indicate that consumption of non-Alectorioid lichens was minimal. These re s u l t s are consistent with those of Bochelle (1978)., who found that Alectoria (s. 1.) , but not other lichens, was a major forage item in winter. As non-lichen l i t t e r f a l l was not separated into components, no conclusions may be drawn about consumption of l i t t e r components other than lichens. The difference between Alectoria quantities inside and outside each exclosure provides an approximation of Alectoria u t i l i z a t i o n at each s i t e (Fig. 24). Using the Mann-Whitney s t a t i s t i c to calculate 95% confidence l i m i t s for the difference TABLE IX. L i t t e r f a l l quantit ies inside 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 Outside exclosure Mann-Whitney U 69.9 44.2 47 1 41.0 37.3 1861 867.5 1108.8 1401 878.4 1190.7 151 1 Plot 4 Inside exclosure Outside exclosure Mann-Whitney U 151.2 71.2 20.5 1 46.5 43.6 186.5 1 868.0 793.6 141.5 1 1065.7 908.5 127 1 Plot 5 Inside exclosure Outside exclosure Mann-Whitney U 31.9 15.2 182 34.2 55.4 1631 1005.0 924.1 1551 1071 .1 994.9 146 1 1 U.Q25 =127 2 U.025 = 20 S denotes s t a t i s t i c a l s ign i f icance at p < .025. 105 | 100 - i 80 H z: O N co 3] TJ 60 H o 3 03 < 5 cr \ 4 o - | (~\ CD o LU _ J < 20H PLOT Figure 24. Alectoria u t i l i z a t i o n on Plots 3, 4, and 5, with 95% confidence l i m i t s . 106 between two means (Conover 1971:238), u t i l i z a t i o n a t P l o t 3 was 25.7 (13.5 t o 35.3) kg/ha/180 days; u t i l i z a t i o n a t P l o t 4 was 80.0 (46.3 to 92.5) kg/ha/180 days; and u t i l i z a t i o n at P l o t 5 was 16.7 (15,.8 to 28.1) kg/ha/180 days. T h i s represented 31%, 53%, and 52% u t i l i z a t i o n of a v a i l a b l e Jklectoria, is. 1,) . Deer use of l i t t e r f a l l p l o t s Results of p e l l e t group counts i n d i c a t e g r e a t e r use of winter range than of non-winter range, d e s p i t e the extremely mild winter. Mean values of 1.12, 0.80, and 0.40 p e l l e t groups per subplot were present on the severe winter range s i t e ( P l o t 3 ) , the mild winter range s i t e ( P l o t 4), and the poor winter range s i t e ( P l o t 5), r e s p e c t i v e l y . The data were found to f i t the negative binomial d i s t r i b u t i o n , 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 of the a n a l y s i s of v a r i a n c e i n d i c a t e d a h i g h l y s i g n i f i c a n t d i f f e r e n c e among p l o t s (p<0.01). B e s u l t s of Duncan's New M u l t i p l e Range Te s t showed t h a t the means of P l o t s 3 and 4 d i d not d i f f e r s i g n i f i c a n t l y , but the mean of P l o t 5 d i f f e r e d from t h e ethers (p<0.05). Two e l k p e l l e t groups were found i n s i d e s u b p l o t s a t P l o t 4, i n d i c a t i n g some winter use of the area by Roosevelt e l k . A few elk p e l l e t groups were a l s o noted i n the v i c i n i t y of P l o t 3, though none appeared to be f r e s h i n s p r i n g 1977. No e l k s i g n was observed at P l o t 5. T h i s low l e v e l o f e l k use i n d i c a t e s t h at i f any l i c h e n removal was due to e l k r a t h e r than to deer, i t was probably minor. 107 The most s i g n i f i c a n t snowfall of the winter occurred cn March 27tfc, 1977, Track counts were made and snow depths measured on March 28 and 29. Results (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 to represent severe winter range, average snow depth at the s i t e was only 7 cm. Plot #, t h e m i l d winter range area, had much deeper snow (53 cm) and no signs of deer use aft e r the snowfall. The area around Plot 5, where snow depth was intermediate (26 cm), was evidently being used by one animal. Lichen abundance and deer use Available information on the status of li c h e n plots as winter deer habitat i s summarized i n Table XI. I t i s d i f f i c u l t to draw firm conclusions from the data available here because arboreal lichens represent only part of the available forage. A v a i l a b i l i t y of rooted forage, p a r t i c u l a r l y shrubs, was not measured as part of t h i s study, but i s undoubtedly an important factor in winter habitat selection., With the exception of Plots 8 and 12, areas used in winter were moderate or high i n lichen abundance, whereas areas not selected i n winter were low i n lichen abundance. Shrubs are high and dense at Plot 8, and are probably the major source of forage for wintering deer. Selection of Plot 12 as winter habitat i s d i f f i c u l t to explain, as neither lichens nor shrubs are abundant. The tendency for deer to use moderate or high lichen areas in winter, coupled with the data on u t i l i z a t i o n of lichens, strongly suggests that li c h e n a v a i l a b i l i t y i s a factor influencing winter habitat selection. TABLE X. Deer use and snow depths at Plots 3, 4, and 5, March 28-29, 1977 Plot Date T r a c k D e e r S n o w Count Seen Depth (cm) March 28 30 4 March 28 0 0 53 March 29 0 1 26 109 TABLE' XI. Habitat se lect ion and l ichen biomass Plot Lichen Biomass (kg/ha) Summary of Habitat Selection Information 3 665 Used in severe winter by OFL 60 1 , good winter range 2. 4 942 Used in mild winter by OFL 60 1 . 5 303 Used in f a l l by OFL 60 1 , poor winter range 2. 9 800 Used in severe winter by OFL 62 1 . 8 289 Used in mild winter by OFL 62 1 . 14 176 Used occasional ly in mild winter by OFL 62 1 . 13 107 Not used in winter by OFL 62^. 10 521 Used in f a l l when snow present by OFL 68 , good winter range 2. 12 21 Used in Winter by OFL 6 1 l . 1 Harestad 1978. 2 Jones 1975 and unpublished data. 110 Discussion L i t t e r f a l l The lichen l i t t e r f a l l t o t a l s obtained in t h i s study are compared with r e s u l t s of other studies in Table XII. The range of values reported here i s somewhat higher than.that reported by Eochelle (1976) and Kale (unpublished data) V in their studies on Vancouver Island, but i n both of the l a t t e r cases l i t t e r f a l l was subject to consumption by herbivores. The l i t t e r f a l l values reported by Denison (1973) for Lobaria oregana and Andre et a l . (1975) for a l l arboreal lichens f a l l within the range of values reported here. The maximum value for a l l arboreal lichens reported by Schroeder (1974) for a subalpine forest i n i n t e r i o r B r i t i s h Columbia exceeds the maximum value found in the study area by more than 75 kg/ha. As Schroeder's annual rates of l i t t e r f a l l were based on summer measurements, they may underestimate true values. L i t t e r f a l l taken as a percentage of standing crop ranged from a minimum of 1.2% (Andre et a l . 1975) to a maximum of 22,9% (Denison 1973). The values reported i n the present study are intermediate. * I am grateful to W. Kale and D. Hebert of the B r i t i s h Columbia Fish and W i l d l i f e Branch for permission to use these data. I l l TABLE XII. Comparison of reported quantit ies of lichen l i t t e r f a l l Source Area Forest Type Lichens Lichen L i t t e r f a l l kg/ha/year Standing Crop (kg/ha) L i t t e r f a l l As % of Standi ng Crop Denison 1973 Western Oregon Pseudotsuga menziesii Lobaria oregana 89.7 392.3-504.4 17.8-22.! Schroeder 1974 Se lk i rk Mountains, B.C. and U.S.A. Picea engelmannii-Abies lasiocarpa; Larix occidentalis A l l arboreal lichens 7.8-273.9 1 Andre et a l . 1975 France Abies alba A l l arboreal l ichens; mainly ^ ^ gg -j Pseudevernia furfuracea 1040.0 7.2- 9.5% W. Kale (unpub-l ished data) Northwest Bay, Vancouver Island Pseudotsuga menziesii A l l arboreal lichens Alectoria, Bryoria, and Usnea spp. 1.4-105.92 0.3- 99.42 Rochelle (1976) and th i s study Nimpkish Va l ley, Vancouver Island Tsuga heterophylla-Pseudotsuga menziesii-Thuja plicata A l l arboreal l ichens Alectoria and Bryoria spp, 40.5-162.6 2 13.2-115.3 2 7.6- 9.4% This study Tsuga heterophylla-Pseudotsuga menziesii-Thuja plicata A l l arboreal lichens 66.1-197.7 2 Alectoria and Bryoria spp. 31.9-151.2 2 10.5-16.1% 1 Based on summer rates extended to 1 year. 2 Based on 180-day winter period. 112 Seme workers have assumed that epiphyte biomass i s i n a steady state i n an old-growth forest; growth i s balanced by l i t t e r f a l l , i n -situ decomposition, and consumption by herbivores (Pike et a l . 1972). I f t h i s i s true, then annual, l i t t e r f a l l provides an estimate of annual turnover and thereby annual growth, since the other pathways are probably r e l a t i v e l y minor. To my knowledge, i t has not been demonstrated empirically that there i s no net change i n epiphyte biomass in old-growth forests. The results of t h i s study provide information 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 relationship to standing crop of Alectoria (s.l.) l i t t e r f a l l i n a single winter. No information i s provided on variation in l i t t e r f a l l among years. A regular increase in l i t t e r f a l l rates during the winter was apparent in the present study, but i t i s not known whether that i s a general pattern. F i n a l l y , no relationship between weather and rates of l i t t e r f a l l could be established. Other studies provide information on variation i n l i t t e r f a l l guantities between years. The value f o r lichen l i t t e r f a l l reported by Andre et a l . (1975) for 1970-1971 was 76% of that measured at the same s i t e i n 1972-1973; t o t a l l i t t e r f a l l in 1970-1971 was 91% of t o t a l l i t t e r f a l l in 1972-1973. In Kale's study area, Alectoria (s.l.) l i t t e r f a l l (including: Usnea ) was consistently lower during the extremely mild winter on 1976-77 than in 1975-1976, averaging only 3H% of the 1975-1976 values. Other lichens and t o t a l l i t t e r f a l l were both lower in 1S76-1S77 on 4 of the 5 s i t e s studied, averaging U7% and 55%, respectively, of the 1975-1976 values. Evidently, variation in 113 lichen l i t t e r f a l l from one year to another can be considerable. If the lower l i t t e r f a l l values reported by Kale for 1976-1977 were associated with the unusually mild winter, then i t i s possible that the values reported i n t h i s study are lower than i s usual for the study area. Timing of l i t t e r deposition, as well as quantity, i s important to wintering deer. I f l i t t e r f a l l rates are highest i n late winter, 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 coincides with the period of maximum stress for the animals. Data of Bochelle (1978) and Kale (unpublished) presented for comparison in Figures 25 and 26 indicate that the pattern may not be general. L i t t e r f a l l rates not only fluctuated e r r a t i c a l l y during the course of winter, but temporal patterns varied from one s i t e to another within a study area. It seems l i k e l y that lichen l i t t e r f a l l rates are a function of standing crop and of weather fac t o r s , which may be highly s i t e - s p e c i f i c , Abee and Lavender (1972) reported that most l i t t e r i n a mature Douglas-fir stand i n Oregon f e l l during winter and attributed t h i s pattern to breakage of branches due to snow. Pike et a l . (1972) suggested that the increased weight of epiphytes when wet might be important i n causing branch breakage. Other authors (Cowan 1945, Gates 1968) attributed lichen l i t t e r f a l l to snow and wind. Bochelle (1978) observed that l i t t e r f a l l i s probably influenced most by intense i n d i v i d u a l storms rather than o v e r a l l weather patterns. 150CH CO •a CO O) ioooH cc LU I— LU o < cc o LL 500-5900 r; CIIDL N - D - J - F - M - A N - D - J - F - M - A N - D - J - F - M - A D - J - F - M - A J - F - M - A S I T E 1 2 3 4 5 Figure 25. Monthly deposition rates of forage l i t t e r (lichens and green conifer fo l iage) in the study area, 1973-1974 (Rochelle 1978). 800H * 6 0 ° -j - 400-j CO 200A A L E C T O R I A g/ha /day O N D J F M 800-CO 600H UJ I— 400-] CO 200-O N D J F M 800-T - 600-^ 400-00 200- J L 800-co 600-1— £ 400-200 O N D J F M 800H 600H 400-200H O T H E R L I C H E N g/ha/day O N D J F M 800-600-400-200-O N D J F M 800 600-400-200-O N D J F M 800-600-400-200-O N D J .F M 30H 20H 10H N O N - L I C H E N kg/ha/day 4 6 O N D J F M 30-1 20-10-O N D J F M 30H 20H 10H O N D J F M 30H 20-ion O N D J F M O N D J F M Figure 26. Monthly deposition rates of three l i t t e r f a l l components at Northwest Bay, 1975-1976 (Kale, unpublished data). 116 The interaction of climatic factors i s probably important in producing lichen l i t t e r f a l l . Von Schrenk (1898) found that Usnea t h a l l i were about 3 times heavier when wet than when dry, and had about 25% as much t e n s i l e strength. In a 20 foot (6.1 m) wind tunnel dry t h a l l i did not break at v e l o c i t i e s of 124 km/hr, but wet 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. wind may 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 i n a rainstorm or snowstorm than under dry conditions. I suspect that detailed, frequent, and s i t e - s p e c i f i c monitoring of weather conditions would be necessary to demonstrate their influence on l i t t e r f a l l rates. Deer use and lichen abundance assessment of deer use of the l i t t e r f a l l plots indicated higher deer use and higher lichen abundance i n the areas thought to represent winter range than i n the area thought to be poor winter range. Evaluation of lichen abundance in areas selected by radio-collared deer i n winter indicated a tendency for animals to use areas having r e l a t i v e l y high lichen abundance. There i s no one-to-one correspondence between winter range and a v a i l a b i l i t y of forage lichens. Rather, I suggest t h a t deer select areas where food a v a i l a b i l i t y i s high, and that arboreal lichens are an important component of winter food. available a l e c t o r i a (s.l.) ranged from 31.9 to 151.2 kg/ha for a 180-day winter period. U t i l i z a t i o n ranged from 16.7 to 80.0 kg/ha f o r the same period. Quantities of available rooted forage were not measured as part of t h i s study, but Rochelle (1978) reported an average of 42 kg/ha on low elevation s i t e s 117 and 149 kg/ha on mi d - e l e v a t i o n s i t e s i n the same study area. R e s u l t s of t h i s study support B o c h e l l e * s c o n c l u s i o n t h a t amounts of forage p o t e n t i a l l y a v a i l a b l e i n l i t t e r f a l l are comparable to a v a i l a b l e r ooted forage, and demonstrate that as much as 53% of a v a i l a b l e A l e c t o r i a l i t t e r f a l l may be u t i l i z e d . 118 VI. SUMMARY AND MANAGEMENT RECOMMENDATIONS Summary 1. A l e c t o r i a ( s . l . ) biomass on ten sampled t r e e s ranged from 436 to 7036 g. 2. The r e g r e s s i o n y = 158.03{A x CL), (8) where y i s l i c h e n biomass, A i s an estimate o f A l e c t o r i a ( s . l . ) cover on a p o r t i o n of the t r e e crown, and CL i s crown l e n g t h , had an r 2 of 0.75 and was used t o estimate l i c h e n biomass on tr e e s t h a t were not sampled. 3. Estimates of A l e c t o r i a ( s . l . ) biomass on the 14 s i t e s s t u d i e d ranged frcm 21-1528 kg/ha. 4. O p t i c a l d e n s i t y v a l u e s and redrgreen f i l t e r r a t i o s f o r t r e e s with high l i c h e n biomass d i d not 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, but d i f f e r e n c e s i n red:green f i l t e r r a t i o s f o r western hemlock approached s t a t i s t i c a l s i g n i f i c a n c e . 5. Photo i n t e r p r e t e r s * r a n k i n g s of t r e e s a c c o r d i n g t o l i c h e n 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 with v i s u a l e s t i m a t e s of l i c h e n abundance i n 39% of cases. 6. Previous experience with photo i n t e r p r e t a t i o n and knowledge 119 of l i c h e n ecology p o s i t i v e l y a f f e c t e d i n t e r p r e t e r s 1 success i n judging l i c h e n abundance. 7. Photo i n t e r p r e t e r s were unable to d i s t i n g u i s h among s i t e s with high and low l i c h e n abundance. 8. Of the f o u r v e g e t a t i o n communities d e s c r i b e d , the Douglas-f i r / S a l a l Community and the T r a n s i t i o n Community appeared to have the g r e a t e s t p o t e n t i a l f o r l i c h e n p r o d u c t i o n . 9. A r e g r e s s i o n of a l e c t o r i a ( s . l . ) abundance on p o t e n t i a l annual s o l a r r a d i a t i o n was s t a t i s t i c a l l y s i g n i f i c a n t and accounted f o r 31% of the v a r i a t i o n i n A l e c t o r i a abundance. 10. A r e g r e s s i o n of A l e c t o r i a ( s . l . ) abundance on e l e v a t i o n was s t a t i s t i c a l l y s i g n i f i c a n t and accounted f o r Hl% of the v a r i a t i o n i n A l e c t o r i a abundance. Much of t h e r e s i d u a l v a r i a t i o n appeared e x p l a i n a b l e by v e g e t a t i o n community. 11. Taken s e p a r a t e l y , s l o p e and aspect were not s i g n i f i c a n t l y r e l a t e d t o A l e c t o r i a ( s . l . ) abundance, but i n combination with e l e v a t i o n , these f a c t o r s accounted f o r 82% of the v a r i a t i o n i n A l e c t o r i a abundance. 12. Crown c l o s u r e as measured by v i s u a l e s t imate and by the moosehorn method were not s i g n i f i c a n t l y r e l a t e d by A l e c t o r i a ( s . l . ) abundance, but densiometer measurements of crown c l o s u r e accounted f o r UQ% of the v a r i a t i o n i n A l e c t o r i a abundance. 120 13. Basal area was not s i g n i f i c a n t l y related to alectoria-(s.l.) abundance. 14. The r a t i o of crown length to tree height appeared to be related to a l e c t o r i a (s.l.) abundance only on Douglas-fir/Salal s i t e s and on Transition s i t e s . 15. Forest productivity, as measured by the height of the codeainant tree layer, accounted for 64% of the v a r i a b i l i t y i n al e c t o r i a (s.l.) abundance. 16. Quantities 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 days; differences among s i t e s 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 . 1-7. Alectoria (s.l.) l i t t e r f a l l represented 10.5-16.1 % of A l e c t o r i a biomass in the tree canopy. 18. Alectoria (s.l.) deposition rates increased from early winter to l a t e winter on a l l three plots, but comparison:with other studies suggested that t h i s pattern may not be general. 19. No relationship could be established between l i t t e r f a l l rates and precipitaton or l i t t e r f a l l rates and wind, as measured by t o t a l i z i n g anemometers. 20. Differences between Alectoria (s.l.) accumulation inside and 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 t h r e e study s i t e s , i n d i c a t i n g consumption of these l i c h e n s by h e r b i v o r e s . 21. D i f f e r e n c e s between accumulation 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 o u t s i d e e x c l o s u r e s were 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 , i n d i c a t i n g t h at consumption of these l i c h e n s by h e r b i v o r e s was minimal. 22. Based on A l e c t o r i a ( s . l . ) 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 , u t i l i z a t i o n was estimated at 25.7 kg/ha/180 days i n a severe winter range a r e a , 80.0 kg/ha/180 days i n a mild winter range area, and 16.7 kg/ha/180 days i n a poor winter range area. 23. Deer use during the winter of 1976-1977, as measured by p e l l e t group counts and t r a c k counts, was g r e a t e r i n the severe and mild winter range areas than i n the poor winter range area. 24. Areas s e l e c t e d by deer as winter range tended to be moderate or high i n l i c h e n abundance, although t h e r e was no one-to-one correspondence. 122 Becommendations Be s u l t s of t h i s study support e a r l i e r f i n d i n g s t h a t A l e c t o r i a ( s . 1 . ) l i t t e r f a l l i s an important component of the s i n t e r d i e t of b l a c k t a i l e d deer, even i n a mild w i n t e r . In areas t h a t are to be managed f o r b l a c k t a i l e d deer, l i c h e n abundance should be considered when mature timber stands are set a s i d e as winter range. I f managers are to c o n s i d e r a r b o r e a l l i c h e n s , they must be able t o i d e n t i f y areas where l i c h e n abundance i s high. B e l a t i o n s h i p s between l i c h e n abundance and s i t e c h a r a c t e r i s t i c s d e s c r i b e d i n Chapter IV 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 such areas w i t h i n the C o a s t a l Western Hemlock Zone ( K r a j i n a 1965). P r e l i m i n a r y s t r a t i f i c a t i o n of s i t e s with south a s p e c t , moderate to steep s l o p e s , r e l a t i v e l y high e l e v a t i o n s , and,low f o r e s t p r o d u c t i v i t y c o u l d be accomplished with topographic maps and 20-or 40-chain photographs. Methods f o r r a p i d f i e l d assessment of A l e c t o r i a ( s . l . ) abundance are needed. Q u a n t i f i c a t i o n of l i c h e n biomass, as c a r r i e d out i n t h i s study, i s too time-consuming f o r i n v e n t o r y purposes but some of the methods d e s c r i b e d here c o u l d be adapted. Estimates of percent cover of A l e c t o r i a ( s . l . ) on a sample of t r e e s c o u l d be used t o o b t a i n i n d i c e s of l i c h e n abundance on s i t e s , s i m i l a r to the measfure "percent A l e c t o r i a " used i n Chapter IV. Measurement of crown lengths and stems/ha would be unnecessary. Such i n d i c e s would permit comparison of s i t e s w i t h i n a study a r e a , but could not be r e l a t e d t o a c t u a l l i c h e n biomass. The r e g r e s s i o n used i n the present study to 123 p r e d i c t l i c h e n biomass on unsampled t r e e s should not be e x t r a p o l a t e d to other areas without t e s t i n g . 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 on methods emerge from the present study. Ideas f o r improving methods c f g u a n t i f y i n g l i c h e n abundance d i s c u s s e d i n Chapter I I I -p a r t i c u l a r l y a p p l i c a t i o n of the 3P method - r e g u i r e t e s t i n g . The u s e f u l n e s s of o b l i q u e a e r i a l photographs f o r l i c h e n i n v e n t o r y could be assessed. C l a r i f i c a t i o n of s e v e r a l a s p e c t s of the b i o l o g y of A l e c t o r i o i d l i c h e n s would provide i n f o r m a t i o n u s e f u l t o managers. S t u d i e s of 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 would have i m p l i c a t i o n s f o r s i z e and spacing of c l e a r c u t s i n areas t o be managed as winter range. W i l d l i f e managers need i n f o r m a t i o n about growth r a t e s of l i c h e n s i n developing f o r e s t s to p r e d i c t changes i n forage a v a i l a b i l i t y d u r i n g s u c c e s s i o n . The hypothesis t h a t slow-growing t r e e s provide b e t t e r s u b s t r a t e s f o r l i c h e n s t than r a p i d l y - g r o w i n g t r e e s should be t e s t e d . 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S c i . 56 (3): 5 31-542. 131 APPENDIX I C a l c u l a t i o n of A l e c t o r i a (sensu l a t o ) Biomass T o t a l s f o r P l o t s Symbols used i n appendix; AF - a m a b i l i s f i r (Abies a m a b i l i s ) DF - d o u g l a s - f i r (Pseudotsuga m e n z i e s i i ) G F - grand f i r JAbies grandis) MH - mountain hemlock (Tsuga mertensianaV WH - western hemlock (Tsuga heterophylla) WP - white pine j g i n u s monticola) WRC - western red cedar (Thuja p l i c a t a ) . -YC - yellow c y p r e s s (Cfaamaecyparis nootkatensis) C - codominant D - dominant I - i n t e r m e d i a t e Footnotes 1/ Based on v i s u a l estimates except as i n d i c a t e d . 2/ Based on biomass sampling. 3/ Species/dcminance c l a s s not sampled; based on most s i m i l a r species/dominance c l a s s 132 PLOT 1 Species/ Lichen x Lichen Number Total Lichen Dominance Tree Biomass/ Biomass/ of Trees Biomass/ Class Number Tree (g) Class (g) in Class Class (kg) WH-D 112 56 92.59 56 92.59 5 28. 46 WH-C 2 4956.33 46 90.89 51 239.24 13 70 36.22 27 4933.80 38 3614.26 45 4716.03 79 2950.93 81 4628.63 WH-I 22 1850.93 2258.85 48 108. 42 31 2666.77 WHC-C 69 1772. 96 2184.87 28 61. 18 70 2596.78 WBC-I 17 95.96 26 5.97 22 5.86 119 435.97 DF-D 84 3686.52 4874.82 6 29.25 101b 6063.12 DF-C 34 502.9 3 1705.34 9 15.35 98 3106.25 101a 1506.85 AF-I 1705.34 1 DF-I 1705.34 1 Total lichen biomass in plot (kg) 1/plot area (ha) Total lichen biomass (kg/ha) 491. 16 x 3. 11 1528.04 133 PLOT 2 Species/ Lichen x Lichen Number Total Lichen Dominance Tree Biomass/ Biomass/ of Trees Biomass/ Class Number Tree(g) Class (g) in Class Class (kg) WBC-I 5 4785.90 4785.90 3 14. 36 WH-C 13 620.89 56 5.97 11 6.23 57 511.04 HH-I 12 90. 14 72.21 26 1.88 26 43.62 41 82.87 SfiC-C 14 947.26 464.46 22 10.22 31 93.78 33 202.09 42 555.21 48 226.80 49 1034.89 59 191.18 DF-D 50 1809.21 2187.86 5 10.94 55 1521.12 61 3233.26 DF-C 54 570.65 1095.50 3 3. 30 6 3 1628.34 8 8C-C 7 0.51 Total l i c h e n biomass i n plot (kg) 1/plot area (ha) Total lichen biomass (kg/ha) 47. 44 x 3.69 175. 25 134 P L O T 3 Species/ Lichen x Lichen Dominance Tree Biomass/ Biomass/ Class Number Tree(g) Class (g) Number Total Lichen of Trees Biomass/ in Class Class (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 5 134.97 152.91 4 .61 17 170.84 Df-D 18 3474. 54 3474.54 2 6.95 DF-C 2 2498.64 3193.49 16 51.10 7 1507.6 4 8 2304.07 10 3631.95 13 4631. 19 28 2498.64 32 3887.96 35 4587.79 DF-I 9 3679.66 3679.66 1 3.68 Total lichen biomass i n plot (kg) 1/plot area (ha) Total l i c h e n biomass (kg/ha) 6 6 . 5 2 x 1 0 . 0 0 6 6 5 . 2 0 PLOT 4 135 Sp e c i e s / L i c h e n x L i c h e n Number T o t a l L i c h e i Dominance Tree Biomass/ Biomass/ of Trees Biomass/ C l ass Number Tree (g) Cl a s s ( g ) i n C l a s s C l a s s (kg) HH-C 17 29 56.89 2851.20 24 68.43 18 3771.89 37 3200.09 53 987.89 56 2655.80 57 3778.69 62 2607.16 HH-I 3 174.73 663.88 12 7.9 7 6 181.94 47 1705. 13 63 593.70 WRC-C 5 459.65 459.65 3 1.38 HRC-I 49 508.09 50 8.09 3 1.52 DI-C 34 3737.75 2487.82 6 14.93 50 1237.88 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 (kg/ha) 94. 23 x 10.0 0 942.27 136 PLOT 5 Species/ Lichen x Lichen Number Total Lichen Dominance Tree Biomass/ Biomass/ of Trees Biomass/ Class Number Tree{g) Class(g) in Class Class (kg) HH-D 8 3495.76 4 080.52 2 8. 16 14 4665.28 HH-C 6 2520.13 2622.04 7 18.35 9 2825.71 11 3562.59 18 1176.86 29 3594.93 30 2052.33 HH-I 15 7.73 695.57 3 2.09 24 1383.40 WBC-C 17 309.30 30 9.30 1 .31 Af-C 1 354.63 435.04 3 1.31 4 146.73 5 803.75 Af-I 20 65.08 65.08 1 .07 Total l i c h e n biomass in plot (kg) 1/plot area (ha) Total lichen biomass (kg/ha) 30.29 x 10.0 0 302.90 Species/ Lichen Dominance Tree Bioniass/ Glass Number Tree(g) WH-D 30 6005.92 WH-C 56 3233. 23 92 2236.49 108 902.11 133 24 91.86 WH-I 5 58. 43 45 1071.46 83 1812.34 127 3677.34 WBC-C 58 825.37 WBC-I 104 1049.47 DF-D 54 3349.41 DF-C 15 1380.61 DF-I 31 51.78 MH-C 113 1230.67 137 PLOT 6 x Lichen Number Total Lichen Biomass/ of Trees Biomass/ Class <g) in Class Class (kg) 6005.92 2 12.01 2215.92 41 90.85 1654.89 30 49.65 825.37 10 8.25 1049.47 12 12.59 3349.41 4 13.40 1380.61 5 6.90 51.78 2 0.10 1230.67 1 1.23 Total lichen biomass in plot (kg) 1/plot area (ha) Total lichen biomass (kg/ha) 194.98 x 5.00 974.90 138 PLOT 7 Species/ Lichen x Lichen Number Total Lichen Dominance Tree Biomass/ Biomass/ of Trees Biomass/ Class Number Tree(g) Class (g) in Class Class (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 lichen biomass in plot (kg) 27.92 1/plot area (ha) x 5.00 Total l i c h e n biomass (kg/ha) 139.60 139 PLOT 8 Species/ Lichen x Lichen Number Total Lichen Dominance Tree Biomass/ Biomass/ of Trees Biomass/ Class Number Tree eg) Class (g) in Class Class (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 Total lichen biomass in plot (kg) 1/plot area (ha) Total lichen biomass (kg/ha) 57.74 x 5.00 288.71 140 PLOT 9 Species/ Lichen x Lichen Number Total Lichen Dominance Tree Biomass/ Biomass/ of Trees Biomass/ Class Number Tree(g) Class (g) i n Class Class (kg) WH-D WH-C WH-I WBC-C WBC-I DF-C DF-D DF-I 20 47 52 78 98 103 106 2 112 16 92 17 41 13 29 38 7238.69 1953.80 5503.25 3727.46 2263. 12 1232.15 3316.15 3801.73 529. 73 266.36 764.74 304.77 154. 18 4 59.28 13 27.93 2040.51 7238.69 3204.68 6 50 43.4 3 160.23 216 5.73 515.55 229.48 1275.91 (1275.91) 1275.913 30 13 38 20 3 2 64. 97 6.70 8.72 25.52 3.83 2.55 Total l i c h e n biotrass in plot (kg) 1/plot area (ha) Total lichen biomass (kg/ha) 315.95 x 2.53 800.46 141 PLOT 10 Species/ Lichen x Lichen Number Total Lichen Dominance Tree Biomass/ Biomass/ of Trees Biomass/ Class Number Tree (g) Class (g) in Class Glass (kg) BH-B 35 3672.86 3672.86 4 14.69 WH-C 9 1880*01 269 8. 36 17 45. 8 7 12 3567.05 13 2326.51 15 3885.22 17 2534.49 33 1996.86 WH-I 30 871. 77 730.27 9 6.57 43 773.24 47 545.81 WBC-C 26 225.37 225.37 3 0.68 HRC-I 42 106.98 106.98 8 1 0.86 DF-C 1 2321.14 1972.94 9 17.76 10 1858. 19 20 1739.49 GF-C 6 146.9 9 146.99 1 0.15 DF-D (1972.94) 2 3.95 DF-I 1972.943 1.97 Total lichen biomass i n plot (kg) 1/plot area (ha) Total lichen biomass (kg/ha) 92.50 x 5.6 3 521.20 142 PLOT 11 Species/ Lichen x Lichen Number Total Lichen Dominance Tree Biomass/ Biomass/ of Trees Biomass/ Class Number Tree(g) Class (g) i n Class Class (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 8. 14 Total lichen biomass i n plot (kg) 258.16 1/plot area (ha) x 5.87 Total lichen biomass (kg/ha) 1516.18 143 PLOT 12 Species/ Lichen x Lichen Number Total Lichen Dominance Tree Biomass/ Biomass/ of Trees Biomass/ Glass Number Tree(g) Class (gj in Class Class (Jig) WH-D 1 2033.86 2033.86 2 4. 07 WH-C 66 79 83 3,94 57.02 63. 90 41.62 9 0.0 4 WH-I 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 A f - I 11 45 50 53 0 2.05 0 0 17 0.01 Total lichen biomass i n plot (kg) 4. 17 1/plot area (ha) x 5.00 Total lichen biomass (kg/ha) 20.84 144 PLOT 13 Species/ Lichen x Lichen Number Total Lichen Dominance Tree Biomass/ Biomass/ of Trees Biomass/ Class Number Tree (g) Class (g) in Class Class (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 Total l i c h e n biomass i n plot (kg) 1/plot area (ha) Total lichen biomass (kg/ha) 21. 30 x 5.0 0 106.55 145 PLOT 14 Species/ Lichen x Lichen Number Total Lichen Dominance Tree Biomass/ Biomass/ of Trees Biomass/ Class Number Tree (g) Class (g) in Class Class (kg) HH-C 9 14 33 35 56 353.08 20 94.49 233.85 6409.88 1304.62 2079.18 11 22. 8 7 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 Total lichen biomass i n plot (kg) 1/plot area (ha) Total lichen biomass (kg/ha) 35.20 x 5.00 175. 95 146 APPENDIX I I D i f f e r e n t i a t e d Table of Vegetation D a t a 1 (Hueller-Dombois and E l l e n b e r g 1974: 177-193) 1 The f i r s t number i n each box r e p r e s e n t s an estimate of percent cover. The second number re 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 only one number i s present, i t r e p r e s e n t s percent cover. Vegetation Community AF/OH Plot number 14 8 15 5 Elevation (m) 603 707 695 573 Aspect N35W S05W N15W N75E Slope (degrees) 24 5 16 23 Abies amabilis 25 15 25 10 Covnus canadensis 2/4 ' 1/5 1/4 3/6 Rubus pedatus '2/6 .1/5 1/4 1/5 Vaccinium ovalifolium 10/5 5/5 15/6 1/2 Menziesia fevruginea 3/4 3/4 1/2 Listera cauvina 1/2 1/2 1/1 Clintonia uniflora 1/4 1/5 Streptopus roseus 1/5 1/5 1/2 Plagiothecium: undulgtum 5/5 1/3 Tsuga mevtensiana 7 2 2 Chamaecyparis nootkatensis 2 5 2/5 Sphagnum sp. 1/5 Khizomnium sp. 1/5 2/5 Coptis asplenifolium 2/6 1/5 Gymnocarpium dryopteris 1/5 2/6 Pseudotsuga menziesii Gaulthevia shallon Eemitomes congestum Stokesiella ovegana Viola sempevvirens Boschniakia hookevi Tsuga heterophylla 35 35 45 60 Listera covdata 1/2 1/2 1/2 1/4 Hylocomium splendens 20/6 15/6 5/5 15/6 Rhytidiadelphus loreus 50/8 15/6 50/8 35/8 Thuja plicata 15 5 2 Vaccinium parvifolium 5/4 1/4 3/4 TRANS DF/S UNCLASSIFIED 6 9 11 13 10 1 3 12 4 7 2 669 646 792 426 469 686 543 487 709 282 442 S60E N82W S60E N87W S10W S15E S30E S22W S60E N80W S40E 30 32 14 25 20 32 54 32 22 20 15 1 1 1 25 1 1 1/1 2/7 1/3 1/3 1/6 1/3 1/6 1/5 1/5 25/6 40/8 1/1 1/2 1/2 1/1 1/1 1/4 1/1 1/1 1/3 1/5 2 1 7 20 20 35 35 7 55 8 5 20 70/7 1/5 5/6 1/3 90/8 40/8 80/8 20/6 1/5 3/3 1/5 1/2 1/2 1/1 1/1 1/2 1/5 3/5 1/1 •"•1/1 3/4 1/3 1/5 1/2 1/3 1/1 1/2 1/1 1/4 55 50 60 40 45 30 9 25 70 80 35 1/4 1/4 2/7 1/2 : i / i 1/4 1/4 1/4 1/6 1/2 3/7 70/8 30/6 20/8 40/8 9/6 30/8 50/8 10/6 50/8 70/8 30/8 10/6 10/5 20/8 20/6 1/4 20/7 5/7 10/6 10/7 10/7 30/8 2 15 5 15 .5 15 7 15 5 4 35 3/6 10/4 5/4 3/5 2/4 15/7 12/7 2/4 15/7 4/7 3/2 Appendix II Continued Vegetation Community AF/OH TRANS DF/S UNCLASSIFIED Vaeciniwn alaskaense 50/6 75/8 40/8 25/8 2/5 35/7 40/8 2/5 1/1 10/6 1/1 2/4 40/7 2/7 Chimaphila menziesii 1/2 1/2 1/2 1/2 1/2 1/2 1/4 1/1 1/5 1/4 1/2 1/2 1/2 Rhytidiopsis robusta 2/5 30/8 20/6 35/8 25/6 40/6 5/6 2/5 15/7 1/2 30/7 2/5 5/7 C o r a l l o r h i z a mevtensiana 1/5 1/5 1/5 1/5 1/5 1/5 1/5 1/2 1 / 2 1 / 2 1 / 2 Goodyera o b l o n g i f o l i a 1/2 1/5 1/2 1/2 1/2 1 / 5 1 / 2 1 / 1 1 / 4 1 / 2 3/2 Hypopitys monotvopa 1/5 1/5 1/2 1/2 1/2 1/2 1/1 1/2 1/2 1/4 1/2 Bleohnum spioant 1/5 1/3 1/5 1/2 1/3 1/2 1/2 Aehlys triphylla 5/6 1/3 1/3 1/3 1/5 3/2 Pinus monticola 1 1 1 1 1 1 Moneses u n i f l o r a 1/2 1/4 1/1 1/3 1/2 Linnaea borealis 1 / 5 1 / 5 1 / 4 1/4 1/3 T i a r e l l a tvifoliata 2/4 1/5 3/3 Laotuaa muralis 1/11/2 3/2 Polystichum lonehitzs 1/1 1/5 1/5 Berberis nervosa 1/3 1/2 1/3 Polystichum. munitum 1/1 2/5 1/3 Athyrium f i l i x - f e m i n a 1/6 1/1 T i a r e l l a trifoliata var. 1/2 1/3 laainiata Chimaphila umbellata 1/6 1/6 Dryopteris austriaca 1/1 1/1 Diaranum sp. 1/5 1/5 Abies grandis 1 Rosa gymnooarpa 1/5 Sambucus raeemosa 1/1 AIlotropa virgata 1/3 Maianthemum dilatatum 1/1 Bromus oarinatus 1/5 Lysiohitum americanum 1/5 Epilobium angustifolium 1/1 Veratrum v i r i d e 1/5 Viola g l a b e l l a 1/5 Plagiomnium sp. 1/5 Monotropa uniflorg. 1/5 

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