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Dispersion of the western winter wren (Troglodytes troglodytes pacificus [Baird]) in coastal western… McLachlin, Roderick Archibald 1983

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DISPERSION OF THE WESTERN WINTER WREN (TROGLODYTES  TROGLODYTES PACIFICUS [BAIRD]) IN COASTAL WESTERN HEMLOCK FOREST AT THE UNIVERSITY OF BRITISH COLUMBIA RESEARCH FOREST IN SOUTH-WESTERN BRITISH COLUMBIA by RODERICK ARCHIBALD MCLACHLIN B.A., U n i v e r s i t y of Western O n t a r i o , 1964 M.Sc, U n i v e r s i t y of A l b e r t a , 1970 A THESIS SUBMITTED IN PARTIAL FULFILMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY in THE FACULTY OF GRADUATE STUDIES (Department of F o r e s t r y ) we accept t h i s t h e s i s as conforming to the r e q u i r e d standard THE UNIVERSITY OF BRITISH COLUMBIA A p r i l 1983 © R.A. McLachlin, 1983 In presenting t h i s thesis i n p a r t i a l f u l f i l m e n t of the requirements for an advanced degree at the University of B r i t i s h Columbia, I agree that the Library s h a l l make i t f r e e l y available for reference and study. I further agree that permission for extensive copying of t h i s thesis for scholarly purposes may be granted by the head of my department or by h i s or her representatives. I t i s understood that copying or publication of t h i s thesis for f i n a n c i a l gain s h a l l not be allowed without my written permission. Department of The University of B r i t i s h Columbia 1956 Main Mall Vancouver, Canada V6T 1Y3 DE-6 (3/81) ABSTRACT I s t u d i e d the d i s p e r s i o n of winter wrens i n 100-year-o l d , second-growth, c o a s t a L western hemlock f o r e s t at the U n i v e r s i t y of B r i t i s h Columbia Research F o r e s t i n southwestern B r i t i s h Columbia from 1978-81. Male winter wrens were t e r r i t o r i a l on non-overlapping t e r r i t o r i e s at an average d e n s i t y of 60 per km2. An average of 8% were polygamous. Females occupied g e n e r a l l y non-overlapping home ranges at l e a s t d u r i n g the breeding p e r i o d , but were not shown t e r r i t o r i a l , although t h i s p o s s i b i l i t y c o u l d not be excluded. Winter wrens were not u n i f o r m l y d i s t r i b u t e d but showed d i f f e r e n t i a l use of v a r i o u s i n d i v i d u a l ecosystems (as mapped by K l i n k a 1976) and ecosystems grouped by f o r e s t f l o o r h a b i t a t s . S u r p l u s , p o t e n t i a l l y t e r r i t o r i a l males were a v a i l a b l e d u r i n g the breeding p e r i o d which c o u l d have occupied the empty or s p a r s e l y occupied a r e a s . I n v e r t e b r a t e food was more abundant i n h a b i t a t s used by winter wrens as compared to avoided h a b i t a t s , and, food i s proposed as a f a c t o r i n h a b i t a t s e l e c t i o n by winter wrens. I propose that winter wrens are spaced by t e r r i t o r i a l i t y and clumped by s u i t a b l e h a b i t a t , and suggest that these two f a c t o r s i n f l u e n c e the p a t t e r n s of d i s p e r s i o n of winter wrens i n c o a s t a l western hemlock f o r e s t , and perhaps elsewhere as w e l l . K l i n k a ' s ecosystems and grouped ecosystems were proposed as i n d i c a t i v e of the d i s t r i b u t i o n of winter wrens, and perhaps of other w i l d l i f e s p e c i e s g e n e r a l l y . I f so, ecosystems can arid should be used as the base f o r the study and management of w i l d l i f e i n the p r o v i n c e of B r i t i s h Columbia, and perhaps elsewhere as w e l l . i v TABLE ABSTRACT TABLE OF CONTENTS LIST OF TABLES ... LIST OF FIGURES . . LIST OF APPENDICES ACKNOWLEDGEMENTS . INTRODUCTION 1 THE STUDY AREAS AND THE HANDLING OF WINTER WRENS• • 6 Study Areas 6 Study area 6 Study p l o t s 13 Removal p l o t s 14 Catc h i n g , Banding, and C o l l e c t i n g Winter Wrens . 14 Sex Determination of Winter Wrens 17 Age Determination of Winter Wrens 27 WINTER WREN DISPERSION AS A FUNCTION OF TERRITORIALITY 33 I n t r o d u c t i o n 33 Methods 34 Winter Wren T e r r i t o r i a l i t y 39 OF CONTENTS Page i i i v v i i x i v x x i x x i i V TABLE OF CONTENTS (continued) Page Males 39 Females 69 D i s t r i b u t i o n of Winter Wrens 82 Removal Experiments 87 WINTER WREN DISPERSION AS A FUNCTION OF HABITAT 9 5 Winter Wren D i s p e r s i o n as a F u n c t i o n of K l i n k a ' s Ecosystems and Ecosystems Grouped by F o r e s t F l o o r H a b i t a t s 9 5 I n t r o d u c t i o n 0,5 Methods -LOl D e s c r i p t i o n and mapping of ecosystems and ecosystems grouped by f o r e s t f l o o r h a b i t a t s . . . 109 Study area and study p l o t s 109 Removal p l o t s 136 D i s t r i b u t i o n s of male t e r r i t o r i e s and female home ranges among the ecosystems and ecosystems grouped by f o r e s t f l o o r h a b i t a t s 160 D i s t r i b u t i o n of winter wren o b s e r v a t i o n s among the ecosystems and ecosystems grouped by f o r e s t f l o o r h a b i t a t s 169 Winter Wren D i s p e r s i o n as a F u n c t i o n of Food 186 I n t r o d u c t i o n 186 v i TABLE OF CONTENTS (continued) Page Methods 187 Winter wren food 193 P o t e n t i a l winter wren food w i t h i n the ecosystems grouped by f o r e s t f l o o r h a b i t a t s 199 CONCLUSIONS 212 REFERENCES 217 APPENDICES 253 v i i LIST OF TABLES No. P a 9 " e 1 Methods of capture and c o l l e c t i o n of winter wrens 15 2 Summary of the g e n e r a l m o r p h o l o g i c a l , b e h a v i o u r a l , and anatomical c r i t e r i a f o r winter wren sex d e t e r m i n a t i o n 18 3 Data summary and s t a t i s t i c a l comparisons of "known" male and female winter wren weights, l e f t f e a t h e r measurements, l e f t wing l e n g t h s , and c o l o u r s . 22 4 Winter wren sex d i s c r i m i n a t i o n c r i t e r i a based on weights, l e f t f e a t h e r measurements, l e f t wing l e n g t h s , and c o l o u r s 24 5 Average weights and f i r s t primary and c e n t r a l r e c t r i x l e ngths from broods of winter wrens of known dates of h a t c h i n g 30 6 Numbers of t e r r i t o r i a l males and male t e r r i t o r i e s on the study area, study p l o t s , and v i i i LIST OF TABLES (continued) Page removal p l o t s 84 7 Removal and replacement of t e r r i t o r i a l males on and around the removal p l o t s D and E d u r i n g the breeding p e r i o d s 1979 and 1980 88 8 R e l a t i o n s h i p s among the taxa of the Linnaean, Canadian s o i l , v e g e t a t i o n , and s y n e c o l o g i c a l c l a s s i f i c a t i o n systems. 97 9 Abridged v e g e t a t i o n p l o t data of the K l i n k a s y n e c o l o g i c a l map u n i t s 1 1 2 10 Areas (m 2) and percentages of the ecosystems on the study area and study p l o t s 1 2 3 11 Percent occurrences of the f o r e s t f l o o r h a b i t a t s on the ecosystems on the study area and the r e s u l t i n g assignments of the ecosystems by f o r e s t f l o o r h a b i t a t s . 1 4 1 12 Areas (m 2) and percentages of the f o r e s t f l o o r h a b i t a t s on the study area and study p l o t s as c a l c u l a t e d from the grouped ecosystems as shown in Table 11 1 4 2 i x LIST OF TABLES (continued) Page 13 Abridged v e g e t a t i o n p l o t data f o r the f o r e s t f l o o r h a b i t a t s 143 14 Areas (m 2) and percentages of the ecosystems on the removal p l o t s D and E 152 15 Percent occurrences of the f o r e s t f l o o r h a b i t a t s on the ecosystems on the removal p l o t s and the r e s u l t i n g assignments of the ecosystems to f o r e s t f l o o r h a b i t a t s 159 16 S t a t i s t i c a l comparisons (X 2) of the expected and observed d i s t r i b u t i o n s of the areas (m 2) of male t e r r i t o r i e s among the ecosystems and ecosystems grouped by f o r e s t f l o o r h a b i t a t s d u r i n g the v a r i o u s census p e r i o d s of the study area, 1978-81 162 17 S t a t i s t i c a l comparisons ( X 2 ) of the expected and observed d i s t r i b u t i o n s of the areas (m 2) of male t e r r i t o r i e s and female home ranges among the ecosystems and ecosystems grouped by f o r e s t f l o o r h a b i t a t s d u r i n g the v a r i o u s census p e r i o d s of the study p l o t s , 1 979-80 163 X LIST OF TABLES (continued) Page 18 S t a t i s t i c a l comparisons (X 2) of the expected and observed d i s t r i b u t i o n s of the areas (m 2) of male t e r r i t o r i e s among the ecosystems f o r the f i r s t censuses on removal p l o t E, 1979-80 164 19 Numbers of census p e r i o d s i n which the observed areas (m 2) of male t e r r i t o r i e s and female home ranges w i t h i n the v a r i o u s ecosystems were g r e a t e r than the expected v a l u e s 165 20 Numbers of census p e r i o d s i n which the observed areas (m 2) of male t e r r i t o r i e s and female home ranges w i t h i n the v a r i o u s ecosystems grouped by f o r e s t f l o o r h a b i t a t s were g r e a t e r than the expected v a l u e s . 166 21 S t a t i s t i c a l comparisons (X 2) of the expected and observed d i s t r i b u t i o n s of the numbers of winter wren o b s e r v a t i o n s among the ecosystems and ecosystems grouped by f o r e s t f l o o r h a b i t a t s d u r i n g the v a r i o u s census p e r i o d s of the study a r e a , 1978-81 172 22 S t a t i s t i c a l comparisons (X 2) of the expected and observed d i s t r i b u t i o n s of the numbers of x i LIST OF TABLES (continued) Page winter wren o b s e r v a t i o n s among the ecosystems and ecosystems grouped by f o r e s t f l o o r h a b i t a t s d u r i n g the v a r i o u s census p e r i o d s of the study p l o t s , 1979-80 173 23 S t a t i s t i c a l comparisons ( X 2 ) of the expected and observed d i s t r i b u t i o n s of the numbers of winter wren o b s e r v a t i o n s among the ecosystems d u r i n g a l l censuses of removal p l o t E, 1979-80... 174 24 Numbers of census p e r i o d s i n which the observed numbers of o b s e r v a t i o n s of winter wrens w i t h i n the v a r i o u s ecosystems were g r e a t e r than the expected values 175 25 Numbers of census p e r i o d s i n which the observed numbers of o b s e r v a t i o n s of winter wrens w i t h i n the v a r i o u s ecosystems grouped by f o r e s t f l o o r h a b i t a t s were g r e a t e r than the expected v a l u e s . . . 176 26 Summary of winter wren p r e f e r e n c e s f o r ecosystems and ecosystems grouped by f o r e s t f l o o r h a b i t a t s 181 27 Summary of the winter wren stomach contents 194 X I 1 LIST OF TABLES (continued) Page 28 Abridged v e g e t a t i o n data of the f o r e s t f l o o r h a b i t a t s 200 29 Summary (means and ranges) of the i n v e r t e b r a t e s e x t r a c t e d from the i n v e r t e b r a t e p l o t samples from the v a r i o u s f o r e s t f l o o r h a b i t a t s d u r i n g the breeding p e r i o d 202 30 Summary (means and ranges) of the i n v e r t e b r a t e s e x t r a c t e d from the i n v e r t e b r a t e p l o t samples from the v a r i o u s f o r e s t f l o o r h a b i t a t s d u r i n g the winter p e r i o d .... 204 31 Rankings and s t a t i s t i c a l comparisons (Spearman's c o e f f i c i e n t of rank c o r r e l a t i o n , r s ) of the mean f r e q u e n c i e s (% occurrences) of s e l e c t e d i n v e r t e b r a t e taxa from the winter wren stomachs, with the products of the mean fr e q u e n c i e s X the mean volumes of the i n v e r t e b r a t e taxa as e x t r a c t e d from the i n v e r t e b r a t e p l o t samples from the v a r i o u s f o r e s t f l o o r h a b i t a t s d u r i n g the breeding p e r i o d 207 32 Rankings and s t a t i s t i c a l comparisons x i i i LIST OF TABLES (continued) Page (Spearman's c o e f f i c i e n t of rank c o r r e l a t i o n , r g ) of the mean f r e q u e n c i e s (% occurrences) of s e l e c t e d i n v e r t e b r a t e taxa from the winter wren stomachs, with the products of the mean fr e q u e n c i e s X the mean volumes of the i n v e r t e b r a t e taxa as e x t r a c t e d from the i n v e r t e b r a t e p l o t samples from the v a r i o u s f o r e s t f l o o r h a b i t a t s d u r i n g the winter p e r i o d . . . 208 33 S t a t i s t i c a l comparisons (Duncan's m u l t i p l e range t e s t ) of the means of the t o t a l volumes (ml) of i n v e r t e b r a t e s from the i n v e r t e b r a t e p l o t samples of the v a r i o u s f o r e s t f l o o r h a b i t a t s d u r i n g the breeding and winter x i v LIST OF FIGURES No. N Page 1 L o c a t i o n map of the U n i v e r s i t y of B r i t i s h Columbia Research F o r e s t i n south-western B r i t i s h Columbia 7 2 L o c a t i o n map of the study area on the U n i v e r s i t y of B r i t i s h Columbia Research F o r e s t . . 9 3 A i r photograph showing the l o c a t i o n of the study area 11 4 Area and extent of the male t e r r i t o r i e s on the study area d u r i n g the breeding p e r i o d , 1978 41 5 Area and extent of the male t e r r i t o r i e s on the study area d u r i n g the breeding p e r i o d , 1979 43 6 Area and extent of the male t e r r i t o r i e s on the study area d u r i n g the breeding p e r i o d , 1980 45 7 Area and extent of the male t e r r i t o r i e s on the study area d u r i n g the breeding p e r i o d , 1981 47 X V LIST OF FIGURES (continued) Page 8 Area and extent of the male t e r r i t o r i e s on the study plots during the winter period, 1979. 4 9 9 Area and extent of the male t e r r i t o r i e s on the study plots during the breeding period, 1979. •• 51 10 Area and extent of the male t e r r i t o r i e s on the study plots during the f a l l period, 1979 53 11 Area and extent of the male t e r r i t o r i e s on the study plots during the winter period, 1979-80... 55 12 Area and extent of the male t e r r i t o r i e s on the study plots during the breeding period, 1980. .. 57 13 Area and extent of the male t e r r i t o r i e s on the removal plots during the early breeding periods, 1979 and 1980. 59 14 Area and extent of the female home ranges on the study plots during the breeding period, 1979 71 1 5 Area and extent of the female home ranges on the study plots during the f a l l period, 1979.... 73 xv i LIST OF FIGURES (continued) Page 16 Area and extent of the female home ranges on the study p l o t s d u r i n g the winter p e r i o d , 1979/80 75 17 Area and extent of the female home ranges on the study p l o t s d u r i n g the breeding p e r i o d , 1980 77 18 Remapped s y n e c o l o g i c a l map of the study area i n c l u d i n g the study p l o t s A, B, and C I l l 19 Schematic diagram of the r e l a t i v e l o c a t i o n s of s e l e c t e d ecosystems on the sl o p e s of the study area . 122 20 (LICHEN) - GAULTHERIA - DF, 11.30, (study a r e a ) : v e r t i c a l mosaic 125 21 GAULTHERIA - WH - DF, 131.30/131.4, (study a r e a ) : v e r t i c a l mosaic 125 22 Mahonia - G a u l t h e r i a - WH - DF, 132.9, (study a r e a ) : v e r t i c a l mosaic 125 23 (LICHEN) - GAULTHERIA - DF, 11.30, (study x v i i LIST OF FIGURES (continued) Page a r e a ) : f o r e s t f l o o r 127 24 GAULTHERIA - WH - DF, 131.30/131.4, (study a r e a ) : f o r e s t f l o o r . . . . . . 127 25 Mahonia - G a u l t h e r i a - WH - DF, 132.9, (study a r e a ) : f o r e s t f l o o r 127 26 Moss - WH, 211.4, (study a r e a ) : v e r t i c a l mosaic 129 27 MOSS - (POLYSTICHUM) - WH - WRC, 31.412, (study a r e a ) : v e r t i c a l mosaic 129 28 POLYPODIUM - GAULTHERIA - DF - WRC, 62.90, (study a r e a ) : v e r t i c a l mosaic 129 29 Moss - WH, 211.4, (study a r e a ) : f o r e s t f l o o r . . . . 131 30 MOSS - (POLYSTICHUM) - WH - WRC, 31.412, (study a r e a ) : f o r e s t f l o o r 131 31 POLYPODIUM - GAULTHERIA - DF - WRC, 62.90, (study a r e a ) : f o r e s t f l o o r 131 x v i i i LIST OF FIGURES (continued) Page 32 TIARELLA - POLYSTICHUM - WRC, 71.4/71.89, (study a r e a ) : v e r t i c a l mosaic 133 33 RUBUS - POLYSTICHUM - WRC, 72.4/72.47, (study a r e a ) : v e r t i c a l mosaic 133 34 Ribes - Oplopanax - WRC, 812.89, (study a r e a ) : v e r t i c a l mosaic. 133 35 TIARELLA - POLYSTICHUM - WRC, 71.4/71.89, (study a r e a ) : f o r e s t f l o o r 135 36 RUBUS - POLYSTICHUM - WRC, 72.4/72.47, (study a r e a ) : f o r e s t f l o o r . 135 37 Ribes - Oplopanax - WRC, 812.89, (study a r e a ) : f o r e s t f l o o r 135 38 Vaccinium - L y s i c h i t u m - WRC, 911.0, (study a r e a ) : v e r t i c a l mosaic 138 39 TIARELLA - POLYSTICHUM - WRC, 71.12/71.4/71.89, (removal p l o t D): v e r t i c a l mosaic 138 40 Moss - WH, 211.23, (removal p l o t E ) : v e r t i c a l xix LIST OF FIGURES (continued) Page mosaic 138 41 Vaccinium - L y s i c h i t u m - WRC, 911.0, (study a r e a ) : f o r e s t f l o o r 140 4 2 TIARELLA - POLYSTICHUM - WRC, 71.12/71.4/71.89, (removal p l o t D): f o r e s t f l o o r 140 43 Moss - WH, 211.23, (removal p l o t E ) : f o r e s t f l o o r 140 44 Remapped s y n e c o l o g i c a l map of the removal p l o t s D and E. ; 149 45 Schematic diagram of the r e l a t i v e l o c a t i o n s of the ecosystems on the s l o p e s of the removal p l o t s D and E 151 46 Mahonia - Moss - WRC - WH, 212.89, (removal p l o t E ) : v e r t i c a l mosaic 155 47 MOSS - (POLYSTICHUM) - WH - WRC, 31.12, (removal p l o t E ) : v e r t i c a l mosaic. 155 48 Mahonia - Moss - WRC - WH, 212.89, (removal X X LIST OF FIGURES (continued) Page p l o t E ) : f o r e s t f l o o r 157 49 MOSS - (POLYSTICHUM) - WH - WRC, 31.12, (removal p l o t E ) : f o r e s t f l o o r 157 50 M o d i f i e d B e r l e s e type f u n n e l s used to e x t r a c t i n v e r t e b r a t e s from the samples of f o r e s t f l o o r v e g e t a t i o n and s u r f a c e l i t t e r . 190 x x i LIST OF APPENDICES No. Page .1 S y n o p s i s of the s y n e c o l o g i c a l c l a s s i f i c a t i o n of the c o a s t a l w e s t e r n hemlock f o r e s t of t h e U n i v e r s i t y of B r i t i s h Columbia R e s e a r c h F o r e s t . . . . 254 2 Summary of the K l i n k a s y n e c o l o g i c a l u n i t s of the study a r e a , t h e i r c l a s s i f i c a t i o n , p a r e n t m a t e r i a l s , n u t r i e n t and m o i s t u r e regimes, and a s s o c i a t e d s o i l s 258 3 L i s t of t h e p l a n t s p e c i e s found on the v e g e t a t i o n and f o r e s t f l o o r sample p l o t s 263 4 L i s t of the i n v e r t e b r a t e t a x a i d e n t i f i e d from the i n v e r t e b r a t e sample p l o t s and the w i n t e r wren stomachs 271 x x i i ACKNOWLEDGEMENTS The committee was as f o l l o w s : Dr. J.V. Thirgood -chairman, Dr. F.L. Bunnell - t h e s i s a d v i s o r , and Dr. C.L. Gass, Dr. J.P. Kimmins, and Dr. J.N.M. Smith -committee members. I was a s s i s t e d i n the f i e l d by Ms. P. W a l l i s and Ms. B. Peterson, and Messrs. B. MacDonald, F. Simpson, and P. Henderson. Mr. MacDonald a l s o a s s i s t e d i n the s o r t i n g and i d e n t i f i c a t i o n of the i n v e r t e b r a t e s . The f i e l d r e s e a r c h was c a r r i e d out at the U n i v e r s i t y of B r i t i s h Columbia Research F o r e s t . L o g i s t i c a l support there was k i n d l y p r o v i d e d by Mr. P.R.W. Sanders, and Ms. J . Jensen. Support d u r i n g the study was p r o v i d e d by the f o l l o w i n g : a K a i s e r Resources L t d . F e l l o w s h i p , an H.R. MacMillan Family F e l l o w s h i p , the Donald S. McPhee F e l l o w s h i p Fund, and a U n i v e r s i t y of B r i t i s h Columbia Graduate Summer S c h o l a r s h i p . The t h e s i s was typed by Ms. P. M i l l s , and Ms. C. van den D r i e s e n . The f i g u r e s were prepared by Ms. L. Koza, and Messrs. F. Yanetta, and D. Waddell. 1 INTRODUCTION Wynne-Edwards (1962, p. l ) d e f i n e d animal d i s p e r s i o n as f o l l o w s : "the placement of i n d i v i d u a l s and groups of i n d i v i d u a l s w i t h i n the h a b i t a t s they occupy and the pr o c e s s e s by which t h i s i s brought about." I examined the phenomenon of d i s p e r s i o n i n a p o p u l a t i o n of western winter wrens, T r o g l o d y t e s t r o g l o d y t e s p a c i f i c u s ( B a i r d ) , by a s k i n g two g e n e r a l q u e s t i o n s : 1) How are the i n d i v i d u a l s and groups of i n d i v i d u a l s d i s t r i b u t e d ? ; and 2) Why are they d i s t r i b u t e d that way? The c e n t r a l i n t e r e s t of t h i s study i s the h y p o t h e s i s that ecosystems and grouped ecosystems (sensu K l i n k a 1976) are i n d i c a t i v e of the d i s t r i b u t i o n of winter wrens. The ecosystems as recognized and mapped by K l i n k a were developed u s i n g the c r i t e r i a of climate, s o i l , and v e g e t a t i o n (Brooke et a l . 1970; Kojima and K r a j i n a 1975; K l i n k a 1976; K l i n k a and Skoda 1977) and i t has always been assumed t h a t the ecosystems a l s o r e f l e c t e d the d i s t r i b u t i o n of animals (K. K l i n k a , p e r s . comm.). To my knowledge, t h i s study of winter wrens i s the f i r s t attempt to t e s t the u t i l i t y of K l i n k a ' s ecosystems u s i n g animals. 2 In the course of my i n v e s t i g a t i o n of t h i s c e n t r a l h y p o t h e s i s , I proposed the f o l l o w i n g q u e s t i o n s : 1) are winter wrens t e r r i t o r i a l , and spaced by t e r r i t o r i a l i t y ? ; 2) are winter wrens u n i f o r m l y d i s t r i b u t e d over the study area?; 3) i f not, are s u r p l u s , p o t e n t i a l l y t e r r i t o r i a l males a v a i l a b l e which c o u l d occupy the empty or s p a r s e l y occupied a r e a s ? ; 4) do winter wrens show d i f f e r e n t i a l use of the v a r i o u s i n d i v i d u a l ecosystems (p. 96) and ecosystems grouped by f o r e s t f l o o r v e g e t a t i o n (p. 105), and i f so, are they clumped by s u i t a b l e h a b i t a t ? ; and 5) i s the abundance of i n v e r t e b r a t e food a f a c t o r i n the d i s p e r s i o n of winter wrens. The n a t u r a l h i s t o r y of the European wren ( T . t . t r o g l o d y t e s ) i s reasonably w e l l documented. Some of the e a r l i e s t s c i e n t i f i c o b s e r v a t i o n s were made i n I r e l a n d by B u r k i t t (1919, 1920) and i n England by B u t t e r f i e l d (1920a, b ) . The f i r s t long term study was th a t of K l u i j v e r et a l . (1940) i n the Netherlands. T h e i r work was fo l l o w e d by Armstrong's s t u d i e s i n England (Armstrong 1944, 1953a, 1955, 1956, 1963; Whitehouse and Armstrong 1953; Armstrong and Whitehouse 1977). More recent s t u d i e s i n England i n c l u d e the s t u d i e s of Williamson (1969a) and Hawthorn and others (Hawthorn 1971, 1974, 1975; Hawthorn et a l . 1971; Hawthorn and Mead 1975). In a d d i t i o n , 3 K e n t i s h (1976) examined s p r i n g and summer home ranges and t e r r i t o r i e s , and Garson (1978) s t u d i e d t e r r i t o r i a l and breeding behaviour (Garson and Hunter 1979; Garson 1980a, b) . On the Continent, Pokrovskaya (1968) and Zimin (1972) s t u d i e d n e s t i n g b i o l o g y i n e a s t e r n Europe while Dallman (1977) r e p o r t e d on breeding i n West Germany. Bremond (1968, 1978) and Kreutzer (1972, 1974a, b) s t u d i e d wren -song i n France and Wesolowski (1981) conducted removal experiments i n Poland. The n a t u r a l h i s t o r y of the v a r i o u s n o r t h A t l a n t i c i s l a n d wren subspecies has a l s o been s t u d i e d . The best known i s the S t . K i l d a wren ( T . t . h i r t e n s i s ) of which the e a r l i e s t s t u d i e s were made by C l a r k e (1915) f o l l o w e d by the more d e t a i l e d s t u d i e s of H a r r i s s o n and Buchan (1934, 1936), Armstrong (1953b, 1959), Bagenal (1958), W i l l i a m s o n (1958), and Waters (1964). The other n o r t h A t l a n t i c wren subspecies have r e c e i v e d l e s s a t t e n t i o n but are a l s o r e l a t i v e l y w e l l known: Faeroe wren (T.t:. b o r e a l i s ) (Williamson 1947); I c e l a n d wren ( T . t . i s l a n d i c u s ) (Armstrong 1950); F a i r I s l e wren ( T . t . f r i q i d a r i e n s i s ) (Williamson 1951 , 1958); Shetland wren (T._t. z e t l a n d i c u s ) (Armstrong 1952; Armstrong and Thorpe 1952), and the Hebridean wren ( T . t . h e b r i d e n s i s ) (Armstrong 1953c; 4 Hawthorn et a l . 1976). More r e c e n t l y , Cody and Cody (1972a, b) compared t e r r i t o r y , c l u t c h s i z e , and food supply among E n g l i s h mainland and n o r t h A t l a n t i c i s l a n d wren su b s p e c i e s . Elsewhere i n the Old World, Haneda and Kosakai (1971) r e p o r t e d on the breeding b i o l o g y of the Japanese wren ( T . t . fumigatus). Regarding my q u e s t i o n s on d i s p e r s i o n , O l d World wrens are t e r r i t o r i a l and are a p p a r e n t l y spaced by t e r r i t o r i a l behaviour (e.g., K l u i j v e r et a l . 1940; Armstrong and Whitehouse 1977; Garson 1978). They are a l s o a p p a r e n t l y clumped by s u i t a b l e h a b i t a t (e.g., Williamson 1969a, b ) . S u r p l u s , p o t e n t i a l l y t e r r i t o r i a l males have been observed d u r i n g the breeding p e r i o d (Wesolowski 1 9 8 1 ) . The abundance of i n v e r t e b r a t e food has been suggested as a f a c t o r i n wren d i s t r i b u t i o n (Cody and Cody 1972a, b ) . However, to my knowledge, ecosystems have not been shown to be i n d i c a t i v e of wren d i s t r i b u t i o n . The New World winter wren i s much l e s s w e l l known than i t s O ld World c o u n t e r p a r t . I t has been customary to s u b s t i t u t e the b i o l o g y of the European wren f o r that of the winter wren (Verner and W i l l s o n 1966; Kroodsma 1977, 1980). However, a b r i e f , s p e c i f i c l i t e r a t u r e on the New World winter wren e x i s t s . Kroodsma (1980) s t u d i e d winter 5 wren s i n g i n g behaviour i n Maine, New York, and Oregon. There are a l s o e a r l y o b s e r v a t i o n s on: the n a t u r a l h i s t o r y of the Alaska winter wren ( T . t . a l a s k e n s i s ) (Heath 1920; P r e b l e and McAtee 1923); decoy n e s t s of the western winter wren (Bowles 1899); western winter wren d i s p l a y (Pearse 1933); and western winter wren w i n t e r - r o o s t i n g (Ehinger 1925) . As t h i s study was, to my knowledge, the f i r s t g e n e r a l study of the winter wren i n North America, I a l s o found i t necessary to work out methods f o r capture, and de t e r m i n a t i o n of sex and age of winter wrens, as w e l l as to develop some understanding of t h e i r n a t u r a l h i s t o r y . 6 THE STUDY AREAS AND THE HANDLING OF WINTER WRENS Study Areas I d i d my study at the U n i v e r s i t y of B r i t i s h Columbia Research F o r e s t (49°N, 122°W). The 5,151 ha Research Fore s t i s l o c a t e d i n south-western B r i t i s h Columbia on the north s l o p e of the F r a s e r River V a l l e y approximately s i x km nor t h of the town of Haney and 50 km east of the c i t y of Vancouver ( F i g u r e 1). The study area was l o c a t e d on the southern or lower end of the Research F o r e s t . The study area c o n t a i n e d three study p l o t s A, B, and C. In a d d i t i o n there were two removal p l o t s D and E remote from the study area ( F i g u r e s 2 and 3). Study Area The roughly t r i a n g u l a r 38.7 ha study area was l o c a t e d on moderately to s t e e p l y s l o p i n g h i l l s i d e and ranged from 55 to 180 m i n e l e v a t i o n . The e a s t e r n base of the t r i a n g l e was about 900 m long and the east-west a x i s was about 625 m (F i g u r e 3 ) . The study area was f o r e s t e d with unmanaged, late-immature, second-growth, c o a s t a l western hemlock f o r e s t . Old f o r e s t was necessary f o r t h i s study, as o n l y i n such f o r e s t s does the understory v e g e t a t i o n of the v a r i o u s f o r e s t ecosystems reach f u l l e x p r e s s i o n . With FIGURE 1 LOCATION MAP OF THE UNIVERSITY OF BRITISH COLUMBIA RESEARCH FOREST IN SOUTH-WESTERN BRITISH COLUMBIA (South Western B r i t i s h Columbia 1979) Oo f J ^ *Wi. fBaker I ty • T.|H gKS.N F I G U R E 2 L O C A T I O N MAP OF STUDY AREA ON THE U N I V E R S I T Y OF B R I T I S H COLUMBIA R E S E A R C H F O R E S T ( C o q u i t l a m , B r i t i s h C o l u m b i a 1 9 7 6 ) FIGURE 3 AIR PHOTOGRAPH SHOWING THE LOCATION OF THE STUDY AREA ( U n i v e r s i t y of B r i t i s h Columbia Research F o r e s t 1979) (Scale: approximately 1:12,000) 13 the e x c e p t i o n of a small t r i a n g u l a r - s h a p e d area of s i m i l a r f o r e s t to the south-east ( F i g u r e 3) the study area was surrounded by r e c e n t l y logged or c l e a r e d areas and young p l a n t a t i o n s which were not g e n e r a l l y i n h a b i t e d by winter wrens. The study area was s t u d i e d as a complete e n t i t y only d u r i n g the breeding p e r i o d . I t was used to augment and check the o b s e r v a t i o n s on the study p l o t s . Study P l o t s Three approximately 4 ha study p l o t s A, B, and C (12.2 ha t o t a l ) were l o c a t e d w i t h i n the study a r e a . They measured 225 x 175 m although p l o t s B and C were i n e f f e c t s l i g h t l y l a r g e r because the area between the p l o t s was a l s o s t u d i e d . These p l o t s were s e l e c t e d to sample as wide a v a r i e t y of ecosystems as p o s s i b l e , t o minimize the e f f e c t of f o r e s t edge, and yet a l s o to provide a s u f f i c i e n t sample of the ecosystems i n which winter wrens were numerous enough that the b i o l o g y of the b i r d s c o u l d be s t u d i e d e f f i c i e n t l y . The p r i n c i p a l research e f f o r t was d i r e c t e d to the study p l o t s which were s t u d i e d year round. 14 Removal P l o t s The removal p l o t s D and E were of the same s i z e and c o n f i g u r a t i o n as the study p l o t s and ranged from 130 to 190 m and 220 to 265 m i n e l e v a t i o n , r e s p e c t i v e l y . They were l o c a t e d 1.3 and 1.7 km from the study area, from which they were separated by e x t e n s i v e d i s t u r b e d areas r a r e l y u t i l i z e d by winter wrens. L i k e the study a r e a , the removal p l o t s were s t u d i e d only d u r i n g the breeding p e r i o d . They served to augment and check the o b s e r v a t i o n s of winter wrens on s i m i l a r ecosystems of the study area and study p l o t s , and to e v a l u a t e the replacement of removed t e r r i t o r i a l males. C a t c h i n g , Banding, and C o l l e c t i n g Winter Wrens I caught most of my winter wrens i n mist nets (Table 1). Of the 94 f i r s t c a p t u r e s and 49 r e c a p t u r e s , 68 (72%) and 38 (78%), r e s p e c t i v e l y were by mist n e t s . T e r r i t o r i a l males were a l s o caught d u r i n g the breeding p e r i o d using recorded winter wren songs (Gunn and K e l l o g g 1975) i n c o n j u n c t i o n with mist nets (Garson 1978). The songs were p l a y e d f o r 10 minutes under mist nets i n the c e n t r e of t e r r i t o r i e s of males. Approximately two t r i a l s were r e q u i r e d f o r each t e r r i t o r i a l male cap t u r e d . 15 TABLE 1 METHODS OF CAPTURE AND COLLECTION OF WINTER WRENS Captured Winter Wrens Method M i s t n e t s Recorded song w i t h m i s t nets Winter r o o s t i n g Feather t r a p s S u b t o t a l N e s t l i n g s T o t a l C o l l e c t e d Winter Wrens Method K i l l e d by a c c i d e n t i n m i s t nets Shot by s t a l k i n g Shot u s i n g recorded songs T o t a l Number o f F i r s t Captures (Recaptures) 68(38) 13(5) 11(6) 2(0) 94(49) 98 192 Number C o l l e c t e d 4 106 16 126 16 Over the winter of 1980, I put small c r o s s e d twigs i n the entrance holes of 14 n e s t s . The seven nests at which the twigs were c o n s i s t e n t l y d i s t u r b e d were checked two to three hours before s u n r i s e on s i x n i g h t s over l a t e December through January. On 30 January, 1980, the c o l d e s t n i g h t of nest i n s p e c t i o n s (-9°C), two nests were found to c o n t a i n e i g h t and 12 communally r o o s t i n g winter wrens, of which two and one b i r d ( s ) , r e s p e c t i v e l y , escaped. The p r o p o r t i o n a l d i s t r i b u t i o n s of males and females i n the two n e s t s were about e q u a l : three males, two females, and one sex unknown i n one nest, and s i x males and f i v e females i n the o t h e r . In a d d i t i o n , on the nig h t of 11 February, 1980, a very m i l d night (+6°C), a s o l i t a r y a d u l t male was captured r o o s t i n g on top of a nest i n a c r a c k i n an o l d , hollow cedar snag. Two females were caught d u r i n g the n e s t i n g p e r i o d using hoop n e t t i n g t r a p s (Pieman 1977, 1980) b a i t e d with f e a t h e r s s u i t a b l e f o r use as nest l i n i n g m a t e r i a l . Due to the low capture r a t e (1 female/910 t r a p h o u r s ) , such t r a p s do not seem p r a c t i c a b l e as a capture method f o r winter wrens. I banded 98 n e s t l i n g s i n 18 nests d u r i n g the study. Many nests were not found, although were known to e x i s t from the a c t i o n s of the parents and/or the subsequent 17 appearance of the broods. A l l c aptured winter wrens were banded on the r i g h t l e g using a numbered aluminum band ( s i z e 0 ). A d u l t , y e a r l i n g , and j u v e n i l e b i r d s were a l s o banded with i n d i v i d u a l , d i s t i n c t i v e l y c o l o u r e d combinations of l i g h t - c o l o u r e d epoxy-painted bands. Over the study, the c o l o u r combinations of about 80% of the b i r d s observed to be banded were read s u c c e s s f u l l y . A t o t a l of 126 winter wrens were c o l l e c t e d d u r i n g the study (Table 1). Four were k i l l e d a c c i d e n t a l l y d u r i n g mist n e t t i n g , 48 were shot on and around the removal p l o t s , and the r e s t were shot i n areas on and around the Research F o r e s t remote from the study area. During the 1980 c o l l e c t i o n s on the removal p l o t s , t e r r i t o r i a l males were a t t r a c t e d to the gun using recorded winter wren songs at approximately twice the e f f i c i e n c y of s t a l k i n g . Sex Determination of Winter Wrens The sex of y e a r l i n g and a d u l t winter wrens was determined i n the f i e l d and at autopsy using general m o r p h o l o g i c a l , b e h a v i o u r a l , and anatomical c r i t e r i a of sex (Table 2 ) . Acceptable c r i t e r i a , any one of which was s u f f i c i e n t to determine sex, i n c l u d e d presence or absence of a brood patch, song, sexual d i s p l a y , egg l a y i n g , i n c u b a t i o n TABLE 2 SUMMARY OF THE GENERAL MORPHOLOGICAL, BEHAVIORAL, AND ANATOMICAL CRITERIA FOR WINTER WREN SEX DETERMINATION Acceptable C r i t e r i a of Sex - any one of which was s u f f i c i e n t to determine sex (period e f f e c t i v e ) . Brood patch: presence - female; absence - male (mid-April through J u l y ) . Song: male ( a l l months but only occurs frequently from early-February through September). Sexual display: male; or i f courted by male before the appearance of juveniles-female (breeding period). Egg laying, incubation and/or the only adult with a young brood - female (breeding period). Autopsy: male or female (less e f f e c t i v e f o r juveniles and young yearlings and outside the breeding period) Corroborative C r i t e r i a of Sex - not s u f f i c i e n t alone to determine sex (period e f f e c t i v e ) . Loud, prolonged, spaced chipping from a r e l a t i v e l y exposed s i t e : male ( f a l l and especially winter but also to a lesser extent during the breeding period). Bold aggressive behavior toward the observer: male ( a l l seasons). Where both parents are feeding the young, does most of the work: female (breeding period). Observed on the t e r r i t o r y of a known male during the breeding period before the appearance of juveniles: female (early breeding period). CO 19 and brood tending, and sex organs a t autopsy. The sex of b i r d s determined under these c r i t e r i a was termed "known". However, i t should be noted that the a c c e p t a b l e m o r p h o l o g i c a l , and b e h a v i o u r a l c r i t e r i a of sex were only a p p l i c a b l e d u r i n g the breeding p e r i o d while autopsy was only p o s s i b l e f o r c o l l e c t e d b i r d s . Using these c r i t e r i a , the "known" sex of 57 (61%) of the colour-banded b i r d s and 96 (76%) of the c o l l e c t e d b i r d s or a t o t a l of 153 (70%) of a l l the study b i r d s was determined. I took ten measurements from captured and c o l l e c t e d winter wrens i n an e f f o r t to determine the sex of the 67 b i r d s of "unknown" sex, and to d e v i s e a method of sex de t e r m i n a t i o n which would a l s o be a p p l i c a b l e o u t s i d e the breeding p e r i o d . These were: weights, lengths of f i r s t p r i m a r i e s and c e n t r a l r e c t r i c e s , t o t a l l e ngths and s h a f t diameters of plucked f i r s t p r i m a r i e s and c e n t r a l r e c t r i c e s , wing l e n g t h s , and crown and b r e a s t c o l o u r s . However, complete measurements were not a v a i l a b l e f o r a l l b i r d s as measurements of the l e n g t h of f i r s t p r i m a r i e s and c e n t r a l r e c t r i c e s , wing l e n g t h s , and c o l o u r s were not i n s t i t u t e d u n t i l mid-study. In a d d i t i o n , f e a t h e r and c o l o u r measurements were not a v a i l a b l e f o r b i r d s taken d u r i n g the moult. 20 Winter wrens were weighed to the nearest 0.1 g at capture or c o l l e c t i o n u s i n g a 1-30 g s p r i n g s c a l e . Weights of j u v e n i l e and young y e a r l i n g s were excluded from c o n s i d e r a t i o n . Maximum lengths of l e f t f i r s t p r i m a r i e s (most proximal) and l e f t c e n t r a l r e c t r i c e s were taken i n p l a c e to the nearest 0.1 mm us i n g a m a c h i n i s t ' s d i a l micrometer. These f e a t h e r measurements d i d not, as hoped, o b v i a t e the n e c e s s i t y of p l u c k i n g the f e a t h e r s f o r l a t e r measurement. In a d d i t i o n , the lengths of 28 p a i r e d r i g h t f i r s t p r i m a r i e s were taken. The l e f t f i r s t p r i m a r i e s and c e n t r a l r e c t r i c e s were then plucked, and a t the completion of the f i e l d work, t o t a l maximum fe a t h e r lengths were taken to the nearest 0.1 mm. The diameters of the f e a t h e r s h a f t s were measured at the s u p e r i o r u m b i l i c u s at r i g h t angles to the u m b i l i c a l scar to the nearest 0.01 mm u s i n g a 40X d i s s e c t i n g microscope with a c a l i b r a t e d e y e p i e c e . In a d d i t i o n , 76 p a i r e d r i g h t f i r s t p r i m a r i e s and 79 p a i r e d c e n t r a l r e c t r i c e s were a l s o plucked and measured. L e f t wing (chord) l e n g t h s (Canadian W i l d l i f e S e r v i c e 1976-77) were measured to the nearest 0.1 mm f o r a l l c o l l e c t e d b i r d s . 21 The p r i m a r i e s and r e c t r i c e s of y e a r l i n g s are grown while the b i r d s are n e s t l i n g s and j u v e n i l e s , and are r e t a i n e d d u r i n g the p o s t - j u v e n a l moult. Although grown while the b i r d s are s m a l l , these f e a t h e r s are of s i m i l a r s i z e to those of a d u l t s and the f e a t h e r and wing l e n g t h measurements of mature f e a t h e r s were i n c l u d e d i n the de t e r m i n a t i o n of sex. F i n a l l y , the average Munsell c o l o u r s (Wood and Wood 1972; Munsell C o l o r 1973) of the crown and lower b r e a s t plumage were measured i n s u n l i g h t . Colour measurements of j u v e n i l e s were excluded from c o n s i d e r a t i o n as j u v e n a l c o l o u r a t i o n d i f f e r s from that of y e a r l i n g s and a d u l t s . The v a r i o u s weights, l e f t f e a t h e r measurements, l e f t wing l e n g t h measurements, and c o l o u r measurements of winter wrens (except j u v e n i l e s or very young y e a r l i n g s of both "known" and "unknown" sex as determined by the ge n e r a l m o r p h o l o g i c a l , b e h a v i o u r a l , and anatomical c r i t e r i a ) were p l o t t e d over a cal e n d a r year. Measurements d i d not appear to vary with e i t h e r the time of year or the age of the b i r d s . The weights and f e a t h e r measurements of "known" males were s t a t i s t i c a l l y g r e a t e r than those of "known" females (Table 3 ) . The crown and bre a s t plumage c o l o u r v a l u e s ( r e l a t i v e b r i g h t n e s s of the s p e c t r a l c o l o u r ) of females were s t a t i s t i c a l l y g r e a t e r than those of males. TABLE 3 DATA SUMMARY AND STATISTICAL COMPARISONS OF "KNOWN" MALE AND FEMALE WINTER WREN WEIGHTS, LEFT FEATHER MEASUREMENTS, LEFT WING LENGTHS AND COLOURS A t t r i b u t e Data Summary x ± SD (range)n Males Females S t a t i s t i c a l Comparisons t or x' value* (df) Weights (g) 9.4±0.3(8.1-11.6)96 9.110.9(7.6-10.9)30 t = 2.37* (124) F i r s t Primary Lengths (mm) Central R e c t r i x Lengths (mm) 35.811.0(32.6-37.7)97 30.311.2(26.5-32.6)87 33.310.9(32.1-35.0)28 28.611.0(26.5-30.8)23 t = 11.68*** (123) t = 6.18*** (108) Plucked F i r s t Primary Lengths (mm) 39.610.9(37.5-41.4)111 Plucked F i r s t Primary Shaft Diameters (mm) 0.5510.02(0.50-0.61)113 37.4+0.7(34.8-39.3)40 0.51+0.02(0.47-0.55)39 t = 11.31*** (149) t = 9.78*** (150) Plucked Central R e c t r i x Lengths (mm) Plucked Central R e c t r i x Shaft Diameters (mm) Wing Lengths (mm) 33.4+1.4(30.3-38.0)98 0.3910.02(0.35-0.43)100 45.811.1(43.1-48.1)76 31.211.1(27.8-33.1)38 0.37+0.02(0.34-0.40)38 42.8+1.0(41.0-44.2)16 t = 8.32*** (134) t = 8.58*** (136) t = 9.92*** (90) Munsell Colours ( a l l hue 10 yr) Crown - Values Crown - Chromae Breast - Values Breast - Chromae 3.010.2(3-4)87 4.3+1.1(2-6)87 5.810.6(5-7)99 5.810.8(4-8)99 3.410.5(3-4)30 4.311.0(3-6)30 6.310.6(5-7)27 6.110.9(4-8)27 x* = 30.73*** (1) x 2 = 1.98 (3) x 2 = 27.64*** (2) x 2 = 1.80 (2) * P < 0.05; *** P < 0.001 23 However, the crown and b r e a s t plumage chromae ( r e l a t i v e p u r i t y of the s p e c t r a l c o l o u r ) of.males and females were not s t a t i s t i c a l l y d i f f e r e n t . A l l crown and b r e a s t plumage hues were 1OYR (dominant s p e c t r a l c o l o u r ) . Because of the o v e r l a p between the sexes, no s i n g l e measurement c o u l d be used t o determine winter wren sex. However, i t was p o s s i b l e to develop r e l i a b l e sex d i s c r i m i n a t i o n c r i t e r i a u s i n g combined measurements. The c r i t e r i a a s s o c i a t e d with each measurement are summarized i n Table 4. They were d e r i v e d from the data of Table 3. I u t i l i z e d the combined measurements as f o l l o w s : 1 ) At l e a s t one feather measurement or wing l e n g t h was necessary to a s s i g n the sex of a b i r d ( c r i t e r i a of Table 4 ) . Otherwise the b i r d was as s i g n e d sex unknown. Weights and/or c o l o u r s alone were not c o n s i d e r e d s u f f i c i e n t l y d i a g n o s t i c . 2) A l l measurements i n the range sex unknown (Table 4) were ign o r e d except when a l l measurements of a b i r d were sex unknown. In t h i s l a t t e r case a b i r d was ass i g n e d as sex unknown. 3) When an i n d i v i d u a l b i r d e x h i b i t e d both male and female c r i t e r i a f o r weight, f e a t h e r measurements, or wing l e n g t h , but not f o r c o l o u r , i t was as s i g n e d as sex unknown. TABLE 4 WINTER WREN SEX DISCRIMINATION CRITERIA BASED ON WEIGHTS LEFT FEATHER MEASUREMENTS, LEFT WING LENGTHS, AND COLOURS' Attributes Weight (g) F i r s t Primary Length (mm) Central Rectrix Length (mm) Plucked F i r s t Primary Length (mm) Plucked F i r s t Primary Shaft Diameter (mm) Wing Length (mm) Crown Colour Value Breast Colour Value Sex Discrimination C r i t e r i a Female unknown -8.0 >8.0 i 32.5 £ 37.4 £ 0.49 Plucked Central Rectrix Length (mm) t 30.2 Plucked Central Rectrix Shaft Diameter (mm) - 0.34 - 43.0 4? £ 7? 32.6-35.0 f 30.8 37.5-39.3 0.55-0.55 30.3-33.1 0.35-0.40 43.1-44.2 A l l A l l Male ^35.1 £30.9 £39.4 >0.56 £33.2 >0.41 >44.3 3? 5 and 6? Although male and female colour values were s t a t i s t i c a l l y d i f f e r e n t (Table 3 ) , the ranees of the colour values overlapped completely and could not be used to distinguish sex - hence ?. 25 4) Where a b i r d had been captured s e v e r a l times, a l l measurements were c o n s i d e r e d t o g e t h e r . 5) P a i r e d r i g h t wing f e a t h e r measurements, although not used i n the development of sex d i s c r i m i n a t i o n c r i t e r i a , were a l s o c o n s i d e r e d i n the assignment of sex. P a i r e d f e a t h e r s g e n e r a l l y had very s i m i l a r measurements. Male and female c r i t e r i a never o c c u r r e d i n the same f e a t h e r p a i r although one measurement might be male or female and the other sex unknown. The sex of the 67 winter wrens of "unknown" sex as determined by m o r p h o l o g i c a l , b e h a v i o u r a l , and anatomical c r i t e r i a was assigned u s i n g the preceding conventions as: 29 males and 12 females, while 26 remained as sex unknown. Of the 26 b i r d s of unknown sex, 18 l a c k e d f e a t h e r or wing measurements because they had been taken e i t h e r d u r i n g moult or very e a r l y i n the study before f e a t h e r measurements were taken, four had only unknown measurements, and four e x h i b i t e d both male and female c r i t e r i a . The sex composition of a l l the colour-banded and c o l l e c t e d winter wrens of the study as determined by using an i n t e g r a t e d method employing both the g e n e r a l m o r p h o l o g i c a l , b e h a v i o u r a l , and anatomical c r i t e r i a (Table 26 2) and the sex d i s c r i m i n a t i o n f u n c t i o n s (Table 4) was as f o l l o w s : 143 males, 51 females, and 26 sex unknown. The e f f i c i e n c y of the i n t e g r a t e d method was 88% (194 of 220 b i r d s ) . I t was s l i g h t l y lower f o r banded than f o r c o l l e c t e d b i r d s because a l a r g e r p o r t i o n of the banded b i r d s were taken d u r i n g moult when not a l l measurements were a v a i l a b l e . Hawthorn (1975) and Garson (1978) used a c l e a r s e p a r a t i o n given by weights p l o t t e d a g a i n s t wing lengths (males > females) to determine the sex of European wrens ( T . t . t r o g l o d y t e s ) . T h e i r data were not a p p l i c a b l e here as European wrens are approximately 10% l a r g e r than the winter wrens of t h i s study. The use of weights and wing l e n g t h s alone would not have given a c l e a r s e p a r a t i o n i n t h i s study. Due to the e x t e n s i v e o v e r l a p of male and female measurements, p a r t i c u l a r l y weights (Table 3), the sex of most b i r d s would have remained unknown. Because s i m i l a r s i z e d i f f e r e n c e s and v a r i a t i o n i n measurements may occur among and w i t h i n the v a r i o u s North American subspecies of winter wrens, the method of sex d e t e r m i n a t i o n developed i n t h i s study should be t e s t e d before being a p p l i e d elsewhere. 27 Age Determination of Winter Wrens I d e f i n e d the f o l l o w i n g age c l a s s e s f o r my p o p u l a t i o n of winter wrens: n e s t l i n g s , j u v e n i l e s , y e a r l i n g s (young y e a r l i n g s , and o l d e r y e a r l i n g s ) a d u l t s , and age unknown. N e s t l i n g s were d e f i n e d as young b i r d s s t i l l i n the n e s t . J u v e n i l e s were young b i r d s which had f l e d g e d but were s t i l l w i t h t h e i r p a r e n t ( s ) and/or s i b l i n g s . Y e a r l i n g s were d e f i n e d as b i r d s which had l e f t t h e i r p a r e n t ( s ) and/or s i b l i n g s u n t i l the general moult i n August of the year f o l l o w i n g h a t c h i n g . Y e a r l i n g s were f u r t h e r d i v i d e d i n t o young y e a r l i n g s u n t i l m i d - f a l l and as o l d e r y e a r l i n g s t h e r e a f t e r . A l l b i r d s o l d e r than y e a r l i n g s were c o n s i d e r e d a d u l t s . Hawthorn (1971) found that a d u l t and y e a r l i n g European wrens c o u l d be d i s t i n g u i s h e d on the b a s i s of c o n t r a s t on the g r e a t e r wing c o v e r t s , and by b a r r i n g on the l a r g e f e a t h e r of the a l u l a and on the f o u r t h primary. However, Garson (1978) noted from a p e r s o n a l communication with E. Meek (p. 11) that i t was o f t e n impossible to c l a s s i f y b i r d s using t h i s scheme d u r i n g m i g r a t i o n along the n o r t h -east c o a s t of England. I was unable to d i s t i n g u i s h o l d e r y e a r l i n g s from a d u l t s u s i n g these methods, nor was I a b l e to d i s c o v e r any other g e n e r a l d i s t i n g u i s h i n g d i f f e r e n c e s between o l d e r y e a r l i n g s and a d u l t s . However, s i n g i n g 28 young y e a r l i n g males c o u l d be d i s t i n g u i s h e d from o l d e r y e a r l i n g s and a d u l t s from l a t e June through October by t h e i r immature songs (Armstrong 1955, p. 85) which were incomplete and u s u a l l y f a i n t and quavery. J u v e n i l e s and very young y e a r l i n g s were d i s t i n g u i s h e d from o l d e r y e a r l i n g s and a d u l t s by the presence of a gape and/or down, being a member of a brood, or as having been banded as a n e s t l i n g . I used average weights and f i r s t primary and c e n t r a l r e c t r i x l e n g t h s from broods of n e s t l i n g s of known hat c h i n g dates to develop a method f o r the age d e t e r m i n a t i o n f o r n e s t l i n g s . Weights of i n d i v i d u a l n e s t l i n g s , except n e s t l i n g s under s i x days of age which were weighed together, were taken to the nearest 0.1 g u s i n g 0-5 and 0-10 g s p r i n g s c a l e s . Maximum lengths of r i g h t f i r s t p r i m a r i e s and c e n t r a l r e c t r i c e s were measured to the nearest 0.1 mm. There was v a r i a t i o n i n the weights and f e a t h e r measurements w i t h i n broods and the v a r i a t i o n i n c r e a s e d as the broods became o l d e r . However, as the extent of the v a r i a t i o n appeared f a i r l y c onstant among broods of s i m i l a r ages t h i s v a r i a t i o n was i g n o r e d . For each group of measurements of a brood of n e s t l i n g s , the average weight and f e a t h e r lengths f o r the brood were c a l c u l a t e d . 29 E i g h t broods of n e s t l i n g s of known hat c h i n g dates were measured from one to f i v e times each. Hatching o c c u r r e d over a 24 hour p e r i o d or l e s s i n a l l f i v e nests i n which i t was observed. Broods were f i r s t measured at from three to s i x days of age and again at three to f i v e day i n t e r v a l s t h e r e a f t e r u n t i l f l e d g i n g or the l o s s of the brood. Average weights and f e a t h e r measurements of i n d i v i d u a l broods of the same age were themselves averaged. The average weights and f e a t h e r measurements of the broods of known age are summarized i n Table 5. G e n e r a l l y , n e s t l i n g s gained weight r a p i d l y f o r approximately the f i r s t nine days a f t e r h a t c h i n g and weights were most u s e f u l f o r age d e t e r m i n a t i o n d u r i n g t h i s p e r i o d . F i r s t p r i m a r i e s i r r u p t e d a t approximately four days of age and grew r a p i d l y u n t i l f l e d g i n g when they were approximately 7/8 of t h e i r mature l e n g t h . C e n t r a l r e c t r i c e s , on the other hand, i r r u p t e d at approximately f i v e days of age and grew more s l o w l y u n t i l f l e d g i n g when they were approximately 1/2 t h e i r mature l e n g t h . Primary and r e c t r i x measurements were most u s e f u l i n age de t e r m i n a t i o n d u r i n g mid- to l a t e - n e s t l i n g stages but became i n c r e a s i n g l y v a r i a b l e among broods d u r i n g l a t e n e s t l i n g stages. TABLE 5 AVERAGE WEIGHTS AND FIRST PRIMARY AND CENTRAL RECTRIX LENGTHS FROM BROODS OF WINTER WREN NESTLINGS OF KNOWN DATES OF HATCHING Known Age Number of Broods (Days) Measured Weights(g) Measurements x (range) F i r s t Primary Lengths (mm) Central Rectrix Lengths (mm) 3 1 3.1 0.1 0.0 4 1 1.8 0.0 0.0 5 2 3.3(3.2-3.3) 0.3(0.1-0.4) 0.0(0.0) 6 2 5.6(5.2-6.1) 4.1(3.2-5.0) 0.2(0.2) 7 1 4.7 2.2 0.3 8 2 6.1(5.4-6.7) 6.1(4.3-7.9) 0.6(0.1-1.0) 9 4 7.6(7.2-7.8) 11.6(9.3-12.9) 2.5(1.3-3.5) 10 1 8.6 17.3 4.9 12 3 8.6(7.6-9.1) 17.2(12.6-21.1) 4.7(1.6-5.2) 14 2 8.7(8.7-8.8) 27.2(26.3-28.2) 12.3(11.6-13.1) 15 3 8.7(8.0-9.4) 25.1(20.4-28.1) 9.6(5.8-12.2) 18 2 8.4(8.3-8.5) 29.2(27.3-31.1) 12.6(9.7-15.5) U) o 31 I estimated the ages of 11 broods of n e s t l i n g s of unknown h a t c h i n g dates by comparing t h e i r measurements with the measurements of the broods of n e s t l i n g s of known hat c h i n g dates (Table 5). Three broods of n e s t l i n g s of unknown h a t c h i n g dates were measured two, three, and three times, r e s p e c t i v e l y . The f i r s t measurements of these broods were used t o determine the ages of the broods and the subsequent measurements corresponded w e l l with the measurements of broods of known hatching dates of the same estimated age. Some n a t u r a l v a r i a t i o n o c c u r r e d among broods i n the r e l a t i v e growth r a t e s of weight and f e a t h e r s . T h i s v a r i a t i o n appeared t o i n c r e a s e i n o l d e r broods. F u r t h e r , I observed that broods grew more slowly d u r i n g p e r i o d s of c o l d , r a i n y weather, and when the male d i d not a s s i s t i n f e e d i n g . However, d e s p i t e t h i s v a r i a b i l i t y , i t appeared that e stimates of the age of broods of n e s t l i n g s were p o s s i b l e w i t h i n an accuracy of one day t o approximately age 12 days. The accuracy probably decreased to two or more days f o r o l d e r broods. I c a l c u l a t e d the dates of c l u t c h and i n c u b a t i o n commencement, h a t c h i n g , and f l e d g i n g where they were not known f o r a l l broods of n e s t l i n g s as f o l l o w s . C l u t c h accumulation appeared t o proceed at a r a t e of one egg per 32 day based on o b s e r v a t i o n s of 10 new eggs i n four n e s t s . The most common c l u t c h s i z e was s i x (5.3 ± 0.9; 3-6; 13), although c l u t c h e s of f i v e were a l s o common (X ± S.D.; range; n ) . C l u t c h e s of l e s s than f i v e only o c c u r r e d i n June and J u l y . I t h e r e f o r e allowed s i x days f o r c l u t c h accumulation except where c l u t c h s i z e was known to be l e s s . The i n c u b a t i o n p e r i o d was d e f i n e d as extending from the day egg l a y i n g ceased u n t i l the day before a l l the eggs had hatched and was taken as 15 days based on o b s e r v a t i o n s on two n e s t s . The n e s t l i n g p e r i o d was d e f i n e d as extending from the day a l l the eggs had hatched u n t i l the day the n e s t l i n g s f l e d g e d . Based on f i v e observed n e s t l i n g p e r i o d s (16, 17, 17, 18, and 18 day s ) , I allowed a n e s t l i n g p e r i o d of 17 days f o r those nests f o r which the n e s t l i n g p e r i o d was not known. The f i r s t and l a s t c l u t c h commencements were c a l c u l a t e d as A p r i l 16 and J u l y 11 while the f i r s t and l a s t f l e d g i n g s were observed on May 23 and August 14. The monthly d i s t r i b u t i o n of c l u t c h commencements d u r i n g the study oc c u r r e d as f o l l o w s : A p r i l - 10, May - 4, June - 4, and J u l y - 2. Six of the May through J u l y c l u t c h commencements were known to be second n e s t i n g s a f t e r both f a i l e d and s u c c e s s f u l f i r s t n e s t i n g s . 33 WINTER WREN DISPERSION AS A FUNCTION OF TERRITORIALITY I n t r o d u c t i o n In t h i s chapter I show male winter wrens to be t e r r i t o r i a l and suggest t e r r i t o r i a l i t y f o r females as w e l l . I propose that winter wrens are spaced by t e r r i t o r i a l behaviour. I d e s c r i b e my p o p u l a t i o n s of winter wrens and show t h a t they were not u n i f o r m l y d i s t r i b u t e d . F i n a l l y , I demonstrate that p o t e n t i a l l y t e r r i t o r i a l males were a v a i l a b l e d u r i n g the breeding p e r i o d s s t u d i e d . These males c o u l d have occupied the empty or s p a r s e l y occupied areas had the h a b i t a t been s u i t a b l e . I use the f o l l o w i n g d e f i n i t i o n of t e r r i t o r y (Noble 1939, p. 267): "any defended a r e a " . To t h i s I add the p r o v i s i o n that the h o s t i l i t y of defense i n c l u d e s the combined outcome of a t t a c k , avoidance, and/or escape as r e c o g n i z e d by Tinbergen (1957). Male Old World wrens are s t r o n g l y t e r r i t o r i a l (e.g., H a r r i s s o n and Buchan 1934, 1936; K l u i j v e r et a l . 1940; Armstrong 1955, 1956; K e n t i s h 1976; Haneda and Kosaki 1971; Armstrong and Whitehouse 1977; Garson 1978; Wesolowski 1981). Male New World winter wrens have always been assumed to be t e r r i t o r i a l (Kendeigh 1947; E r s k i n e 34 1977; Holmes e t a l . 1979; Sabo 1980, and o t h e r s ) but i n f a c t t e r r i t o r i a l i t y has never been shown c o n c l u s i v e l y . However, Heath's (1920) account of the n e s t i n g h a b i t s of the A l a s k a w i n t e r wren (T.t_. a l a s k e n s i s ) s t r o n g l y i n d i c a t e d t e r r i t o r i a l i t y , a l t h o u g h h i s account was w r i t t e n b e f o r e Howard's (1920) i n t r o d u c t i o n of the concept of t e r r i t o r i a l i t y t o E n g l i s h - s p e a k i n g o r n i t h o l o g y . To show t h a t w i n t e r wrens a r e t e r r i t o r i a l , I must demonstrate t h e c r i t e r i a of s i t e attachment t o l o c a l i z e d a r e a s and de f e n s e of t h e s e a r e a s , i n c l u d i n g a t t a c k , a v o i d a n c e and/or escape. S i t e attachment w i t h a v o i d a n c e a l o n e would p r o v i d e o n l y c i r c u m s t a n t i a l e v i d e n c e of t e r r i t o r i a l i t y . Methods The s t u d y a r e a ( i n c l u d i n g t h e s t u d y p l o t s ) , and the removal p l o t s were s u r v e y e d i n t o square g r i d s ( s q u a r e s were 50 x 50 m). The g r i d p o i n t s were marked w i t h w h i t e p a i n t e d s t a k e s . The top s of the g r i d s t a k e s were marked on a l l f o u r s i d e s w i t h numbers t h a t were c l e a r l y l e g i b l e u s i n g 7 X f i e l d g l a s s e s a t more than 50 m. I e s t i m a t e d the l o c a t i o n s of w i n t e r wren o b s e r v a t i o n s i n the f i e l d by l o c a t i n g a nearby g r i d c o r n e r s t a k e o r s t a k e s . Then, u s i n g a h a n d - s i g h t i n g compass, I d e t e r m i n e d the g r i d 35 o r i e n t a t i o n . F i n a l l y , I estimated the c o o r d i n a t e s of the o b s e r v a t i o n p o i n t to the nearest m from the nearest north east g r i d s take. Observations of winter wrens were p l o t t e d on 1:787 s c a l e (6.35 cm to 50 m) maps ( I n t e r n a t i o n a l B i r d Census Committee 1970). I s t u d i e d winter wrens on the e n t i r e study area and on the removal p l o t s by walking 50 m spaced t r a n s e c t s ; 25 m spaced t r a n s e c t s were walked on the study p l o t s . The l o c a t i o n s and ti m i n g of the t r a n s e c t s were s y s t e m a t i c a l l y scheduled so that a l l areas w i t h i n the study area, the study p l o t s , and the i n d i v i d u a l removal p l o t s r e c e i v e d equal study from the i n d i v i d u a l o b s e r v e r s . F i e l d s t u d i e s were begun j u s t a f t e r dawn and proceeded u n t i l b i r d a c t i v i t y slackened, u s u a l l y at mid-day duri n g the s p r i n g and summer, and i n the l a t e a f t e r n o o n d u r i n g the f a l l and winter. Work co n t i n u e d i n l i g h t r a i n or snow but was g e n e r a l l y stopped by heavy r a i n or blowing snow. During the s i x - to seven-hour study p e r i o d each day an observer c o u l d cover about o n e - t h i r d of the study area, one t o two study p l o t s or two removal p l o t s . I d i v i d e d the year i n t o t h r e e p e r i o d s : breeding p e r i o d from March through August, the f a l l p e r i o d from September through November, and the winter p e r i o d from December through February. 36 The t r a n s e c t s on the study area were done 10, 11, 18, and 14 times d u r i n g the breeding p e r i o d s of 1978-81 r e s p e c t i v e l y . The t r a n s e c t s on the study p l o t s were done 6, 38, and 30 times d u r i n g the w i n t e r , breeding and f a l l p e r i o d s of 1979, and 28 and 65 times d u r i n g the winter and breeding p e r i o d s of 1980, r e s p e c t i v e l y . The t r a n s e c t s on the removal p l o t s were done d u r i n g the e a r l y breeding p e r i o d s of 1979-80 as f o l l o w s : D - 22 and 20 times and E -11 and 18 times, r e s p e c t i v e l y . I estimated the area and extent of the male t e r r i t o r i e s and female home ranges f o r the i n d i v i d u a l b reeding, f a l l , and winter p e r i o d s on the study a r e a , study p l o t s , and removal p l o t s by co n n e c t i n g the outermost p o i n t s of o b s e r v a t i o n of i n d i v i d u a l b i r d s . Maximum polygons (Odum and Kuenzler 1955) were developed with the f o l l o w i n g e x c e p t i o n s . Polygons were drawn f o r male t e r r i t o r i e s to represent the r e l a t i v e l y s t a b l e s t a t u s at mid-period with e x c e p t i o n of the study area d u r i n g the breeding p e r i o d s of 1979 and 1980 when the work was done in the e a r l y breeding p e r i o d and the polygons were drawn a c c o r d i n g l y . In the cases of mid-period polygons, the s h i f t i n g of t e r r i t o r i e s and t e r r i t o r y boundaries which o c c u r r e d d u r i n g the beginning of the winter p e r i o d , at the ju n c t u r e of the winter and breeding p e r i o d s , and again at 37 the j u n c t u r e of the breeding and f a l l p e r i o d s were ignored. However, due to the low numbers of o b s e r v a t i o n s of colour-banded females, i t was necessary to use a l l o b s e r v a t i o n s , and the polygons f o r female home ranges i n c l u d e the v a r i a t i o n avoided f o r male t e r r i t o r i e s by us i n g mid-period polygons. Adjacent r a t h e r than extreme outermost o b s e r v a t i o n p o i n t s were connected where the extreme p o i n t s were more than 50 m a p a r t , or where an ex t e n s i v e area not otherwise used by the b i r d would have been i n c l u d e d , or where an area of a neighbouring t e r r i t o r y or home range which was not used by the b i r d would have been i n c l u d e d i n the maximum polygon. While t h i s p r a c t i c e r e s u l t e d i n s l i g h t l y s m a l l e r polygons than would have been o b t a i n e d using the maximum polygon technique, the r e s u l t s are probably more accurate measures of the extent and areas of male t e r r i t o r i e s and female home ranges. I estimated the area and extent of the male t e r r i t o r i e s of the study area independently of the study p l o t data, although i n p r a c t i c e i t was very d i f f i c u l t not to use the e x t r a knowledge from the study p l o t s while i n t e r p r e t i n g the study area data. On the other hand, the data from the study area were used when e s t i m a t i n g the male t e r r i t o r i e s and female home ranges of the study p l o t s . 38 I t was r e l a t i v e l y easy to d i s t i n g u i s h i n d i v i d u a l males and determine t e r r i t o r i e s d u r i n g the breeding p e r i o d by a g o n i s t i c c o u n t e r s i n g i n g and c o u n t e r c h i p p i n g , and to a l e s s e r extent by a g o n i s t i c c o u n t e r c h i p p i n g d u r i n g the f a l l and winter p e r i o d . In a d d i t i o n , n e a r l y a l l of the t e r r i t o r i a l males on and around the study p l o t s were colour-banded i n 1979-80 and were thus d i s t i n g u i s h a b l e from one another. However, i t was im p o s s i b l e to i d e n t i f y i n d i v i d u a l females unless colour-banded, except i n one case, where an i n d i v i d u a l unhanded female was surrounded by other colour-banded females. The d e t e r m i n a t i o n of female home ranges was only p o s s i b l e on the study p l o t s 1979-80, when some of the females were colour-banded. The areas of the t e r r i t o r i e s and home ranges were estimated i n m2 by counting squares (1 square = 4 m 2) at the working s c a l e of 1 :787. The numbers of t e r r i t o r i a l males and male t e r r i t o r i e s on the study area d u r i n g the breeding p e r i o d were i d e n t i c a l , as no t e r r i t o r i e s overlapped the study area boundaries. However, as the male t e r r i t o r i e s on the study and removal p l o t s extended beyond the p l o t boundaries, t h e r e were more winter wrens present than complete t e r r i t o r i e s . To o b t a i n estimates of p o p u l a t i o n s on the p l o t s , I determined the complete areas of these 39 t e r r i t o r i e s . The p r o p o r t i o n of each t e r r i t o r y on each p l o t was c a l c u l a t e d to the nearest 0.1 t e r r i t o r y . These valu e s were summed to g i v e a p r o p o r t i o n a l estimate of the number of t e r r i t o r i e s on each p l o t . However, I was unable to estimate numbers of females or female home ranges on the study p l o t s as many females remained u n i d e n t i f i e d . The i n i t i a l numbers of t e r r i t o r i a l males and the gen e r a l extent of t h e i r t e r r i t o r i e s on the removal p l o t s D and E were determined d u r i n g the e a r l y breeding p e r i o d s of 1979 and 1980. These males were then shot. Up to f i v e weeks, but u s u a l l y a s h o r t e r p e r i o d , were allowed to elap s e before the process was repeated. There were two removal p e r i o d s i n 1979 and four i n 1980. Winter Wren T e r r i t o r i a l i t y Males The area and extent of the male t e r r i t o r i e s are shown as f o l l o w s : the study area d u r i n g the breeding p e r i o d s , 1978-81 i n F i g u r e s 4 to 7; the study p l o t s d u r i n g the winter , breeding and f a l l p e r i o d s 1979-80 i n F i g u r e s 8 to 12; and, the removal p l o t s d u r i n g the e a r l y breeding p e r i o d s 1979-80 i n F i g u r e 13. A l l males o l d e r than young y e a r l i n g s and which were observed r e g u l a r l y , were FIGURE 4 AREA 1 AND EXTENT OF THE MALE TERRITORIES 2 ON THE STUDY AREA 3 DURING THE BREEDING PERIOD, 1978 Areas i n ha Banded b i r d s a r e i d e n t i f i e d by c i r c l e d code numbers; ages a r e a l s o g i v e n : A - a d u l t ; Y - y e a r l i n g ; U - age unknown Study p l o t s A, B and C a r e shown i n dashed l i n e s on the study a r e a FIGURE 5 AREA AND EXTENT OF THE MALE TERRITORIES ON THE STUDY AREA DURING THE BREEDING PERIOD, 1979 FIGURE 6 AND EXTENT OF THE MALE TERRITORIES ON THE STUDY AREA DURING THE BREEDING PERIOD, 1980 FIGURE 7 AND EXTENT OF THE MALE TERRITORIES ON THE STUDY AREA DURING THE BREEDING PERIOD, 1981 FIGURE 8 AREA AND EXTENT OF THE MALE TERRITORIES ON THE STUDY PLOTS DURING THE WINTER PERIOD, 1979 FIGURE 9 AREA AND EXTENT OF THE MALE TERRITORIES ON THE STUDY PLOTS DURING THE BREEDING PERIOD, 1979 FIGURE 10 AREA AND EXTENT OF THE MALE TERRITORIES ON THE STUDY PLOTS DURING THE FALL PERIOD, 1979 FIGURE 11 AREA AND EXTENT OF THE MALE TERRITORIES ON THE STUDY PLOTS DURING THE WINTER PERIOD, 1979-80 FIGURE 12 AREA AND EXTENT OF THE MALE TERRITORIES ON THE STUDY PLOTS DURING THE BREEDING PERIOD, 1980 FIGURE 13 AREA AND EXTENT OF THE MALE TERRITORIES ON THE REMOVAL PLOTS DURING THE EARLY BREEDING PERIODS, 1979-80 Removal Plots Breeding Period Male Territories 60 l o c a l i z e d and demonstrated s i t e attachment. The p a t t e r n s of s i t e attachment of males were very s i m i l a r on the study area, study p l o t s , and removal p l o t s d u r i n g the breeding p e r i o d s , and throughout the year on the study p l o t s . A t t a c k and f i g h t i n g among winter wrens appears t o occur o n l y r a r e l y and was observed only three times. A l l were d u r i n g the breeding p e r i o d (15, 16 March, and 3 A p r i l , 1980). A l l were d i r e c t e d a g a i n s t one t e r r i t o r i a l male which took up a t e r r i t o r y i n e a r l y March, 1980, i n a s m a l l , a p p a r e n t l y unoccupied area among f i v e e s t a b l i s h e d t e r r i t o r i a l males (male 24 among males 2, 5, 8, 23, and 25 as shown i n F i g u r e 12). The a t t a c k s were made on male 24 by males 2, 5, and 23 along or j u s t o u t s i d e the boundaries of male 24's t e r r i t o r y . A l l three a t t a c k s r e s u l t e d i n short f i g h t s on the ground f o l l o w e d by escapes by male 24 toward the c e n t r e of h i s t e r r i t o r y . The best observed a t t a c k was by male 2 on 15 March, 1980 and as t h i s a t t a c k was g e n e r a l l y t y p i c a l of the o t h e r s , a short d e s c r i p t i o n f o l l o w s : Males 24 and 2 ( F i g u r e 12) were heard c o u n t e r s i n g i n g at about 08:20 hours on the warm, sunny morning of 15 March, 1980. Male 2 was l o c a t e d near the c e n t r e of h i s t e r r i t o r y while male 24 was l o c a t e d i n a d i s p u t e d 61 area between h i s t e r r i t o r y and that of male 2. As I approached male 24, he continued to s i n g i n a small clump of salmonberry and western sword f e r n . Male 2 ceased s i n g i n g and immediately flew up to a l o g about 2 m from male 24 and about 0.5 m above him, but s t i l l between male 24 and the c e n t r e of h i s own t e r r i t o r y . Male 24 continued to s i n g . Male 2 assumed an upr i g h t posture i n which he stood up on extended l e g s with h i s f e a t h e r s s l i g h t l y compressed. His head was extended upwards at an angle of about 60° from h o r i z o n t a l while h i s t a i l was compressed and p o i n t e d s l i g h t l y downwards at an angle of about 25° from h o r i z o n t a l . His wings were then l i f t e d and extended about 60° above h o r i z o n t a l to about 4/5 t h e i r l e n g t h above h i s body. The wings were slowly q u i v e r e d through about 10° of a r c . Male 2 d i d not s i n g but immediately a t t a c k e d male 24 dropping on him from the l o g and clawing and t u s s e l l i n g with him f o r about four seconds i n the understory v e g e t a t i o n . One of the p a i r or perhaps both, gave a s e r i e s of short sharp c r i e s w h i l e t u s s e l l i n g . Male 24 then flew back along the ground toward the c e n t r e of h i s own t e r r i t o r y and remained s i l e n t f o r 6 2 at l e a s t 10 minutes. Male 2 sang n e a r l y continuous songs at the p o i n t of a t t a c k f o r approximately 1.5 minutes at a height of about 20 cm above the ground i n the understory v e g e t a t i o n and then r e t u r n e d toward the c e n t r e of h i s t e r r i t o r y where he c o n t i n u e d to s i n g . About 10 minutes l a t e r the same two males were heard c o u n t e r s i n g i n g t o one another from the c e n t r e areas of t h e i r r e s p e c t i v e t e r r i t o r i e s . Throughout the r e s t of the breeding p e r i o d n e i t h e r male was ever observed to advance beyond the p o i n t of the a t t a c k , although both were observed w i t h i n a few metres of the p o i n t but always on t h e i r own s i d e . Attack and f i g h t i n g i n European wrens i s a l s o a p p a r e n t l y r a r e . Armstrong and Whitehouse (1977, p. 246) noted o n l y three or four a t t a c k s . Garson (1978) r e p o r t e d two i n s t a n c e s i n which o t h e r s had observed b i r d s clawing at each other on the ground. K l u i j v e r et a l . (1940) d i d not r e c o r d a t t a c k or f i g h t i n g . The a g g r e s s i v e u p r i g h t posture with r a i s e d wings d e s c r i b e d i n t h i s a t t a c k resembled the t h r e a t d i s p l a y p i c t u r e d i n Armstrong (1955, p. 108) but without song. 63 However, a t t a c k should not be confused with male sexual p u r s u i t of the female or the pounce (Armstrong 1955, p. 110-113), both of which I observed o c c a s i o n a l l y d u r i n g the breeding p e r i o d f o r the winter wren and which appear a n t o g o n i s t i c and d e f e n s i v e but are not. I a l s o observed a n t a g o n i s t i c behaviour by t e r r i t o r i a l males d u r i n g the breeding p e r i o d i n response to the recorded songs played on t h e i r t e r r i t o r i e s i n c o n j u n c t i o n with mist net s e t s and c o l l e c t i o n s . Most males a t t r a c t e d to the recorded songs adopted the a g g r e s s i v e p o s t u r e d e s c r i b e d f o r male 2, although u s u a l l y without r a i s e d wings. A l l the t e r r i t o r i a l males a t t r a c t e d to the recorded songs and not immediately caught or shot, r e s o r t e d t o song a f t e r s e a r c h i n g f o r the supposed i n t r u d e r . The t e r r i t o r i a l song and the accompanying d i s p l a y s of the male winter wrens of t h i s study appeared very s i m i l a r to those d e s c r i b e d f o r the European wren ( K l u i j v e r et a l . 1940; Armstrong 1944, 1955). I heard the f i r s t t e r r i t o r i a l songs of the year i n l a t e January and e a r l y February (3 February, 1978; 9 February, 1979; 2 February, 1980; 27 January, 1981) and they became gen e r a l i n the l a s t week of February. T e r r i t o r i a l song decreased g r a d u a l l y i n June and J u l y and was r a r e l y heard i n August 64 and e a r l y September d u r i n g moult. There was a short resurgence of song i n l a t e September and e a r l y October a f t e r which i t d e c l i n e d and was very r a r e l y heard ( f o u r o c c a s i o n s only) i n December and January. The t e r r i t o r i a l song of the European wren has long been thought t o r e p e l other males, among other f u n c t i o n s ( K l u i j v e r et a l . 1940; Armstrong 1944, 1955). The r e p e l l i n g e f f e c t of song advertisement on other males has been demonstrated f o r the great t i t (Parus major) (Krebs 1971, 1977; Krebs et a l . 1978). In a s i m i l a r way, the t e r r i t o r i a l song of the winter wren probably f u n c t i o n s t o discourage t r e s p a s s e r s and p o t e n t i a l t r e s p a s s e r s , t o i n c r e a s e the d i s t a n c e at which a t e r r i t o r i a l male may defend h i s t e r r i t o r y , and t o reduce the amount of d i r e c t i n t e r a c t i o n and f i g h t i n g among males. There was c o n s i d e r a b l e s h i f t i n g of boundaries at the beginning of the winter p e r i o d , at the j u n c t u r e of the winter and breeding p e r i o d s , and again a t the j u n c t u r e of the breeding and f a l l p e r i o d s . An i n d i c a t i o n of the changes i n t e r r i t o r i e s and t e r r i t o r y boundaries d u r i n g these p e r i o d s of i n s t a b i l i t y can be o b t a i n e d i n two ways. F i r s t , changes can be seen i n a p e r i o d t o p e r i o d comparison of the s t a b l e mid-period t e r r i t o r y boundaries on the study p l o t s 1979-80 ( F i g u r e s 8 t o 12). Second, 65 changes are a l s o demonstrated i n a comparison of the t e r r i t o r y boundaries of the study p l o t s as estimated f o r the study area i n i t s e n t i r e t y d u r i n g the very e a r l y b r e e d i n g p e r i o d s i n 1979 and 1980 ( F i g u r e s 5 and 6) as opposed to the corresponding t e r r i t o r y boundaries as estimated f o r the study p l o t s d u r i n g the more s t a b l e mid-breeding p e r i o d s ( F i g u r e s 9 and 12), although a d m i t t e d l y i n t h i s second comparison the l o c a t i o n s of the t e r r i t o r y boundaries were o b t a i n e d by d i f f e r e n t methods. T e r r i t o r i e s as estimated d u r i n g the s t a b l e mid-periods d i d not g e n e r a l l y o v e r l a p . Nor d i d i t appear t h a t much o v e r l a p o c c u r r e d d u r i n g the p e r i o d s of i n s t a b i l i t y but r a t h e r t h at t e r r i t o r y boundaries changed. The two r e c o r d e d cases of o v e r l a p of t e r r i t o r i e s i n v o l v e d u n u s u a l l y unaggressive males. The t e r r i t o r y of male 9 ( F i g u r e 9) overlapped m a r g i n a l l y on the t e r r i t o r i e s of males 7 and 11 where he was o f t e n observed f o r a g i n g . However, as he was not observed with the other t e r r i t o r y h o l d e r s , nor was he observed to s i n g on the other t e r r i t o r i e s , h i s o v e r l a p may be a simple case of undetected t r e s p a s s . Male 9 sang l e a s t and was the shyest of a l l the t e r r i t o r i a l males and i t i s a l s o p o s s i b l e t h a t he may have been t o l e r a t e d by h i s neighbours due to h i s l a c k of a g g r e s s i v e n e s s . Nonetheless male 9 a p p a r e n t l y 6 6 v. obtained a mate and helped f l e d g e a brood and care f o r the j u v e n i l e s . The t e r r i t o r y of male 27 ( F i g u r e 12) on the other hand, overlapped e x t e n s i v e l y on the t e r r i t o r i e s of males 12 and 28 d u r i n g the e a r l y breeding p e r i o d . He a l s o was shy and seldom sang, but was observed to s i n g on both t e r r i t o r i e s . However, he was not observed c o u n t e r s i n g i n g with the t e r r i t o r y h o l d e r s . He d i d not appear to a c q u i r e a mate although both other males were mated and he disappeared about mid-breeding p e r i o d while the other males p e r s i s t e d to the end of the study at the end of the breeding p e r i o d . Male 27 may have been a c t i v e only when males 23 and 28 c o u l d be avoided, although he may a l s o not have been t e r r i t o r i a l and/or was t o l e r a t e d due to h i s l a c k of a g g r e s s i v e n e s s . I observed t r e s p a s s by neighbouring t e r r i t o r i a l males only three times d u r i n g the breeding p e r i o d and once d u r i n g the winter p e r i o d . Two of the breeding p e r i o d o b s e r v a t i o n s were of males caught i n nets while the other was of a male a p p a r e n t l y f o l l o w i n g a female. In view of the r e l a t i v e r a r i t y with which winter wrens were caught i n nets, t r e s p a s s may be more common than these few o b s e r v a t i o n s suggest. During the dry months of June, J u l y , and August when there was o f t e n no water i n the f o r e s t except i n S p r i n g Creek, t e r r i t o r i a l males remote 67 from the creek were o f t e n observed wet as i f they had been b a t h i n g . To v i s i t the creek to d r i n k and bath i n many cases would have n e c e s s i t a t e d t r e s p a s s . The t r e s p a s s i n g male observed d u r i n g the winter p e r i o d was a p p a r e n t l y e x p l o r i n g and r e t r e a t e d q u i c k l y when the t e r r i t o r y holder appeared. Tresspass i n winter should be d i s t i n g u i s h e d from communal winter r o o s t i n g when the h o s t i l i t y of defense must disappear or a t . l e a s t d i m i n i s h g r e a t l y . The evidence f o r the d i s c r e t e n e s s of male t e r r i t o r i e s i n the European wren i s c o n t r a d i c t o r y . Weso'lowski (1981) found that males stayed w i t h i n the boundaries of t h e i r t e r r i t o r i e s d u r i n g the whole s p r i n g ( A p r i l - J u l y ) with only s m a l l adjustments of borders between neighbouring males. Garson (1978) found a ge n e r a l l a c k of simultaneous o v e r l a p and noted only two i n t r u s i o n s of t e r r i t o r i a l males on the t e r r i t o r i e s of o t h e r s . Armstrong (1955) found o c c a s i o n a l s i l e n t t r e s p a s s d u r i n g the breeding p e r i o d and open t r a n s g r e s s i o n of t e r r i t o r i a l boundaries i n wi n t e r . On the other hand, K e n t i s h (1976) d i s t i n g u i s h e d t e r r i t o r i e s (defended) and home ranges (non-defensive a c t i v i t i e s ) f o r t e r r i t o r i a l males d u r i n g the breeding p e r i o d . He found that males foraged on one another's t e r r i t o r i e s and home ranges, although g r e a t e r o v e r l a p o c c u r r e d f o r home ranges than f o r t e r r i t o r i e s . 68 In summary, male winter wrens p a r t l y f u l f i l the c r i t e r i a of t e r r i t o r i a l i t y set out e a r l i e r i n t h i s c h a p t er. S i t e attachment t o l o c a l i z e d areas was demonstrated throughout the year. Defense through a t t a c k and escape was shown f o r the breeding p e r i o d , although a d m i t t e d l y f o r only a few males, but was not shown f o r the f a l l and winter p e r i o d s . Defense through avoidance of one another's l o c a l i z e d areas o c c u r r e d throughout the year. Thus, male winter wrens f u l f i l a l l the c r i t e r i a f o r t e r r i t o r i a l i t y d u r i n g the breeding p e r i o d but l a c k the c r i t e r i a of a t t a c k and escape f o r the f a l l and winter p e r i o d s . However, i n my o p i n i o n the complete l a c k of o v e r l a p of l o c a l i z e d areas d u r i n g the f a l l and winter p e r i o d s i s such s t r o n g c i r c u m s t a n t i a l evidence of t e r r i t o r i a l defense through avoidance that I suggest male winter wren t e r r i t o r i a l i t y d u r i n g these p e r i o d s as w e l l . F i n a l l y , I suggest that the non-random and non-overlapping p a t t e r n of spacing of the male winter wren t e r r i t o r i e s as shown i n F i g u r e s 4 to 13 i s a r e s u l t of the t e r r i t o r i a l i t y of male winter wrens. 69 Females The area and extent of the home ranges of c o l o u r -banded females determined on the study p l o t s are shown from the bree d i n g p e r i o d 1979, through the' breeding p e r i o d 1980, i n F i g u r e s 14 to 17. Only three home ranges were determined f o r the breeding p e r i o d 1979, although other females, both colour-banded and unbanded were present on and around the study p l o t s . A l l the colour-banded females of the study p l o t s of the breeding p e r i o d 1979 disappeared d u r i n g the l a t e breeding and e a r l y f a l l p e r i o d s although three reappeared the f o l l o w i n g b r e e d i n g p e r i o d . The one colour-banded female present on the study p l o t s d u r i n g the f a l l and winter p e r i o d s 1979-80 a l s o appeared to occupy a home range although she was observed very seldom. She moved to the study p l o t s d u r i n g the e a r l y f a l l p e r i o d from o u t s i d e the study a r e a . However, based on c o l o u r bandings at the communal winter r o o s t s , other females were present on the study p l o t s , at l e a s t d u r i n g the winter p e r i o d . Most but not a l l female home ranges were determined on the study p l o t s f o r the breeding p e r i o d , 1980. A l l of the colour-banded females o l d e r than young y e a r l i n g s and which were observed r e g u l a r l y were l o c a l i z e d . Attack and escape were never observed f o r females. Females were never observed to s i n g or otherwise FIGURE 14 AREA 1 AND EXTENT OF THE FEMALE HOME RANGES 2 ON THE STUDY PLOTS 3 DURING THE BREEDING PERIOD, 1979 1 Areas i n ha 2 Banded b i r d s a r e i d e n t i f i e d by c i r c l e d code numbers; ages a r e a l s o g i v e n : A - a d u l t ; U - age unknown 3 Study p l o t s A, B and C a r e shown i n dashed l i n e s on t h e st u d y a r e a to FIGURE 15 AREA AND EXTENT OF THE FEMALE HOME RANGES ON THE STUDY PLOTS DURING THE FALL PERIOD, 1979 FIGURE 16 AREA AND EXTENT OF THE FEMALE HOME RANGES ON THE STUDY PLOTS DURING THE WINTER PERIOD, 1979-80 CTl FIGURE 17 AREA AND EXTENT OF THE FEMALE HOME RANGES ON THE STUDY PLOTS DURING THE BREEDING PERIOD, 1980 78 a d v e r t i s e . However, the home ranges of colour-banded females d i d not g e n e r a l l y o v e r l a p d u r i n g the breeding p e r i o d . Nor were other colour-banded females f o r which there were i n s u f f i c i e n t o b s e r v a t i o n s to determine home ranges observed on other female home ranges. However, the numbers of o b s e r v a t i o n s of such b i r d s were very s m a l l . The home ranges of females d u r i n g the breeding p e r i o d s g e n e r a l l y l a y w i t h i n i n d i v i d u a l male t e r r i t o r i e s although some o v e r l a p between t e r r i t o r i e s was observed (1979 -F i g u r e s 9 and 14; 1980 - F i g u r e s 12 and 17). Observations of females o u t s i d e the i n d i v i d u a l male t e r r i t o r i e s u s u a l l y o c c u r r e d d u r i n g the e a r l y breeding p e r i o d before n e s t i n g , or while females were e s c o r t i n g j u v e n i l e s . Most t e r r i t o r i a l male winter wrens were c l e a r l y monogamous but a few were polygamous: three cases of bigamy and one case of trigamy were recorded. Polygamy as used here i s d e f i n e d as f o r a male being s i m u l t a n e o u s l y mated to two or more females d u r i n g c l u t c h accumulation, i n c u b a t i o n , or care of the n e s t l i n g s and j u v e n i l e s . Males and females were c o n s i d e r e d mated i f they c o n s o r t e d and the female nested on the male's t e r r i t o r y (although i n some cases the exact nest s i t e was not d i s c o v e r e d ) , and i f the male helped e s c o r t or care f o r the n e s t l i n g s and/or j u v e n i l e s on h i s t e r r i t o r y . 79 Although polygamy d i d not occur on the study p l o t s proper, two t e r r i t o r i a l males there had second mates o u t s i d e the study p l o t s . These two polygamous males, rep r e s e n t e d polygamy r a t e s among t e r r i t o r i a l males on the study p l o t s of 8% (1 i n 13) i n both 1979 and 1980. However, these r a t e s are probably c o n s e r v a t i v e as polygamy was suspected f o r one other male i n 1979 and two other males i n 1980. In a l l cases of polygamy found i n t h i s study, the nest of o n l y one female of any polygamous male was found w h i l e the l o c a t i o n s of the other n e s t s were known only approximately. I t was t h e r e f o r e i m p o s s i b l e to c a l c u l a t e the exact time i n t e r v a l or d i s t a n c e between nests on the i n d i v i d u a l t e r r i t o r i e s . I t appeared however, based on o b s e r v a t i o n s of the j u v e n i l e s , t h a t winter wren polygamy i s s u c c e s s i v e as the nests were separated by approximately one to three weeks and that polygamous males probably a c q u i r e mates a d d i t i v e l y . K l u i j v e r et a l . (1940), Armstrong (1955), and Armstrong and Whitehouse (1977) found t h a t European wrens were extremely polygamous. K l u i j v e r et a l . rep o r t e d r a t e s of polygamy i n c l u d i n g both bigamy and trigamy at almost 50% of t e r r i t o r i a l males, which g r e a t l y exceeded that of t h i s study. Haneda and Kosakai (1971) a l s o found 80 s i m i l a r l y h i g h r a t e s of polygamy i n c l u d i n g tetragamy i n the Japanese wren. Wesolowski (1981) on the other hand r e p o r t e d lower r a t e s of polygamy (below 10%) f o r the European wren i n Poland. In summary, female winter wrens do not f u l f i l the c r i t e r i a f o r t e r r i t o r i a l i t y set out e a r l i e r i n t h i s c h a p t e r . S i t e attachment to l o c a l i z e d areas was demonstrated f o r the breeding p e r i o d but was not c o n c l u s i v e l y shown over the r e s t of the year. Only one colour-banded female f o r which there were few o b s e r v a t i o n s was observed d u r i n g the f a l l and winter p e r i o d s . She appeared l o c a l i z e d d u r i n g both p e r i o d s . Attack and escape were not observed, nor were females observed to s i n g or otherwise a d v e r t i s e . Areas of female l o c a l i z a t i o n d i d not o v e r l a p d u r i n g the breeding p e r i o d . However, because females were l o c a l i z e d w i t h i n male t e r r i t o r i e s , t h i s apparent mutual avoidance and spacing of females may have been due to the t e r r i t o r i a l i t y of males. T h e r e f o r e , I r e f e r t o the areas of female l o c a l i z a t i o n as home ranges. On the other hand, i t appeared t h a t the spacing of nests used s i m u l t a n e o u s l y f o r breeding w i t h i n the t e r r i t o r i e s of polygamous males was not random, but r a t h e r t h a t the nests were widely i f not maximally spaced. F u r t h e r , the home ranges of females w i t h i n the t e r r i t o r i e s 81 of polygamous males, d i d not o v e r l a p . Because i t was only p o s s i b l e to i d e n t i f y i n d i v i d u a l unbanded females while they were fee d i n g young at the nest, the numbers of o b s e r v a t i o n s f o r unbanded females were very s m a l l . Females g e n e r a l l y used nests b u i l t f o r them by males ( e i g h t nests observed b u i l t by males and l a t e r used, and f i v e n e s t s b u i l t but not used). Second n e s t s were b u i l t by both monogamous and polygamous males, u s u a l l y a f t e r a f i r s t mate had been o b t a i n e d and i n c u b a t i o n had begun. These second nests were u s u a l l y remote from the f i r s t nest w i t h i n the t e r r i t o r y . Thus, the a d d i t i o n a l mates of a polygamous male may be spaced by the male h i m s e l f , although females may a l s o be a b l e to space themselves by choosing nests remote from other females. U n l i k e the female European wren ( K l u i j v e r 1940; Armstrong 1955; Garson 1978) however, female winter wrens a l s o b u i l t and used t h e i r own nests (two nests observed b u i l t and used, and one nest b u i l t but not used). Females may thus be a b l e to choose t h e i r own nest s i t e s independently of the t e r r i t o r i a l male. At l e a s t two a c t i v e nests on the study area and two on and around removal p l o t E were c l e a r l y o u t s i d e male t e r r i t o r i e s , both i n terms of o b s e r v a t i o n s of the nearest t e r r i t o r i a l male and i n terms of h a b i t a t that was o r d i n a r i l y u t i l i z e d by males f o r t h e i r t e r r i t o r i e s . 82 These females a l s o may have b u i l t t h e i r own n e s t s . In a l l four cases the nearest t e r r i t o r i a l males appeared to have mates, who may have been a f a c t o r i n the remote l o c a t i o n of these females and t h e i r n e s t s . I suggest that t h i s s l e nder evidence of mutual avoidance by females, independent of male t e r r i t o r i e s , supports the p o s s i b i l i t y of female t e r r i t o r i a l i t y d u r i n g the breeding p e r i o d . F i n a l l y , i f females are t e r r i t o r i a l , then the non-random and non-overlapping spacing of female home ranges as shown i n F i g u r e s 14 and 17 may be a r e s u l t of t h e i r own as w e l l as male t e r r i t o r i a l behaviour. D i s t r i b u t i o n of Winter Wrens Odum and Kuenzler (1955) proposed the one percent, l e v e l f o r the v a l i d d e t e r m i n a t i o n of t e r r i t o r y and home range s i z e . At that l e v e l , each a d d i t i o n a l o b s e r v a t i o n p r o v i d e s l e s s than an a d d i t i o n a l one percent i n c r e a s e i n area. The areas (ha) of the i n d i v i d u a l male t e r r i t o r i e s and female home ranges as estimated on the study area, study p l o t s , and removal p l o t s d u r i n g the winter breeding and f a l l p e r i o d s are shown i n F i g u r e s 4 to 17. Only the . areas of the male t e r r i t o r i e s as determined on the study p l o t s d u r i n g the breeding p e r i o d s f u l f i l l e d the one percent c r i t e r i o n : 1979 (1.38 ± 0.51; 0.65 - 2.21; 14), 83 and 1980 (1.23 ± 0.50; 0.48 - 2.16; 12), (X ± S.D.; range; n ) . The other estimates of t e r r i t o r y and home range s i z e are probably c o n s e r v a t i v e ( F i g u r e s 4 to 8, 10, 11, 13, and 14 to 17). Garson (1978) found average male European wren t e r r i t o r y s i z e s d u r i n g the breeding p e r i o d over the three years of h i s study, 1975 to 1977, of 0.37 ha (n = 22), 0.85 ha (n = 6), and 0.62 ha (n = 14), r e s p e c t i v e l y , with an o v e r a l l range of 0.15 to 1.16 ha. These v a l u e s are s m a l l e r than those found i n t h i s study. The numbers of t e r r i t o r i a l male winter wrens on the study area d u r i n g the breeding p e r i o d s 1978-81 remained f a i r l y c o n s t a n t , ranging from 21 to 24 (54 - 62 per km 2) (Table 6). Although the t o t a l numbers of males present decreased d u r i n g the f a l l and winter p e r i o d s , the numbers of male t e r r i t o r i e s on the study p l o t s a l s o remained f a i r l y constant among the v a r i o u s p e r i o d s , ranging from 6.9 d u r i n g the breeding p e r i o d 1979, to 8.3 d u r i n g the breeding p e r i o d 1980 (57 to 68 per km2 r e s p e c t i v e l y ) . S i m i l a r l y , the numbers of male t e r r i t o r i e s and t e r r i t o r i a l males on the i n d i v i d u a l removal p l o t s d u r i n g the breeding p e r i o d s 1979-80, remained f a i r l y c o n s t a n t . TABLE 6 NUMBERS OF TERRITORIAL MALES AND MALE TERRITORIES ON THE STUDY AREA, STUDY PLOTS, AND REMOVAL PLOTS Year P e r i o d Study Area A Study P l o t s B C T o t a l (A,B,C) Removal P l o t s D E 1978 Breeding 22J22; 1 57 1979 Winter 0;0 4;3.1 4;4.0 8;7.1; 58 Breeding 21;21; 54 4;0.7 6;3.0; 7;3.2 13;6.9; 57 5;3.6 2;1.6 F a l l 0;0 6;3.6 6;3.8 11;7.4 61 1980 Winter 0;0 6;3.3 5;4.5 9;7.8 64 Breeding 21;21; 54 2;0.7 7;3.2 8;3.5 13;8.3 68 5;4.0 1;0.9 1981 Breeding 24;24; 62 Number o f t e r r i t o r i a l males; c a l c u l a t e d number o f male t e r r i t o r i e s ; c a l c u l a t e d d e n s i t y o f male t e r r i t o r i e s per km. C a l c u l a t i o n s per km 2 were based on the f o l l o w i n g areas: study area - 386,520m study p l o t s A, B and C - 121,941 m2. 85 The breeding b i r d d e n s i t i e s of B r i t i s h Columbia west coast f o r e s t s were reviewed by Buckner et a l . (1975). Winter wren d e n s i t i e s (per km2) were given f o r mature f o r e s t s as 41 f o r p l o t s near Port A l i c e (Buckner at a l . 1975), 49 f o r a p l o t near T e r r a c e (Webster 1969), 37 f o r a p l o t near Goldstream (Edwards and S t i r l i n g 1962, 1963) and 99 f o r a p l o t near Hope (Horvath 1963). These d e n s i t i e s g e n e r a l l y agree with the d e n s i t i e s found i n t h i s study. Winter wren d e n s i t i e s i n medium aged f o r e s t on a p l o t near M i r a c l e Beach averaged 49 (Edwards et a l . 1962; S t i r l i n g and Westerborg 1963; Campbell and Westerborg 1964) . D e n s i t i e s on p l o t s i n immature f o r e s t s near Port A l i c e (Buckner et a l . 1975) averaged 31, while Dow and Dow (1963) found winter wrens only as v i s i t o r s on a p l o t i n very young, open canopy western hemlock f o r e s t i n Golden Ears P r o v i n c i a l Park w i t h i n 5 km of t h i s study. E r s k i n e (1977) reviewed the b i r d p o p u l a t i o n s of the Canadian b o r e a l f o r e s t g e n e r a l l y . He gave d e n s i t i e s (per km2) of t e r r i t o r i a l males of 55 i n western c o a s t a l f o r e s t s . Winter wren d e n s i t i e s were g e n e r a l l y much lower elsewhere with maximum d e n s i t i e s as f o l l o w s : 13 i n spruce f o r e s t s ; 22 i n balsam f i r f o r e s t s ; and 14 i n other c o n i f e r o u s f o r e s t s . I t would appear from the p r e c e d i n g data, that i n Canada winter wrens are most numerous and do 86 best i n mature- (as i n t h i s study) and perhaps medium-aged west coast f o r e s t s . The lower ranges of the d e n s i t i e s of t e r r i t o r i a l males found i n two of three i n t e n s i v e s t u d i e s of the European wren were comparable to those found i n t h i s study while the upper ranges were much h i g h e r . K l u i j v e r et a l . (1940) worked with a p o p u l a t i o n of 8 to 12 on a study area of about 7 ha (114-171 per km 2). Armstrong (1955) found from 1 t o 3 t e r r i t o r i a l males on a study area of about 1.6 ha (63 - 188 per km 2). Garson (1978) found 6 to 22 on h i s study area of about 8.5 ha (71 to 259 per km 2). D e n s i t i e s of t e r r i t o r i a l male European wrens per km2 f o r these authors are my c a l c u l a t i o n s . Although the d e n s i t i e s of male t e r r i t o r i e s d u r i n g the breeding p e r i o d s on the study area and the study p l o t s together were very s i m i l a r both among and w i t h i n y e a r s , c o n s i s t e n t l y fewer, t e r r i t o r i a l males and male t e r r i t o r i e s were observed on study p l o t A as opposed to study p l o t s B and C (Table 6 ) . The numbers of male t e r r i t o r i e s on study p l o t A were 0.7 d u r i n g both breeding p e r i o d s while the p l o t was d e s e r t e d d u r i n g the f a l l and winter p e r i o d s . On the other hand, the numbers of male t e r r i t o r i e s on study p l o t s B and C ranged from 3.0 to 3.6 and 3.2 to 4.5, r e s p e c t i v e l y , throughout the v a r i o u s p e r i o d s . S i m i l a r l y , 87 the numbers of t e r r i t o r i a l males and male t e r r i t o r i e s on removal p l o t E d u r i n g the breeding p e r i o d were c o n s i s t e n t l y l e s s than those of removal p l o t D. In summary, t e r r i t o r i a l males and male t e r r i t o r i e s were not d i s t r i b u t e d u n i f o r m l y over the study area or among the study and removal p l o t s . Empty and s p a r s e l y occupied areas o c c u r r e d ( F i g u r e s 4 to 13). Removal Experiments The e x i s t e n c e of e x t r a , p o t e n t i a l l y t e r r i t o r i a l males i n b i r d p o p u l a t i o n s has o f t e n been demonstrated (e.g., Stewart and Aldrich 1951; Hensley and Cope 1951; Brown 1969; Knapton and Krebs 1974; Edwards 1977; Smith 1978; Wesolowski 1981). Such males are g e n e r a l l y assumed to represent a s u r p l u s " f l o a t i n g " p o p u l a t i o n from which l o s s e s of t e r r i t o r i a l males can be r e p l a c e d . I removed t e r r i t o r i a l male winter wrens from the removal p l o t D and E d u r i n g the breeding p e r i o d t o i n v e s t i g a t e the p o s s i b l e a v a i l a b i l i t y of e x t r a , p o t e n t i a l l y t e r r i t o r i a l males. G e n e r a l l y , the removed t e r r i t o r i a l males were r e p l a c e d by other t e r r i t o r i a l males (Table 7 ) . The numbers of the replacements i n n e a r l y a l l cases appeared to be about equal t o , or s l i g h t l y fewer than the numbers of the removed t e r r i t o r i a l males. TA3LE 7 Removal P l o t REMOVAL AND REPLACEMENT OF TERRITORIAL MALES ON AND AROUND THE REMOVAL PLOTS D AND E DURING THE BREEDING PERIODS, 1979 AND 1980 O r i g i n a l Population 1 F i r s t Removal (24-25 A p r i l ) Replacements Second Removal (13-16 May) Replacements 1979 4 Year O r i g i n a l Population F i r s t Removal (14 A p r i l ) Replacements Second Removal (13 May) Replacements T h i r d Removal (23-24 May) Replacements Fourth Removal (24-29 J u l y ) 1980 S 2? 4 s i n g i n g young y e a r l i n g s O r i g i n a l Population 2 F i r s t Removal (17 May) 2 Replacements 2 Second Removal (27 May - 1 June) 2 O r i g i n a l Population F i r s t Removal (16-18 A p r i l ) Replacements Second Removal (12 May) Replacements Third Removal (23-28 June) Replacements Fourth Removal (21-24 J u l y ) co co O r i g i n a l populations are shown i n Figure 13. 89 Removal p l o t D was l o c a t e d i n w e l l - p o p u l a t e d winter wren h a b i t a t and had o r i g i n a l p o p u l a t i o n s of four and f i v e t e r r i t o r i a l males duri n g the e a r l y breeding p e r i o d s of 1979 and 1980 r e s p e c t i v e l y (Table 7; F i g u r e 13). I t was a d j o i n e d on two s i d e s by s i m i l a r h a b i t a t which c o n t a i n e d other t e r r i t o r i a l male winter wrens. On the second day of the f i r s t removal, 1979, an e x t r a s i n g i n g male, probably a neighbour, was shot on the t e r r i t o r y of a b i r d shot the day b e f o r e . S i m i l a r l y i t appeared that s e v e r a l of the f i r s t replacements i n both 1979 and 1980 were a l s o neighbouring b i r d s which had expanded t h e i r t e r r i t o r i e s or at l e a s t v i s i t e d the emptied t e r r i t o r i e s of the removal p l o t . When these b i r d s sang t h e r e was no c o u n t e r s i n g i n g from the a d j o i n i n g area beside the removal p l o t as there had been with the o r i g i n a l p o p u l a t i o n . In a d d i t i o n , when chased d u r i n g s t a l k i n g these b i r d s f l e d to areas o u t s i d e the removal p l o t and were shot t h e r e . I t i s l i k e l y that the s i n g i n g of the v i s i t i n g , annexing, and/or neighbouring males on and around the removal p l o t s was i n t i m i d a t i n g enough to prevent or r e t a r d s e t t l i n g by new males u n t i l the neighbouring males themselves were removed. In r e t r o s p e c t i t would have been b e t t e r to have had l a r g e r removal p l o t s , such as those of Hensley and Cope (1951) -40 ac (16.2 ha) and Wesolowski (1981) - 48 ha, so as to 90 reduce the e f f e c t of neighbouring males, and to have c o l o u r banded both the males on the p l o t s (Wesolowski 1981) as w e l l as the neighbouring males before removing b i r d s from the p l o t as a check on v i s i t i n g and annexation. The second replacement i n 1979 and the second and t h i r d replacements i n 1980 on removal p l o t D probably i n c l u d e d very few neighbouring b i r d s as they had e i t h e r been shot or c o u l d be heard s i n g i n g on t h e i r t e r r i t o r i e s around the removal p l o t . The t h i r d replacement i n June, 1980 appeared t o be composed of o n l y two b i r d s n e i t h e r of which appeared very l o c a l i z e d on the p l o t . I t may be that with the removal of a l l t e r r i t o r i a l males from the p l o t and immediate surrounding area, the replacement b i r d s c o u l d move with impunity over much l a r g e r areas, and h e l d , or appeared t o h o l d , very l a r g e t e r r i t o r i e s . Both these b i r d s were almost c e r t a i n l y a d u l t s or o l d e r y e a r l i n g s but d u r i n g the f o u r t h removal i n l a t e J u l y , n e i t h e r of these replacement b i r d s was encountered. They may have l e f t the p l o t or c o u l d p o s s i b l y have entered moult and been s i l e n t . Rather, d u r i n g the f o u r t h removal, four s i n g i n g , young y e a r l i n g s were shot. They were spaced about the p l o t about as evenly as t e r r i t o r i a l males had been i n e a r l i e r removals. I t appeared that the four s i n g i n g young y e a r l i n g s were r e p l a c e d subsequently by other s i n g i n g 91 y e a r l i n g s , but as young y e a r l i n g s sang i n f r e q u e n t l y and were very shy and seldom observed, f u r t h e r d e t e r m i n a t i o n of i n d i v i d u a l replacement and removals were i m p o s s i b l e . Removal p l o t E had o r i g i n a l p o p u l a t i o n s of two and one t e r r i t o r i a l males dur i n g the e a r l y breeding p e r i o d of 1979 and 1980 r e s p e c t i v e l y (Table 7; F i g u r e 13). It was composed mostly o f , and a d j o i n e d by, h a b i t a t s t h a t were not g e n e r a l l y u t i l i z e d by winter wrens on the study area, study p l o t s , and elsewhere on the Research F o r e s t . In both 1979 and 1980 no neighbouring t e r r i t o r i a l males were heard on three s i d e s of the p l o t , w h ile the c l o s e s t t e r r i t o r y boundary to the south was at about 100 m d i s t a n t and a c r o s s a'road. T h i s neighbouring male may have been one of the two b i r d s taken i n the second removal i n 1979. However, the other replacements i n both years were almost c e r t a i n l y not neighbours. R e l a t i v e to most other t e r r i t o r i a l males, the t e r r i t o r i a l males of the t h i r d and f o u r t h removals on removal p l o t E i n 1980 were unaggressive and shy, both to the observer and to recorded song. Although these replacements were both a d u l t s or o l d e r y e a r l i n g s , they sang weakly and i n f r e q u e n t l y . T h i s l a c k of a g g r e s s i o n may have c o n t r i b u t e d to t h e i r i n a b i l i t y to o b t a i n t e r r i t o r i e s e a r l i e r i n the breeding p e r i o d . S i n g i n g young y e a r l i n g s 92 were not observed as replacements on removal p l o t E. On the other hand, the four colour-banded t e r r i t o r i a l males which disappeared May through e a r l y J u l y from on and around the study p l o t s were not r e p l a c e d by t e r r i t o r i a l a d u l t s and o l d e r y e a r l i n g s . Other t e r r i t o r i a l males that a p p a r e n t l y disappeared May through J u l y on the r e s t of the study area a l s o were not r e p l a c e d immediately, although replacement of some of these m i s s i n g b i r d s by s i n g i n g young y e a r l i n g s began i n August. However, a l l the t e r r i t o r i a l males which disappeared from the study p l o t s and study area were surrounded d i r e c t l y by other s u r v i v i n g t e r r i t o r i a l males. In a l l such cases of disappearance, the neighbouring t e r r i t o r i a l males annexed, or at l e a s t v i s i t e d and sang on, a d j o i n i n g areas of the empty t e r r i t o r i e s , and may have prevented new males from r e p l a c i n g the absent males. These o b s e r v a t i o n s are s i m i l a r t o , and support, those made durin g the f i r s t removals on the removal p l o t s . The apparent expansion of neighbouring t e r r i t o r i e s upon the disappearance or removal of an i n d i v i d u a l t e r r i t o r i a l male suggest that the t e r r i t o r i e s were s l i g h t l y compressed (Huxley 1934). P r e v e n t i o n of the establishment of new t e r r i t o r i a l males by neighbouring males may reduce s t r i f e w i t h i n the p o p u l a t i o n d u r i n g the mid- and l a t e - b r e e d i n g p e r i o d while the young 93 are being tended. Appearance of new males at t h i s time, such as o c c u r r e d i n e a r l y March by male 24 among f i v e neighbours ( F i g u r e 12), would l i k e l y have been extremely d i s r u p t i v e and c o u n t e r p r o d u c t i v e to the breeding e f f o r t s of i n d i v i d u a l males. Elsewhere i n North America, Stewart and A l d r i c h (1951), found no replacement of two male e a s t e r n winter wrens ( T . t . h i e m a l i s ) d u r i n g g e n e r a l b i r d removal s t u d i e s . However, the next year on the same p l o t , Hensley and Cope (1951) found that the o r i g i n a l p o p u l a t i o n of 1.5 males was r e p l a c e d by an a d d i t i o n a l 3.5 males over three c o l l e c t i n g p e r i o d s ( f r a c t i o n s of b i r d s r e f e r t o f r a c t i o n s of t e r r i t o r i e s w i t h i n the p l o t ) . The replacement b i r d s were co n s i d e r e d as pa r t of a s u r p l u s " f l o a t i n g " p o p u l a t i o n . Wesolowski (1981) demonstrated p o s t - m i g r a t o r y replacement of t e r r i t o r i a l male European wrens d u r i n g the breeding p e r i o d . H i s f i r s t removal of seven t e r r i t o r i a l males i n m i d - A p r i l r e s u l t e d i n replacement by f i v e new males while the second removal of s i x males ( i n c l u d i n g the replacements) r e s u l t e d i n a f u r t h e r replacement by f i v e new males. Wesolowski suggested t h a t a non-breeding element of e x t r a , p o t e n t i a l l y t e r r i t o r i a l males was present i n h i s p o p u l a t i o n . 94 S i m i l a r l y , I suggest that the replacement of t e r r i t o r i a l males removed from p l o t s i n t h i s study i n d i c a t e s that e x t r a , p o t e n t i a l l y t e r r i t o r i a l males were a v a i l a b l e i n the p o p u l a t i o n d u r i n g the e a r l y and mid-breeding p e r i o d . However, i n c o n t r a s t to Wesolowski who thought a l l h i s replacements were po s t - m i g r a t o r y , i n t h i s study some of the replacement b i r d s d u r i n g e a r l y s p r i n g may gave been migrants w a i t i n g f o r the higher mountain f o r e s t s and the i n t e r i o r f o r e s t s t o c l e a r of snow. At that time male winter wrens were o f t e n observed s i n g i n g i n f o r e s t h a b i t a t s t h a t , as the s p r i n g advanced, became s i l e n t and empty as i f these b i r d s had moved on. On the other hand, some of the replacement b i r d s may have been p a r t of a r e s i d e n t , f l o a t i n g p o p u l a t i o n of e x t r a , p o t e n t i a l l y t e r r i t o r i a l male winter wrens. In e i t h e r case, however, e x t r a , t e r r i t o r i a l male winter wrens were a v a i l a b l e d u r i n g the breeding p e r i o d . These e x t r a , t e r r i t o r i a l males c o u l d have occupied the empty and s p a r s e l y occupied areas of the study area, study p l o t s , and removal p l o t s . 9 5 WINTER WREN DISPERSION AS A FUNCTION OF HABITAT In t h i s c hapter I demonstrate that winter wrens show a d i f f e r e n t i a l use of the v a r i o u s i n d i v i d u a l ecosystems and of ecosystems grouped by f o r e s t f l o o r v e g e t a t i o n . I propose that winter wrens are clumped by s u i t a b l e h a b i t a t . F o l l o w i n g from t h i s , I propose that ecosystems and grouped ecosystems (sensu K l i n k a 1976) are i n d i c a t i v e of the d i s t r i b u t i o n of winter wrens. F i n a l l y , I demonstrate that the volume of p o t e n t i a l winter wren food i s r e l a t i v e l y g r e a t e r i n the grouped ecosystems p r e f e r r e d by winter wrens and propose that the abundance of food i s a f a c t o r i n winter wren d i s p e r s i o n . Winter Wren D i s p e r s i o n As A F u n c t i o n Of K l i n k a ' s Ecosystems And Ecosystems Grouped By F o r e s t F l o o r H a b i t a t s „ I n t r o d u c t i o n K l i n k a (1976), and K l i n k a and Skoda (1977), b u i l d i n g on the ideas and work of Sukachev (1944), Sukachev and D y l i s (1968), K r a j i n a (1960, 1965a, b, 1969, 1972), Lesko (1961), Lacate (1965), O r l o c i (1961, 1964, 1965), Brooke et a l . (1970), and Kojima and K r a j i n a (1975) d e v i s e d a s y n e c o l o g i c a l c l a s s i f i c a t i o n and map f o r the c o a s t a l 96 western hemlock f o r e s t of the Research F o r e s t . . The i n d i v i d u a l s i n the s y n e c o l o g i c a l c l a s s i f i c a t i o n are the b a s i c ecosystems or biogeocoenoses ( K l i n k a 1976), or as they are now known, ecosystem types (e.g., I n s e l b e r g et a l . 1982) but which h e r e a f t e r are r e f e r r e d to simply as ecosystems (Table 8 ) . In a d d i t i o n , the term ecosystem a l s o a p p l i e s to grouped, s i m i l a r ecosystems at the l e v e l of ecosystem s u b - a s s o c i a t i o n (Table 8 ) . Sukachev and D y l i s (1968), as t r a n s l a t e d i n K l i n k a et a l . (1979, p.2), d e f i n e d a f o r e s t ecosystem as f o l l o w s : "that p a r t of the f o r e s t uniform over a c e r t a i n area i n the composition, s t r u c t u r e , and p r o p e r t i e s of i t s components, and i n the i n t e r r e l a t i o n s h i p s among them; that i s , uniform i n the p l a n t s , animals, and microorganisms i n h a b i t i n g i t , i n the parent m a t e r i a l , i n i t s h y d r o l o g i c a l , m i c r o c l i m a t i c (atmospheric), and s o i l environments and the i n t e r a c t i o n s among them; and i n the kind of matter and energy exchange between these components and other n a t u r a l phenomena i n nature". Ecosystems were grouped h i e r a r c h i c a l l y by K l i n k a i n t o ecosystem a s s o c i a t i o n s which are the b a s i c taxon of h i s s y n e c o l o g i c a l c l a s s i f i c a t i o n system. A synopsis of the s y n e c o l o g i c a l c l a s s i f i c a t i o n of the c o a s t a l western hemlock f o r e s t of the Research F o r e s t ( s o i l parent m a t e r i a l omitted) i s shown i n Appendix 1. A summary of the K l i n k a s y n e c o l o g i c a l u n i t s of the study area and removal p l o t s i s shown i n Appendix 2. TABLE 8 RELATIONSHIPS AMONG THE TAXA OF THE LINNAEAN, CANADIAN SOIL, VEGETATION, AND SYNECOLOGICAL CLASSIFICATION SYSTEMS ( a f t e r K l i n k a (1976), and K l i n k a and Skoda (1977)) .Linnaean C l a s s i f i c a t i o n Canadian S o i l C l a s s i f i c a t i o n V e g e t ation C l a s s i f i c a t i o n S y n e c o l o g i c a l C l a s s i f i c a t i o n Species S o i l S e r i e s P l a n t A s s o c i a t i o n Ecosystem A s s o c i a t i o n I n d i v i d u a l Organism I n d i v i d u a l Organism Polypedon Pedon P l a n t Community Sample P l o t Ecosystem Su b - a s s o c i a t i o n Ecosystem (Biogeocoenose) Sample P l o t ( i n c l u d i n g pedons) 98 To i n t e r p r e t the 1976 K l i n k a c l a s s i f i c a t i o n system at the ecosystem l e v e l , i t should be noted that i n d i v i d u a l ecosystems and as grouped at the l e v e l of ecosystem s u b - a s s o c i a t i o n are w r i t t e n p a r t l y i n lower case l e t t e r s while the ecosystem a s s o c i a t i o n s are w r i t t e n e n t i r e l y i n upper case l e t t e r s . Where an ecosystem a s s o c i a t i o n i s not d i s t i n g u i s h e d i n t o two or more subordinate ecosystems, the ecosystem a s s o c i a t i o n name a l s o serves as t h a t of the ecosystem s u b - a s s o c i a t i o n but i s s t i l l w r i t t e n i n upper case l e t t e r s . On maps, the ecosystems are l a b e l l e d by a numerical symbol i d e n t i f y i n g the ecosystem a s s o c i a t i o n or s u b - a s s o c i a t i o n and the a s s o c i a t e d s o i l parent m a t e r i a l s as i n the ecosystem s u b - a s s o c i a t i o n G a u l t h e r i a - WH - DF, 131.30 (Appendix 2 ) . The s y n e c o l o g i c a l c l a s s i f i c a t i o n i s g e n e t i c and h i e r a r c h i c a l ( K l i n k a 1976) and i s a n a t u r a l c l a s s i f i c a t i o n ( L a v k u l i c h 1972). On the other hand, the v a r i o u s u n i t s of any s y n e c o l o g i c a l taxa and/or the taxa themselves may be c o n s t i t u t e d , named, and arranged to s u i t s p e c i f i c o b j e c t i v e s . T h e r e f o r e , under these circumstances the s y n e c o l o g i c a l c l a s s i f i c a t i o n can be i n t e r p r e t i v e r a t h e r than n a t u r a l ( L a v k u l i c h 1972). S p e c i f i c a l l y , the taxa of the present s y n e c o l o g i c a l c l a s s i f i c a t i o n (Appendix 1) have been i n t e r p r e t i v e l y o r g a n i z e d and named to serve the 99 o b j e c t i v e s of botany and f o r e s t r y . For example, K l i n k a has regrouped h i s ecosystems to serve the o b j e c t i v e s of f o r e s t management ( K l i n k a 1976, p.96; K l i n k a e t a l . 1980, p.21) and, i n p a r t i c u l a r , s p e c i e s s e l e c t i o n and p r e s c r i b e d burning ( K l i n k a 1977). The d i s p e r s i o n of the European wren (e.g., W i l l i a m s o n 1969a, b, 1974; F e r r y and Frochot 1970; F e r r y 1974; T o m i a l o j c ' e t a l . 1977; Walankiewicz 1977), the Japanese wren (Nakamura 1974, 1975) and the winter wren (e.g. Horvath 1963; Roe 1974; Buckner et a l . 1975; E r s k i n e 1977; S c o u l l a r 1980) has long been c o n s i d e r e d to be a f u n c t i o n of h a b i t a t . In t h i s study, I compare winter wren d i s t r i b u t i o n with the d i s t r i b u t i o n of ecosystems (sensu K l i n k a , 1976). I a l s o compare winter wren d i s t r i b u t i o n w i t h the d i s t r i b u t i o n of ecosystems which have been grouped i n t e r p r e t i v e l y a c c o r d i n g to the dominant f o r e s t f l o o r v e g e t a t i o n . T h i s grouping i s done i n the same gene r a l way t h a t K l i n k a grouped h i s ecosystems a c c o r d i n g to c r i t e r i a of f o r e s t management. Hilde'ri ( 1965) a p p l i e d d i s t i n c t i o n s drawn by Baker (1938) and c a t e g o r i z e d the f a c t o r s determining h a b i t a t s e l e c t i o n i n b i r d s as u l t i m a t e and proximate. U l t i m a t e f a c t o r s are e s s e n t i a l to the s u r v i v a l of the s p e c i e s and are the u n d e r l y i n g reasons f o r h a b i t a t s e l e c t i o n by b i r d s . 100 They i n c l u d e : food, s h e l t e r from enemies and adverse weather, and the requirements imposed by the s t r u c t u r a l and f u n c t i o n a l c h a r a c t e r i s t i c s of the s p e c i e s . Proximate f a c t o r s on the other hand are the s t i m u l a e , both p o s i t i v e and n e g a t i v e , by which b i r d s s e l e c t h a b i t a t and i n c l u d e : landscape, t e r r a i n , nest-, song-, look-out-, f e e d i n g - , and d r i n k i n g - s i t e s , other animals, food, and i n t e r n a l m o t i v a t i o n . Proximate f a c t o r s are d i f f i c u l t to d i s t i n g u i s h from u l t i m a t e f a c t o r s . Proximate f a c t o r s are a l s o g e n e r a l l y more s p e c u l a t i v e and l e s s amenable to study than u l t i m a t e f a c t o r s and t h e r e f o r e are not as w e l l p e r c e i v e d and understood (Hilde'n 1965). However, t h i s s e p a r a t i o n of f a c t o r s i s probably somewhat a r t i f i c i a l as most f a c t o r s appear to a c t i n c o n c e r t and to p r o v i d e a continuum of i n f l u e n c e s on h a b i t a t s e l e c t i o n . In t h i s study, without s e p a r a t i n g proximate and u l t i m a t e f a c t o r s , I compare gen e r a l a t t r i b u t e s of the cover and nature of the f o r e s t f l o o r v e g e t a t i o n of the v a r i o u s ecosystems and grouped ecosystems as p o s s i b l e e x p l a n a t i o n s f o r winter wren d i s p e r s i o n . I a l s o compare the volumes of i n v e r t e b r a t e s on the v a r i o u s f o r e s t f l o o r h a b i t a t s as a f u r t h e r p o s s i b l e f a c t o r i n winter wren d i s p e r s i o n . 101 The d e f i n i t i o n of ecosystem (Sukachev and D y l i s 1968) used i n t h i s study (p.96) p r e s c r i b e s c h a r a c t e r i s t i c u n i f o r m i t y w i t h i n i n d i v i d u a l ecosystems f o r p l a n t s , animals, microorganisms, and parent m a t e r i a l i n i t s h y d r o l o g i c a l , m i c r o c l i m a t i c and s o i l environments. Of these a t t r i b u t e s , K l i n k a (1976) used only p l a n t s , parent m a t e r i a l s , s o i l s , and h y d r o l o g i c and n u t r i e n t regimes to d e f i n e h i s ecosystems. However, i t has always been assumed t h a t a s i m i l a r , c h a r a c t e r i s t i c u n i f o r m i t y a l s o occurs f o r the animal component i n a broad sense w i t h i n these ecosystems (K. K l i n k a , p e r s . comm.). To my knowledge such u n i f o r m i t y has never been shown. In my comparisons of winter wren d i s t r i b u t i o n with the d i s t r i b u t i o n of ecosystems and ecosystems grouped by the dominant f o r e s t f l o o r v e g e t a t i o n , I t e s t the u t i l i t y of K l i n k a ' s ecosystems as i n d i c a t o r s of the d i s t r i b u t i o n of one animal, the winter wren. Methods I remapped the study area d u r i n g the v e g e t a t i o n p l o t work at a working s c a l e of 1:787, us i n g K l i n k a ' s s y n e c o l o g i c a l c l a s s i f i c a t i o n and h i s 1:10,000 s c a l e map ( U n i v e r s i t y of B r i t i s h Columbia Research F o r e s t 1976). In most cases the mapping was s t r a i g h t f o r w a r d as the 102 t r a n s i t i o n from one ecosystem to another was r e l a t i v e l y abrupt, o c c u r r i n g over 5 m or l e s s . My remapping, with a few e x c e p t i o n s , g e n e r a l l y agreed with the o r i g i n a l K l i n k a mapping. The areas (m 2) of the i n d i v i d u a l ecosystems were c a l c u l a t e d by c o u n t i n g map squares (1 square = 4 m 2). I d e s c r i b e d the v e g e t a t i o n of the ecosystems of the study areas using v e g e t a t i o n cover p l o t s . Because i t was thought that v e g e t a t i o n was much more important to winter wren d i s p e r s i o n than s o i l parent m a t e r i a l s , the ecosystems i n the study area with s i m i l a r p l a n t communities but which d i f f e r e d only s l i g h t l y i n s o i l parent m a t e r i a l s were grouped (e.g., RUBUS - POLYSTICHUM - WRC, 72.4/72.47). These groupings by v e g e t a t i o n were used f o r d e s c r i p t i o n , and f o r l a t e r comparisons a g a i n s t winter wren d i s t r i b u t i o n . The ecosystems of the study area and study p l o t s were not grouped with the ecosystems of the removal p l o t s . I t i s important to note, however, that to maximize the p r e c i s i o n of mapping, the s o i l parent m a t e r i a l s and s o i l s a long with the other s y n e c o l o g i c a l f a c t o r s were c o n s i d e r e d to h e l p d e f i n e and map the ecosystems of the study a r e a . 103 V e g e t a t i o n p l o t s were o b j e c t i v e l y s e l e c t e d on the ecosystem map u n i t s to g i v e r e p r e s e n t a t i v e d e s c r i p t i o n s of the ecosystems as they o c c u r r e d on the study a r e a s . The l a r g e r ecosystem map u n i t s were d e s c r i b e d by f i v e v e g e t a t i o n p l o t s while s m a l l e r map u n i t s were d e s c r i b e d by three p l o t s . The r e s u l t i n g v e g e t a t i o n d e s c r i p t i o n s of the ecosystems are v a l i d only f o r the study a r e a s . The ecosystems as they apply to the Research F o r e s t g e n e r a l l y are d e f i n e d i n K l i n k a (1976). V e g e t a t i o n p l o t s i z e and shape v a r i e d and were o f t e n l i m i t e d by the extent of the ecosystems. P l o t s i z e was not g e n e r a l l y l e s s than 0.05 ha and was o f t e n as l a r g e as 1.0 ha. The v e g e t a t i o n p l o t s employed to estimate the t r e e and shrub l a y e r s were o f t e n much l a r g e r than those employed to d e s c r i b e the understory l a y e r s . A t o t a l of 52 p l o t s were sampled. Ten ecosystem map u n i t s were d e s c r i b e d f o r the study area while four were d e s c r i b e d f o r the removal p l o t s . Slope (percent) and aspect (compass bearing) were measured f o r each p l o t . V e g e t a t i o n cover (percent) was estimated v i s u a l l y f o r each p l a n t s p e c i e s and as a t o t a l f o r a l l p l a n t s p e c i e s combined i n the f o l l o w i n g v e g e t a t i o n l a y e r s : 104 A - o v e r s t o r y l a y e r A^ - dominant and co-dominant t r e e s A 2 - i n t e r m e d i a t e t r e e s A^ - suppressed t r e e s over 10 m T - t o t a l f o r A l a y e r B - m i d d l e s t o r y l a y e r B i - s m a l l t r e e s and shrubs between 2 and 10 m i n height B 2 - shrubs and woody p l a n t s between 0.15 and 2.0 m i n h e i g h t T - t o t a l f o r B l a y e r C - a l l herbaceous p l a n t s and a l l small woody p l a n t s under 0.15 m i n height D - bryophytes and l i c h e n s The e x c e p t i o n to t h i s scheme was salmonberry which, although o c c a s i o n a l l y higher than 2 m, was always c o n s i d e r e d i n the B 2 l a y e r . The t o t a l s (T) f o r v a r i o u s l a y e r s were not simple a r i t h m e t i c sums but were s e p a r a t e l y estimated v a l u e s which took i n t o account o v e r l a p of the v a r i o u s s p e c i e s and l a y e r components. Ground cover was estimated (percent) f o r humus, decayed wood i n c l u d i n g l o g s and s t i c k s , rocks and stones, mineral s o i l , and open water. 105 In each ecosystem map u n i t , 10 pri s m p l o t s were sampled u s u a l l y at a r a t e of 2 per v e g e t a t i o n p l o t where 5 p l o t s were sampled and 3 or 4 per v e g e t a t i o n p l o t where 3 v e g e t a t i o n p l o t s were sampled. Wherever p o s s i b l e , the common names of p l a n t s are used i n the t e x t and t a b l e s . The s c i e n t i f i c names and a u t h o r i t i e s are given i n Appendix 3. I observed that winter wren d i s t r i b u t i o n appeared to be c o r r e l a t e d w i t h the f o r e s t f l o o r v e g e t a t i o n i n which the b i r d s l i v e d . Four f o r e s t f l o o r h a b i t a t s f o r winter wrens were d i s t i n g u i s h e d : m o s s / l i t t e r , s a l a l , sword f e r n , and salmonberry. D i s t i n g u i s h i n g c r i t e r i a were the s p e c i e s composition and s t r u c t u r e of the understory and bryophyte-l i c h e n v e g e t a t i o n l a y e r s , and the ground cover. Areas with 20% or g r e a t e r cover of s a l a l , western sword f e r n , or salmonberry were assigned t o the s a l a l , sword f e r n , and salmonberry f o r e s t f l o o r h a b i t a t s r e s p e c t i v e l y . The few areas of f o r e s t f l o o r i n which the cover of two of s a l a l , western sword f e r n , or salmonberry exceeded 20% were ass i g n e d to f o r e s t f l o o r h a b i t a t s a c c o r d i n g to the g r e a t e s t cover v a l u e . Areas without s i g n i f i c a n t shrub or herb coverages were a s s i g n e d to the m o s s / l i t t e r f o r e s t f l o o r h a b i t a t . 106 The study area and p l o t s were mapped to f o r e s t f l o o r h a b i t a t s i r r e s p e c t i v e of K l i n k a ' s ecosystems. As f o r the mapping of ecosystems, the mapping of f o r e s t f l o o r h a b i t a t s was s t r a i g h t f o r w a r d as most t r a n s i t i o n s between f o r e s t f l o o r map u n i t s o c c u r r e d over 5 m or l e s s . The K l i n k a ecosystems were then grouped to the four f o r e s t f l o o r h a b i t a t s , a c c o r d i n g to the dominant f o r e s t f l o o r h a b i t a t of each ecosystem. The areas (m 2) of the f o r e s t f l o o r h a b i t a t s were c a l c u l a t e d from the areas of the component ecosystems. The mapping of the f o r e s t f l o o r h a b i t a t s i r r e s p e c t i v e of K l i n k a ' s ecosystems would have given a more ac c u r a t e d i s t r i b u t i o n and area (m 2) of the f o r e s t f l o o r h a b i t a t s than i s p r o v i d e d by the ecosystems grouped by f o r e s t f l o o r h a b i t a t s . However, the same c o u l d a l s o be s a i d of independent mappings of the d i s t r i b u t i o n s of many other a t t r i b u t e s of ecosystems (e.g., p l a n t s p e c i e s , parent m a t e r i a l s , s o i l s , humus forms, n u t r i e n t s and m o i s t u r e ) . I compared the mapping of ecosystems grouped by f o r e s t f l o o r h a b i t a t s with winter wren d i s t r i b u t i o n . G e n e r a l l y , the advantage of u s i n g the mapping of ecosystems and a p p r o p r i a t e l y grouped ecosystems i n t h i s way, as opposed to independent mapping of i n d i v i d u a l ecosystem a t t r i b u t e s , i s that although s l i g h t l y l e s s a c c u r a t e , the s i n g l e 107 mapping can be used f o r most a t t r i b u t e s . F u r t h e r , the g e n e r a l i z a t i o n s inherent i n the mapping of ecosystems and in grouped ecosystems can be used f o r f u r t h e r i n t e r p r e t i v e s t u d i e s and i n land management. Each v e g e t a t i o n p l o t was a s s i g n e d i n the f i e l d t o one of the four f o r e s t f l o o r h a b i t a t s . D e s c r i p t i o n s of the f o r e s t f l o o r h a b i t a t s were o b t a i n e d by grouping the v e g e t a t i o n p l o t s a s s i g n e d to the v a r i o u s f o r e s t f l o o r h a b i t a t s . I determined the s p a t i a l d i s t r i b u t i o n s (m 2) of male t e r r i t o r i e s and female home ranges i n terms of t h e i r aggregated areas w i t h i n i n d i v i d u a l ecosystems and ecosystems grouped to f o r e s t f l o o r h a b i t a t s . These d i s t r i b u t i o n s of t e r r i t o r i e s were c a l c u l a t e d f o r a l l census p e r i o d s on the study a r e a , study p l o t s , and removal p l o t s , while the d i s t r i b u t i o n of home ranges was c a l c u l a t e d only f o r the 1980 breeding p e r i o d on the study p l o t s . D i s t r i b u t i o n s were assessed by o v e r l a p p i n g F i g u r e s 4 to 17 at the working s c a l e of 1:787 on the ecosystem maps of the study area and removal p l o t s ( F i g u r e s 18 and 44). The areas (m 2) of each t e r r i t o r y and home range w i t h i n each ecosystem were c a l c u l a t e d by cou n t i n g map squares. The r e s p e c t i v e areas were summed to gi v e the t o t a l areas 108 of t e r r i t o r i e s and home ranges w i t h i n the v a r i o u s ecosystems, and w i t h i n the ecosystems grouped t o f o r e s t f l o o r h a b i t a t s . T h i s approach i s j u s t i f i e d as the s t u d y a r e a and p l o t s were censused s y s t e m a t i c a l l y (p. 3 5 ) . I a l s o d e t e r m i n e d the numbers of w i n t e r wren o b s e r v a t i o n s w i t h i n i n d i v i d u a l ecosystems and ecosystems grouped t o f o r e s t f l o o r h a b i t a t . The d i s t r i b u t i o n s of o b s e r v a t i o n s were a s s e s s e d f o r a l l census p e r i o d s on the st u d y a r e a , s t u d y p l o t s , and removal p l o t s by o v e r l a y i n g the w i n t e r wren o b s e r v a t i o n s on the ecosystem maps of the st u d y a r e a and removal p l o t s a t the w o r k i n g s c a l e of 1:787 and by c o u n t i n g the o b s e r v a t i o n s on the i n d i v i d u a l ecosystems. The number of w i n t e r wren o b s e r v a t i o n s on t h e a p p r o p r i a t e ecosystems were summed t o g i v e t h e numbers of w i n t e r wren o b s e r v a t i o n s on ecosystems grouped t o f o r e s t f l o o r h a b i t a t s . W i n t e r wren o b s e r v a t i o n s f e l l i n t o t h r e e c a t e g o r i e s : t e r r i t o r i a l m ales, o t h e r b i r d s , and t o t a l wren o b s e r v a t i o n s . T e r r i t o r i a l males i n c l u d e d a l l o b s e r v a t i o n s of s i n g i n g or t e r r i t o r i a l l y c h i p p i n g males, i n c l u d i n g the few s i n g i n g y e a r l i n g s o b s e r v e d , and a l l o t h e r o b s e r v a t i o n s of known c o l o u r - b a n d e d t e r r i t o r i a l m a l es. Other b i r d s i n c l u d e d a l l f e m a l e s and a l l s i l e n t or n o n t e r r i t o r i a l l y c h i p p i n g b i r d s not i n c l u d e d i n the p r e v i o u s c a t e g o r y . 109 Total wren observations was the sum of the previous categories. Description and mapping of the ecosystems and ecosystems grouped by forest floor habitats Study Area and Study Plots The study area, including the study plots was composed of 13 ecosystems. By considering only the vegetation component (plant community) and ignoring similar or c l o s e l y related s o i l components these were reduced to 10. My remapping included a l l 13 ecosystems (Figure 18) as map units but henceforward I consider only ecosystems grouped to plant communities. In addition, I excluded the map units R (non-forested ecosystems on rocks) and the areas within the clearcut (Figure 18) from further consideration as these areas were rarely u t i l i z e d by winter wrens. An abridged summary of the vegetation data for the ecosystems of the study area i s shown in Table 9. Only those plant species which occurred within ecosystems at 10% or more average cover are included, with the exception of the following diagnostic species from the Klinka synecological c l a s s i f i c a t i o n names: d e v i l ' s club, d u l l Oregon-grape, stink-currant, American skunk-cabbage, 110 FIGURE 18 REMAPPED SYNECOLOGICAL MAP OF THE STUDY AREA INCLUDING THE STUDY PLOTS A, B, AND C (See Table 9 and Appendix 2 for an explanation of the Klinka map units; forest f l o o r habitat map units are as follows: M/L - moss/litter; SAL - s a l a l ; SWD - sword fern; SAM - salmonberry) TABLE 9 ABRIDGliD VEGETATION PLOT DATA OF THE KLINKA SYNECOLOGICAL MAP UNITS (means) C l a s s i f i c a t i o n Remapped K l i n k a Map Unit Number 11.30 131.30/131.4 132.9 211.23 211.4 212.89 31.12 Remapped K l i n k a Map Unit Name (LICHEN)-GAULTI1ERIA-DF Gaultheria-WH-DF Mahonia-Gaultheria-WH-DF" Moss-WH Moss-WH Mahonia-Moss-WRC-WH MOSS-(POLYSTICHUM)-WH - WRC Forest Floor Habitat Map Unit S a l a l S a l a l ; Moss/1itter M o s s / l i t t e r ; S a l a l M o s s / l i t t e r Moss Moss Sword fern Location Study Area Study Area Study Area Removal P l o t E Study Area Removal Pl o t E Removal Pl o t E Number o f Sample P l o t s 3 S 3 3 5 3 3 Physiography Slope (%) 41 19 30 8 22 44 17 Vegetation Cover (1) A ( A i ; A 2; A 3; T ) a Coast Douglas f i r 47;0;0;47 a 38;0;0;38 70;0;0;70 6;0;0;6 47;0;0;47 23;0;0;23 12;0;0;12 Red alder 8;0;0;8 Western hemlock 6;2;2;9 24;3;4;31 2;3;7;11 33;10;0;43 14;7;5;26 46;17; 1;64 13;9;4;27 r s t o r y l a y e r (Ai - dominant and co-dominant tr e e s ; A2 - intermediate t r e e s ; A3 - suppressed trees over 10m; T - t o t a l f o r A l a y e r ) . Table 9 (continued) Map Unit Number 11.30 131.30/131.4 132.9 211.23 211.4 212.89 31.12 Western red cedar 15;3;4;22 9;5;11;25 2;9;4;15 30;23;2;S2 5;4;8;17 5;8;1;14 22;13;2;33 Totals 66;4;6;72 70;8;15;8S 73;12;10;85 68;36;2;88 64;11;12;79 75;27;1;88 S7;24;6;73 B(B 1;B 2;T) b Devil's-club D u l l Oregon-grape 0;3;3 b 0; 1; 1 0; 1; 1 0; 1; 1 0; 1; 1 Red alder 1;0; 1 Red huckleberry 0;6;6 0;3;3 0;3;3 0;1;1 0;3;3 0; 1; 1 0;2;2 S a l a l 0;40;40 0;35;35 0;16;16 0;2;2 0;4;4 0;3;3 0;1;1 Salmonberry 0; 1; 1 Sti n k currant Vine maple 1;0; 1 1;0; 1 8;0;8 1;0;1 27;0;27 Western hemlock 1;2;3 2; 1;2 5;0;5 3;2;5 7;2;9 2;1;3 4;1;5 - t o S i e f t 0 1 B } a y e r 1 ( B l " S m a 1 1 t r e e s ^ s h r u b s b e t w e e n 2 and 10m in height; B 2 - shrubs and woody plants between 0.15 and 2.0 i n height; Table 9 (continued) Map Unit Number 11.30 131.30/131.4 132.9 211.23 211.4 212.89 31.12 Western red cedar 12;1;13 6; 1;7 2; 1;2 l ; l ; 1 l ; l ; l 1; 1;2 1;0;1 Totals 13;48;55 8;37;42 6;20;27 12;5;15 9;8;15 5;5;8 31;4;33 C American skunk-cabbage L i c o r i c e f e r n 1 T r i f o l i a t e - l e a v e d foam flower Western sword fern 1 1 2 6 1 1 19 Totals 1 1 5 6 2 1 21 D Hylocomium splendens 25 20 9 5 9 1 19 Isothecium stoloniferum 3 1 1 1 2 Leucolepis m e n z i e s i i 1 Table 9 (continued) Map Unit Number 11.30 131.30/131.4 132.9 211.23 211.4 212.89 31.12 Plagiothecium undulatum 6 9 7 10 1 2 . Pleurozium schreberi 1 Rhytidiadelphus loreus 6 1 1 5 2 S t o k e s i e l l a oregana 27 35 25 5 23 9 16 Totals 68 68 43 20 46 12 44 Ground Cover ($) Humus 32 30 55 77 48 83 48 Decayed wood 5 7 11 6 13 5 10 Prism P l o t Data (means of a l l trees) Tree density (stems/ha) Basal area (mz/ha) Height (m)-(V) = Veteran Diameter breast height (cm) Age (yrs.)-(V) = Veteran 1098 46 31 29 100 1015 52 34 32 97 627 64 49 47 106 1096 57 29 31 51 432 38 43 43 102 1199 44 31 27 50 806 36 31 31 47 Table 9 (continued) C l a s s i f i c a t i o n Remapped K l i n k a Map Unit Number 31.412 62.90 71.4/71.89 71.12/71.4/ 71.89 72.4/72.47 812.89 911.0 Remapped K l i n k a Map Unit Name MOSS-(POLYSTICHUM)-POLYPODIUM-GAULTHHRIA-TIARELLA-POLYSTICHUM-TIARELLA-POLYSTICHUM-RUBUS-POLYSTIOIUM-Ribes -Oplopanax-Vaccinium-Lysichitum-WRC WH-WRC DF-WRC WRC WRC WRC WRC Forest Floor Habitat Map Unit Sword fern S a l a l Sword f e r n ; Salmonberry Sword f e r n Sword f e m ; Salmonberry Salmonberry Salmonberry Location Study Area Study Area Study Area Removal P l o t D Study Area Study Area Study Area Number of Sample P l o t s 5 3 5 5 3 3 3 Physiography Slope (%) 22 85 15 29 9 64 8 Vegetation Cover (4) A ( A 1 ; V V T ) Coast Douglas f i r 31;0;0;31 35;0;0;35 24;0;0;24 S9;0;0;59 1;0;0;1 2;0;0;2 Red alder 0;3;2;5 5;10;4;19 0;0;2;2 Western hemlock 28;6;4;37 3;0;0;3 26;4;4;34 2;0;2;4 38;6;9;53 10;3;2;14 7;0;3;10 Table 9 (continued) Map Unit Number 31.412 62.90 71.4/71.89 71.12/71.4/ 71.89 72.4/72.47 812.89 911.0 Western red cedar 7;2;2;11 10;4;1;15 13;1;4;18 4;9;7;19 18;1;0;20 13;2;0;14 12;1;0;13 Totals 65;9;7;73 46;4;1;50 64;8;8;71 6S;12;9;71 58;11;10;70 30;14;8;43 18;1;3;22 B(B i :B 2;T) Dev i l ' s - c l u b 0; 1; 1 0;1;1 0;1;1 0;2;2 0;8;8 0;9;9 D u l l Oregon-grape 0; 1; 1 Red a l d e r 3;0;3 8; 1;8 10;0;10 Red huckleberry 0;10;10 0;8;8 0;3;3 0;2;2 0;3;3 0;S;5 0;5;5 S a l a l 0;1;1 0;27;27 Salmonberry 0; 1; 1 0; 1; 1 0;6;6 0;1;1 0;22;22 0;48;48 0;57;57 St i n k currant 0;8;8 0; 1; 1 Vine maple 24;0;24 1;0;1 41;0;41 28; 1;28 S0;26;65 14;0;14 48;0;48 Western hemlock 6;S;10 2;1;3 5;1;6 2; 1;3 6;1;7 3;2;S 5; 1;6 Table 9 (continued) Map Unit Number 31.412 62.90 71.4/71.89 71.12/71.4/ 71.89 72.4/72.47 812.89 911.0 Western red cedar 1; 1; 1 4;4;8 1;0; 1 1;0; 1 l ; i ; i i ; i ; 2 2;0;2 Totals 31;16;41 7;39;43 49;11;54 30;4;32 53;26;65 25;68;74 67;62;92 C American skunk-cabbage 1 1 1 6 L i c o r i c e f e r n 1 1 1 T r i f o l i a t e - l e a v e d foamflower 1 3 2 2 3 4 Western sword f e r n 33 7 53 52 25 S 13 Totals 41 10 62 60 35 22 31 D Hylocomium splendens 24 30 12 2 20 9 10 Isothecium stoloniferum 1 2 5 2 9 2 10 Leucolepis m e n z i e s i i 1 1 1 10 7 Table 9 (continued) Map Unit Number 31.412 62.90 71.4/71.89 71.12/71.4/ 71.89 72.4/72.47 812.89 911.0 Plagiothecium undulatum 14 4 7 4 8 3 8 Pleurozium schreberi 1 • Rhytidiadelphus loreus 11 8 4 2 5 16 3 S t o k e s i e l l a oregana 16 14 10 7 12 4 5 Totals 69 72 43 20 60 60 53 Ground Cover (%) Humus 30 27 52 69 37 17 33 Decayed wood 6 6 10 8 9 4 17 Prism P l o t Data (means of a l l trees) Tree density (stems/ha) Basal area (m^/ha) Height (m)-(V) = Veteran Diameter breast height (cm) Age (yrs.)-(V) = Veteran 271 565 190 232 52 36 46 44 56;61(V) 30 51;49(V) 50 69 37 80 69 107;250+(V) 95 99;170(V) 81 368 86 87 41 17 17 39 37;49(V) 17;49(V) 57 65 82 99 83;221(V) 41;202(V) 120 l i c o r i c e f e r n , and t r i f o l i a t e - l e a v e d foamflower. A schematic diagram of the r e l a t i v e l o c a t i o n s of s e l e c t e d ecosystems on the study area i s shown i n F i g u r e 19. The areas and percentages occupied by the ecosystems of the study area and of the study p l o t s are shown i n Table 10. The areas are expressed i n m2 as they are l a t e r used f o r comparisons with observed winter wren d i s t r i b u t i o n s . The v e g e t a t i o n of the study area and study p l o t s dates from a severe w i l d f i r e i n 1859 which, with the exc e p t i o n of a few veteran coast Douglas f i r , d e s t r o y e d the o r i g i n a l f o r e s t cover (Sanders 1981). The f o r e s t v e g e t a t i o n of the ecosystems of the study area and study p l o t s i s shown i n v e r t i c a l , photographic mosaics and i n c l o s e ups of the f o r e s t f l o o r as f o l l o w s : (LICHEN) - GAULTHERIA - DF, 11.30 - F i g u r e s 20 and 23; G a u l t h e r i a - WH - DF, 131.30/131.4 -F i g u r e s 21 and 24; Mahonia - G a u l t h e r i a - WH - DF, 132.9 -F i g u r e s 22 and 25; Moss - WH, 211.4 - F i g u r e s 26 and 29; MOSS - (POLYSTICHUM) - WH - WRC, 31.412 - F i g u r e s 27 and 30; POLYPODIUM - GAULTHERIA - DF - WRC, 62.90 - F i g u r e s 28 and 31; TIARELLA - POLYSTICHUM - WRC, 71.4/71.89 - F i g u r e s 32 and 35; RUBUS - POLYSTICHUM - WRC, 72.4/72.47 - F i g u r e s 33 and 36; Ribes - Oplopanax - WRC, 812.89 - F i g u r e s 34 and 37; and Vaccinium - L y s i c h i t u m - WRC - 911.0 - F i g u r e s FIGURE 19 SCHEMATIC DIAGRAM OF THE RELATIVE LOCATIONS OF SELECTED ECOSYSTEMS ON THE SLOPES OF THE STUDY AREA (not drawn to scale) Non-forested Ecosystems on Rocks, R (LICHEN) - GAULTHERIA-DF, 11-30 Goultherio-WH-DF, 131 3 0 Gaultherio-WH-DF. 131-4 Moss-WH, 211-4 POLYPODIUM-GAULTHERIA-DF-WRC, 6 2 9 0 Mohonio-Goultherio-WH-DF. 132-9 MOSS-(POLYSTICHUM)-WH-WRC, 31-412 T I A R E L L A - POLYSTICHUM-WRC. 71-4 RUBUS-POLYSTICHUM-WRC T 72-4 Voccinium-Lysichitum-WRC. 9 1 1 0 TIARELLA-POLYSTICHUM-WRC. 71-89 Ribes-Oploponox-WRC. 812 8 9 TABLE 10 AREAS (m2) AND PERCENTAGES OF THE ECOSYSTEMS ON THE STUDY AREA AND STUDY PLOTS Area i n m (%) Ecosystem Study Area Study P l o t s (LICHEN) - GAULTHERIA - DF, 11.30 15,335 (3.97) 10,832 (8.88) G a u l t h e r i a - WH - DF. 131.30/131.4 44,769 (11.58) 16,130 (13.23) Mahonia - G a u l t h e r i a - WH - DF. 132.9 24,159 (6.25) 6,779 (5.56) Moss - WH, 211.4 19,009 (4.92) 1,589 (1.30) MOSS - (POLYSTICHUM) - WH - WRC. 31.412 62,000 (16.04) 14,361 (11.77) POLYPODIUM - GAULTHERIA - DF - WRC. 62.90 18,805 (4.86) 3,689 (3.03) TIARELLA - POLYSTICHUM - WRC, 71.4/71.89 131,527 (34.03) 56,088 (46.00) RUBUS - POLYSTICHUM - WRC. 72.4/72.47 43,255 (11.19) 7,901 (6.48) Ribes - Oplopanax - WRC, 812.89 24,470 (6.33) 1,653 (1.36) Vaccinium - Lysichitum - WRC. 911.0 3,191 (0.83) 2,919 (2.39) T o t a l Area 386,520(100.00) 121,941(100.00) 124 FIGURE 20 ( l e f t ) (LICHEN) - GAULTHERIA - DF, 11.30, (STUDY AREA) : VERTICAL MOSAIC ( s t a f f shows 0.25 m i n t e r v a l s ) F o r e s t f l o o r h a b i t a t : s a l a l FIGURE 21 ( c e n t r e ) GAULTHERIA - WH - DF, 131.30/131.4, (STUDY AREA): VERTICAL MOSAIC ( s t a f f shows 0.25 m i n t e r v a l s ) F o r e s t f l o o r h a b i t a t : s a l a l FIGURE 22 ( r i g h t ) Mahonia - G a u l t h e r i a - WH - DF, 132.9, (STUDY AREA): VERTICAL MOSAIC ( s t a f f shows 0.25 m i n t e r v a l s ) F o r e s t f l o o r h a b i t a t : m o s s / l i t t e r 12 5 126 FIGURE 23 ( t o p ) (LICHEN) - GAULTHERIA - DF, 11.30, (STUDY AREA) : FOREST FLOOR ( s t a f f shows 0.25 m i n t e r v a l s ) F o r e s t f l o o r h a b i t a t : s a l a l FIGURE 24 ( c e n t r e ) GAULTHERIA - WH - DF, 131.30/131.4, (STUDY AREA): FOREST FLOOR ( s t a f f shows 0.25 m i n t e r v a l s ) F o r e s t f l o o r h a b i t a t : s a l a l FIGURE 25 (bottom) Mahonia - G a u l t h e r i a - WH - DF, 132.9, (STUDY AREA): FOREST FLOOR ( s t a f f shows 0.25 m i n t e r v a l s ) F o r e s t f l o o r h a b i t a t : m o s s / l i t t e r 127 128 FIGURE 26 ( l e f t ) MOSS - WH, 211.4, (STUDY AREA) : VERTICAL MOSAIC ( s t a f f shows 0.25 m i n t e r v a l s ) F o r e s t f l o o r h a b i t a t : m o s s / l i t t e r FIGURE 27 ( c e n t r e ) MOSS - (POLYSTICHUM) - WH - WRC, 31.412, (STUDY AREA) : VERTICAL MOSAIC ( s t a f f shows 0.25 m i n t e r v a l s ) F o r e s t f l o o r h a b i t a t : sword f e r n FIGURE 28 ( r i g h t ) POLYPODIUM - GAULTHERIA - DF - WRC, 62.90, (STUDY AREA) : VERTICAL MOSAIC ( s t a f f shows 0.25 m i n t e r v a l s ) F o r e s t f l o o r h a b i t a t : m o s s / l i t t e r 1 3 0 FIGURE 29 ( t o p ) MOSS - WH, 211.4, (STUDY AREA) : FOREST FLOOR ( s t a f f shows 0.25 m i n t e r v a l s ) F o r e s t f l o o r h a b i t a t : m o s s / l i t t e r FIGURE 30 ( c e n t r e ) MOSS - (POLYSTICHUM) - WH - WRC, 31.412, (STUDY AREA) : FOREST FLOOR ( s t a f f shows 0.25 m i n t e r v a l s ) F o r e s t f l o o r h a b i t a t : sword f e r n FIGURE 31 (bottom) POLYPODIUM - GAULTHERIA - DF - WRC, 62.90, (STUDY AREA) : FOREST FLOOR ( s t a f f shows 0.25 m i n t e r v a l s ) F o r e s t f l o o r h a b i t a t : m o s s / l i t t e r 131 132 FIGURE 32 ( l e f t ) TIARELLA - POLYSTICHUM - WRC, 71.4/71.89, (STUDY AREA) : VERTICAL MOSAIC ( s t a f f shows 0.25 m i n t e r v a l s ) F o r e s t f l o o r h a b i t a t : sword f e r n FIGURE 33 ( c e n t r e ) RUBUS - POLYSTICHUM - WRC, 72.4/72.47, (STUDY AREA) : VERTICAL MOSAIC ( s t a f f shows 0.25 m i n t e r v a l s ) F o r e s t f l o o r h a b i t a t : sword f e r n FIGURE 34 ( r i g h t ) H i b e s - Oplopanax - WRC, 812.89, (STUDY AREA) : VERTICAL MOSAIC ( s t a f f shows 0.25 m i n t e r v a l s ) F o r e s t f l o o r h a b i t a t : s a l m o n b e r r y 133 134 FIGURE 35 ( t o p ) TIARELLA - POLYSTICHUM - WRC, 71.4/71.89, (STUDY AREA) : FOREST FLOOR ( s t a f f shows 0.25 m i n t e r v a l s ) F o r e s t f l o o r h a b i t a t : sword f e r n FIGURE 36 ( c e n t r e ) RUBUS - POLYSTICHUM - WRC, 72.4/72.47, (STUDY AREA) : FOREST FLOOR ( s t a f f shows 0.25 m i n t e r v a l s ) F o r e s t f l o o r h a b i t a t : sword f e r n FIGURE 37 (bottom) R i b e s - Oplopanax - WRC, 812.89, (STUDY AREA) : FOREST FLOOR ( s t a f f shows 0.25 m i n t e r v a l s ) F o r e s t f l o o r h a b i t a t : sa l m o n b e r r y 136 38 and 41. The mapped d i s t r i b u t i o n s of the f o r e s t f l o o r h a b i t a t s on the study area and study p l o t s are shown i n F i g u r e 18. The percent occurrences of the f o r e s t f l o o r h a b i t a t s on the v a r i o u s ecosystems, and the r e s u l t i n g assignments of i n d i v i d u a l ecosystems to f o r e s t f l o o r h a b i t a t s based on the dominant f o r e s t f l o o r h a b i t a t are shown i n Table 11. The areas (m 2) of the f o r e s t f l o o r h a b i t a t s of the study area and the study p l o t s as c a l c u l a t e d from the ecosystems grouped by f o r e s t f l o o r h a b i t a t are shown i n Table 12. An a b r i d g e d summary of the v e g e t a t i o n p l o t data grouped to as s i g n e d f o r e s t f l o o r h a b i t a t s i s shown i n Table 13. The f o r e s t f l o o r h a b i t a t s of the study area and study p l o t s are i l l u s t r a t e d among the v a r i o u s ecosystems as f o l l o w s : m o s s / l i t t e r - F i g u r e s 22, 25, 26, 28, 29, and 31; s a l a l - F i g u r e s 20, 21, 23, and 24; sword f e r n -F i g u r e s 27, 30, 32, 35, and 38; and salmonberry - F i g u r e s 34, 37, 38, and 41. Removal P l o t s Removal p l o t D was comprised of th r e e ecosystems which, when grouped a t the parent m a t e r i a l s l e v e l , were reduced to one. There were three ecosystems on removal p l o t E. I remapped the removal p l o t s as f o r the study 137 FIGURE 38 ( l e f t ) V a c c i n i u m - L y s i c h i t u m - WRC, 911.0, (STUDY AREA) : VERTICAL MOSAIC ( s t a f f shows 0.25 m i n t e r v a l s ) F o r e s t f l o o r h a b i t a t : sa l m o n b e r r y i n f o r e g r o u n d and sword f e r n i n t h e background FIGURE 39 ( c e n t r e ) TIARELLA - POLYSTICHUM - WRC, 71.12/71.4/71.89, (REMOVAL PLOT D) : VERTICAL MOSAIC ( s t a f f shows 0.25 m i n t e r v a l s ) F o r e s t f l o o r h a b i t a t : sword f e r n FIGURE 40 ( r i g h t ) Moss - WH, 211.23, (REMOVAL PLOT E) : VERTICAL MOSAIC ( s t a f f shows 0.25 m i n t e r v a l s ) F o r e s t f l o o r h a b i t a t : m o s s / l i t t e r 158 139 FIGURE 41 ( t o p ) V a c c i n i u m - L y s i c h i t u m - WRC, 911.0, (STUDY AREA) : FOREST FLOOR ( s t a f f shows 0.25 m i n t e r v a l s ) F o r e s t f l o o r h a b i t a t : salmonberry FIGURE 42 ( c e n t r e ) TIARELLA - POLYSTICHUM - WRC, 71.12/71.4/71.89, (REMOVAL PLOT D) : FOREST FLOOR ( s t a f f shows 0.25 m i n t e r v a l s ) F o r e s t f l o o r h a b i t a t : sword f e r n FIGURE 43 (bottom) Moss - WH, 211.23, (REMOVAL PLOT E) : FOREST FLOOR ( s t a f f shows 0.25 m i n t e r v a l s ) F o r e s t f l o o r h a b i t a t : m o s s / l i t t e r TABLE 11 PERCENT OCCURRENCES OF THE FOREST FLOOR HABITATS1 ON THE ECOSYSTEMS ON THE STUDY AREA AND THE RESULTING ASSIGNMENTS OF THE ECOSYSTEMS BY FOREST FLOOR HABITATS Ecosystem Occurrence o f Forest F l o o r H a b i t a t i n Percent Assignment (each ecosystem 100 percent) Moss/ Sword Salmon-l i t t e r S a l a l f e r n b e r r y (LICHEN) - GAULTHERIA - DF. 11.30 100 S a l a l G a u l t h e r i a - WH - DF. 131.30/131.4 47 53 S a l a l Mahonia - G a u l t h e r i a - WH - DF. 132.9 70 21 9 M o s s / l i t t e r Moss - WH, 211.4 73 27 M o s s / l i t t e r MOSS - (POLYSTICHUM) - WRC - WH. 31.412 2 98 Sword f e r n POLYPODIUM - GAULTHERIA - DF - WRC. 62.90 76 24 S a l a l TIARELLA - POLYSTICHUM - WRC. 71.4/71.89 100 1 Sword f e r n RUBUS - POLYSTICHUM - WRC. 72.4/72.47 13 80 7 Sword f e r n Ribes - Oploponax - WRC. 812.89 100 Salmonberry Vaccinium - Lysi c h i t u m - WRC. 911.0 100 Salmonberry Forest f l o o r h a b i t a t s as mapped i r r e s p e c t i v e o f K l i n k a ' s s y n e c o l o g i c a l u n i t s TABLE 12 AREAS(m2) AND PERCENTAGES OF THE FOREST FLOOR HABITATS ON THE STUDY AREA AND THE STUDY PLOTS AS CALCULATED FROM THE ECOSYSTEMS GROUPED AS SHOWN IN TABLE 11 Forest F l o o r Habitats Area i n m 2(%) Study Area Study P l o t s M o s s / l i t t e r 43,168 (11.17) 8,368 (6.86) S a l a l 78,909 (20.41) 30,651 (25.14) Sword f e r n 236,782 (61.26) 78,350 (64.25) Salmonberry 27,661 (7.16) 4,572 (3.75) T o t a l 386,520(100.00) 121,941(100.00) TABLE 13 ABRIDGED VEGETATION PLOT DATA FOR THE FOREST FLOOR HABITATS (means) C l a s s i f i c a t i o n Forest Floor Habitats M o s s / l i t t e r S a l a l Sword fern Salmonberry Sword fern M o s s / l i t t e r Sword f e r n Location Study Area Study Area Study Area Study Area Removal P l o t D Removal P l o t E Removal P l o t E Number o f Vegetation P l o t s 9 10 11 8 5 6 3 Physiography Slope (*) 21 46 18 8 29 26 17 Vegetation Cover (1) Coast Douglas f i r 50;0;0;50 a 42;0;0;42 2S;0;( );25 1;0;0;1 59;0;0;S9 14;0;0;14 12;0;0;12 Red alder 0; 1; l ; l 2;4;2;8 8;0;0;8 Western hemlock 12;5;5;22 11;2;2;16 29;5;J i;39 15;2;3;19 2;0;2;4 40;13; 1;53 13;9;4;27 Western red cedar 5;6;10;21 10;4;4;18 12;2;2 !;16 12;1;1;14. 4;9;7;19 17;16;1;33 22;13;2;33 Totals 65;11;14;82 63;6;6;70 66;9;7 ';74 30;7;7;39 65;12;9;71 72;31;1;88 57;24;6;73 B ( B i ; B 2 ; T ) b A - overstory layer (Aj - dominant and co-dominant trees; A 2 - intermediate trees; AT - suppressed trees over 10m; T - t o t a l f o r A l a y e r ) . j u t B - middlestory layer (Bj - small trees and shrubs between 2 and 10m i n height; B? - shrubs and woody plants between 0.1S and 2.0m i n height; T - t o t a l f o r B l a y e r ) . Table 13 (continued) Forest Floor Habitats M o s s / l i t t e r S a l a l Sword fern Salmonberry Sword fern M o s s / l i t t e r Sword f e r n Devil's-club 0 ; l ; l b 0;7;7 0;1;1 D u l l Oregon-grape 0; 1; 1 0; 1; 1 0; 1; 1 0; 1; 1 Red a l d e r 1;0; 1 1;0; 1 8; 1;8 Red huckleberry 0;2;2 0;6;6 0;6;6 0;4;4 0;2;2 0;1;1 0;2;2 S a l a l 0;6;6 0;39;39 0; 1; 1 0;2;2 0;1;1 Salmonberry 0; 1; 1 0;4;4 0;46;46 0;1;1 Stink currant 0;3;3 Vine maple 1;0; 1 1;0; 1 35;0;35 34;0;34 28; 1;28 4;0;4 27;0;27 Western hemlock S;l;6 2;1;3 6;3;8 4; 1;5 2; 1;3 3;2;4 4;1;5 Western red cedar 2; 1;2 7;2;9 l ; l ; i l ; i ; i i 1;0; 1 l ; 1; 1 1;0;1 Totals 7;9;15 9;47;52 37;13;46 48;57;78 30;4;32 8;5;12 31;4;33 C American skunk-cabbage Table 13 (continued) Forest Floor Habitats Moss/litter Salal Sword fern Salmonberry Sword fern Moss/litter Sword fern Licorice fern 1 1 1 Trifoliate-leaved foamflower 2 3 2 Western sword fern 1 2 43 10 52 3 19 Totals 3 4 52 26 60 4 21 D Hylocomium splendens 9 26 19 11 2 3 19 Isothecium stoloniferum 1 1 3 7 2 1 2 Leucolepis menziesii 1 7 1 1 Plagiothecium undulatum 9 7 10 7 4 4 2 Rhytidiadelphus loreus 3 6 7 8 2 1 2 Stokesie11a oregana 28 23 13 7 7 7 16 Totals 48 69 57 56 20 16 44 Table 13 (continued) Forest Floor Habitats M o s s / l i t t e r S a l a l Sword f e r n Salmonberry Sword f e r n M o s s / l i t t e r Sword fern Ground Cover (%) Humus 47 20 40 29 69 80 48 Decayed wood 11 6 7 12 8 S 10 147 area to show both p l a n t communities and s o i l parent m a t e r i a l s ( F i g u r e 44) although h e n c e f o r t h the s o i l component w i l l be ignored and the ecosystems w i l l be grouped to p l a n t community. The abridged v e r s i o n of the v e g e t a t i o n data f o r these ecosystems i s shown as f o r the study area i n Table 9. F i g u r e 45 pr e s e n t s a schematic drawing of the r e l a t i v e l o c a t i o n s of the ecosystems on the sl o p e s of the removal p l o t s . The areas (m 2) and percentages of the removal p l o t s occupied by the ecosystems are shown (Table 14). The v e g e t a t i o n of removal p l o t D dates from the same 1859 f i r e as the v e g e t a t i o n of the study area and study p l o t s . The v e g e t a t i o n of removal p l o t E, however, i s much younger and dates from l o g g i n g and s l a s h f i r e s i n 1929 (Sanders 1981). The f o r e s t v e g e t a t i o n of the ecosystem TIARELLA - POLYSTICHUM - WRC, 71.12/71.4/71.89 of removal p l o t D i s shown i n F i g u r e s 39 and 42. G e n e r a l l y the v e g e t a t i o n of t h i s ecosystem was very s i m i l a r to that of the c o r r e s p o n d i n g ecosystem, TIARELLA - POLYSTICHUM - WRC, 71.4/71.89, of the study area ( F i g u r e s 32 and 35; F i g u r e s 39 and 42; Table 9 ) . Given that s i m i l a r i t y , the d i s t r i b u t i o n s of winter wrens on removal, p l o t D were compared as a g e n e r a l check of the f i n d i n g s on the corr e s p o n d i n g ecosystem and ecosystems grouped by f o r e s t 148 FIGURE 44 REMAPPED SYNECOLOGICAL MAP OF THE REMOVAL PLOTS D AND E (see Table 9 and Appendix 2 f o r an e x p l a n a t i o n of the K l i n k a map u n i t s ; f o r e s t f l o o r h a b i t a t map u n i t s are as f o l l o w s : M/L - m o s s / l i t t e r ; SWD - sword f e r n ) 149 LEGEND Ecosystem Boundary Forest Floor Habitat Boundary FIGURE 4 5 SCHEMATIC DIAGRAM OF THE RELATIVE LOCATIONS OF THE ECOSYSTEMS ON THE SLOPES OF THE REMOVAL PLOTS D AND E (not drawn t o s c a l e ) Note: t h e removal p l o t s were l o c a t e d on d i f f e r e n t h i l l s i d e s w i t h s i m i l a r e x p o s u r e s 1 S T TABLE 14 NTAGES OF THE REMOVAL PLOTS D AND E AREAS(m2) AND PERCE THE ECOSYSTEMS ON Ecosystem Area i n m2 (%)  Removal P l o t D TIARELLA - POLYSTICHUM - WRC, 71.12/71.4/71.89 39,375(100.00) Removal P l o t E Moss - WH, 211.23 23,897 (60.69) Mahonia - Moss - WRC - WH, 212.89 8,922 (22.66) MOSS - (POLYSTICHUM) - WH - WRC, 31.12 6,556 (16.65) T o t a l 39,375(100.00) 153 f l o o r h a b i t a t s on the study area and study p l o t s . The f o r e s t v e g e t a t i o n of the ecosystems of removal p l o t E are shown as f o l l o w s : Moss - WH, 211.23 - F i g u r e s 40 and 43; Mahonia - Moss - WH - WRC, 212.89 - F i g u r e s 46 and 48; and MOSS - POLYSTICHUM - WH - WRC, 31.12 - F i g u r e s 47 and 49. G e n e r a l l y , the m i d d l e s t o r y and understory v e g e t a t i o n and ground cover of the ecosystems Moss - WH, 211.23 and Mahonia - Moss - WH - WRC, 212.89 of removal p l o t E resembled areas on the study area and study p l o t s i n the correspo n d i n g and/or c l o s e l y r e l a t e d ecosystems ( F i g u r e s 22, 25, 26, and 29; F i g u r e s 40, 43, 46, and 48; Table 9 ) . A l l were ecosystems which had heavy o v e r s t o r y cover and much reduced lower v e g e t a t i o n l a y e r s . However, due to the age d i f f e r e n c e i n the v e g e t a t i o n of the two areas (53 versus 123 y e a r s ) , t h e i r v e g e t a t i o n c o u l d not be s a i d t o be as s i m i l a r as was the case f o r the preceding c o r r e s p o n d i n g ecosystems of removal p l o t D and the study a r e a . The v e g e t a t i o n of the ecosystem MOSS - (POLYSTICHUM) -WH - WRC, 31.12 of removal p l o t E a l s o g e n e r a l l y resembled the c o r r e s p o n d i n g ecosystem of the study area and ecosystem of the study area and removal p l o t s ( F i g u r e 27, and 30; F i g u r e s 47, and 49). However, the v e g e t a t i o n 154 FIGURE 46 ( l e f t ) Mahonia - Moss - WRC - WH, 212.89, (REMOVAL PLOT E) : VERTICAL MOSAIC ( s t a f f shows 0.25 m i n t e r v a l s ) F o r e s t f l o o r h a b i t a t : m o s s / l i t t e r FIGURE 47 ( r i g h t ) MOSS - (POLYSTICHUM) - WH - WRC, 31.12, (REMOVAL PLOT E) : VERTICAL MOSAIC ( s t a f f shows 0.25 m i n t e r v a l s ) F o r e s t f l o o r h a b i t a t : sword f e r n 156 FIGURE 48 ( t o p ) Mahonia - Moss - WRC - WH, 212.89, (REMOVAL PLOT E) : FOREST FLOOR ( s t a f f shows 0.25 m i n t e r v a l s ) F o r e s t f l o o r h a b i t a t : m o s s / l i t t e r FIGURE 49 (bottom) MOSS - (POLYSTICHUM) - WH - WRC, 31.12, (REMOVAL PLOT E) : FOREST FLOOR ( s t a f f shows 0.25 m i n t e r v a l s ) F o r e s t f l o o r h a b i t a t : sword f e r n 1 5 ? 158 cover of the m i d d l e s t o r y and understory of the removal p l o t E ecosystem were g e n e r a l l y r e l a t i v e l y lower, due probably to a lack of m a t u r i t y of the younger v e g e t a t i o n of the removal p l o t (Table 9 ) . In s p i t e of these minor d i f f e r e n c e s , the d i s t r i b u t i o n s of winter wrens on the ecosystems of removal p l o t E were compared as a very g e n e r a l check with the f i n d i n g s on the corresponding ecosystems on the study area and study p l o t s . The mapped d i s t r i b u t i o n s of the f o r e s t f l o o r h a b i t a t s w i t h i n the v a r i o u s ecosystems and the r e s u l t i n g assignments of i n d i v i d u a l ecosystems to f o r e s t f l o o r h a b i t a t s based on the m a j o r i t y f o r e s t f l o o r h a b i t a t are shown i n Table 15. Removal p l o t D was comprised e n t i r e l y of sword f e r n f o r e s t f l o o r h a b i t a t . A f t e r grouping the i n d i v i d u a l ecosystems of removal p l o t E by f o r e s t f l o o r h a b i t a t s based on the dominant f o r e s t f l o o r h a b i t a t , removal p l o t E was c l a s s i f i e d as e n t i r e l y m o s s / l i t t e r f o r e s t f l o o r h a b i t a t . The abridged v e r s i o n of the v e g e t a t i o n data f o r these f o r e s t f l o o r h a b i t a t s i s shown as f o r the study area i n Table 13. As f o r the ecosystems, the d i s t r i b u t i o n s of winter wrens on the ecosystems grouped by f o r e s t f l o o r h a b i t a t s of the removal p l o t s were compared as a very g e n e r a l check with the f i n d i n g s on the TABLE 15 PERCENT OCCURRENCES OF THE FOREST FLOOR HABITATS ON THE ECOSYSTEMS OF THE REMOVAL PLOTS AND THE RESULTING ASSIGNMENTS OF THE ECOSYSTEMS TO FOREST FLOOR HABITATS Occurrence o f Forest F l o o r  H a b i t a t i n Percent Ecosystem (each ecosystem 100 percent) Assignment M o s s / l i t t e r Sword f e r n Removal P l o t D TIARELLA - POLYSTICHUM - WRC, 71.12/71.4/71.89 100 Sword f e r n Moss - WH, 211.23 Mahonia - Moss - WRC - WH, 212.89 MOSS - (POLYSTICHUM) - WH - WRC, 31.12 cn Removal P l o t E 100 M o s s / l i t t e r 100 M o s s / l i t t e r 61 39 M o s s / l i t t e r 160 c o r r e s p o n d i n g grouped ecosystems on the study area and study p l o t s . The f o r e s t f l o o r h a b i t a t s of the removal p l o t s are i l l u s t r a t e d among the v a r i o u s ecosystems as f o l l o w s : m o s s / l i t t e r (removal p l o t E) - F i g u r e s 40, 43, 46, and 48; and sword f e r n (removal p l o t s D and E) -F i g u r e s 39, 42, 47, and 49. D i s t r i b u t i o n s of male t e r r i t o r i e s and female home ranges among the ecosystems and ecosystems grouped by f o r e s t f l o o r h a b i t a t s Here I compare the d i s t r i b u t i o n s of male t e r r i t o r i e s and female home ranges with the d i s t r i b u t i o n s of K l i n k a ' s ecosystems and ecosystems grouped by f o r e s t f l o o r h a b i t a t s . Winter wrens are shown to u t i l i z e the ecosystems and grouped ecosystems d i f f e r e n t i a l l y , and to u t i l i z e c o n s i s t e n t l y some ecosystems and ecosystems grouped by f o r e s t f l o o r h a b i t a t s over o t h e r s . The observed d i s t r i b u t i o n s (m 2) of male t e r r i t o r i e s and female home ranges on the v a r i o u s ecosystems and ecosystems grouped by f o r e s t f l o o r h a b i t a t s were compared us i n g X 2 t e s t s with the co r r e s p o n d i n g expected val u e s as repr e s e n t e d by the areas (m 2) of the v a r i o u s ecosystems and ecosystems grouped by f o r e s t f l o o r h a b i t a t s (Tables 10, 12, and 14). Comparisons were made f o r the study 161 area, study p l o t s , and removal p l o t E f o r the v a r i o u s i n d i v i d u a l census p e r i o d s (Tables 16, to 18). Comparisons were not p o s s i b l e f o r ecosystems or grouped ecosystems on removal p l o t D which was composed of only one ecosystem, or f o r grouped ecosystems on removal p l o t E f o r which the ecosystems were grouped t o only one f o r e s t f l o o r h a b i t a t (Table 15). As a l l su r v e y i n g and f i e l d o b s e r v a t i o n s were made i n m and a l l c a l c u l a t e d areas were counts of m2, u n i t s of m2 are employed i n the s t a t i s t i c a l a n a l y s e s . The s t a t i s t i c a l comparisons of the expected and observed d i s t r i b u t i o n s of the areas of t e r r i t o r i e s and home ranges were a l l h i g h l y s i g n i f i c a n t (P < 0.001) f o r the study a r e a , study p l o t s , and removal p l o t E (Tables 16 to 18) i n d i c a t i n g a d i f f e r e n t i a l u t i l i z a t i o n of the v a r i o u s ecosystems and ecosystems grouped by f o r e s t f l o o r h a b i t a t s by winter wrens. Expected t o t a l areas (m 2) of male t e r r i t o r i e s and female home ranges w i t h i n the v a r i o u s ecosystems and ecosystems grouped by f o r e s t f l o o r h a b i t a t s were c a l c u l a t e d from t o t a l areas of the v a r i o u s ecosystems and grouped ecosystems based on p r o p o r t i o n a t e random d i s t r i b u t i o n . The observed areas of t e r r i t o r i e s and home ranges found d u r i n g the censuses of the study areas d i f f e r e d from the expected v a l u e s (Tables 19, and 20). TABLE 16 STATISTICAL COMPARISONS (X 2) OF THE EXPECTED AND OBSERVED DISTRIBUTIONS OF THE AREAS (m2) OF MALE TERRITORIES AMONG THE ECOSYSTEMS AND ECOSYSTEMS GROUPED BY FOREST FLOOR HABITATS DURING THE VARIOUS CENSUS PERIODS OF THE STUDY AREA 1978-81 Census P e r i o d Breeding /78 Breeding /79 Breeding /80 Breeding /81 (169,561) a (186,491) (182,089) (211,225) Ecosystems (9d£) 14,185 b 21,707 12,709 13,875 Forest F l o o r Habitats from . „ , Grouped Ecosystems (3df) 9 » 3 7 6 11,458 8,586 4,502 T o t a l areas occupied by t e r r i t o r i e s (m ) X 2 v a l u e ; P f o r a l l t e s t s < 0.001 TABLE 17 STATISTICAL COMPARISONS (X 2) OF THE EXPECTED AND OBSERVED DISTRIBUTIONS OF THE AREAS (m^) OF MALE TERRITORIES AND FEMALE HOME RANGES AMONG THE ECOSYSTEMS AND ECOSYSTEMS GROUPED BY FOREST FLOOR HABITATS DURING THE VARIOUS CENSUS PERIODS OF THE STUDY PLOTS, 1979-80 Census Period Units Male T e r r i t o r i e s Winter /79 (21,130) a Breeding /79 F a l l /79 Winter /79-80 Breeding /80 (87,453) (55,703) (48,880) (84,446) Female Home Ranges Breeding /80 (52,443) Ecosystems (9df) Forest Floor Habitats from Grouped Ecosystems (3df) 10,543 u 10,224 11,235 9,162 22,830 21,819 20,417 18,842 10,037 9,256 2,445 121 Tot a l areas occupied by t e r r i t o r i e s or home ranges (m ) X 2 value; P f o r a l l t e s t s < 0.001 TABLE 18 STATISTICAL COMPARISONS (X 2) OF THE EXPECTED AND OBSERVED DISTRIBUTIONS OF THE AREAS (m2) OF MALE TERRITORIES AMONG THE ECOSYSTEMS FOR THE FIRST CENSUSES ON REMOVAL PLOT E, 1979-80 Uni t s Census P e r i o d Breeding /79 Breeding /80 (13,909) a (6,408) Ecosystems (2df) 4,880 b 4,532 T o t a l areas occupied by t e r r i t o r i e s (m ) 2 X v a l u e ; P f o r a l l t e s t s < 0.001. TABLE 19 iJSJf I^ FDR 1 ^,S9 , S I N W 1 I I C " ™ H OBSERVED AREAS (m2) OF MALE TERRITORIES AND FEMALE HOME RANGES WITHIN THE VARIOUS ECOSYSTEMS WERE GREATER THAN THE EXPECTED VALUES Ecosystems Locations o IO rH O IO Q 3 a i in oo IA in io co to rH IO CJ to to o Ch CJ Q 00 Study Area T e r r i t o r i e s (4 census periods) Study P l o t s T e r r i t o r i e s (5 census periods) Home Ranges (1 census period) Removal P l o t E T e r r i t o r i e s (2 census periods) TABLE 20 NUMBERS OF CENSUS PERIODS IN WHICH THE OBSERVED AREAS Cm2) OF MALE TERRITORIES AND FEMALE HOME RANGES WITHIN THE VARIOUS ECOSYSTEMS GROUPED BY FOREST FLOOR HABITATS WERE GREATER THAN THE EXPECTED VALUES Locations Forest Floor Habitats Moss/litter Study Area Territories (4 census periods) Salal Sword fern Salmonberry Study Plots Territories (5 census periods) Home Ranges (1 census period) 0 1 0 0 5 0 4 1 0"> 167 D i s p r o p o r t i o n a t e l y g r e a t e r areas of t e r r i t o r i e s o c c u r r e d on c e r t a i n ecosystems and grouped ecosystems. G e n e r a l l y the p a t t e r n s of d i f f e r e n c e s from the expected v a l u e s f o r the study area, study p l o t s , and removal p l o t E were s i m i l a r . However, because the study p l o t s were pa r t of the study area and because the ecosystems and f o r e s t f l o o r h a b i t a t s of removal p l o t E were d i f f e r e n t from those of the study a r e a , the r e s u l t s of the censuses of the d i f f e r e n t areas were not a d d i t i v e but served as checks on one another. Male t e r r i t o r i e s on the study area and study p l o t s occupied d i s p r o p o r t i o n a t e l y l a r g e r areas i n descending order of rank of the ecosystems TIARELLA - POLYSTICHUM -WRC, 71.4/71.89, MOSS - (POLYSTICHUM) - WH - WRC, 31.412, RUBUS - POLYSTICHUM - WRC, 72.4/72.47, Vaccinium -L y s i c h i t u m - WRC, 911.0, and Ribes - Oplopanax - WRC, 812.89 suggesting p r e f e r e n c e f o r these ecosystems by winter wrens f o r t h e i r t e r r i t o r i e s . Observed u t i l i z a t i o n of Mahonia - G a u l t h e r i a - WH - DF, 132.9 i n d i c a t e d no marked p r e f e r e n c e f o r t h i s ecosystem on the study area but i n d i c a t e d avoidance on the study p l o t s (see f o l l o w i n g s e c t i o n ) . The ecosystems POLYPODIUM - GAULTHERIA - DF -WRC, 62.90, Moss - WH, 211.4, G a u l t h e r i a - WH - DF, 131.30/131.4, and (LICHEN) - GAULTHERIA - DF, 11.30 were 168 u t i l i z e d f o r t e r r i t o r i e s much l e s s than expected than i f the ecosystems had been u t i l i z e d randomly. Winter wren t e r r i t o r i e s on removal p l o t E occupied d i s p r o p o r t i o n a t e l y l a r g e r areas of the ecosystem MOSS - (POLYSTICHUM) - WH -WRC, 31.12, showed no marked pr e f e r e n c e f o r Moss - WH, 211.23 and u t i l i z e d the Mahonia - Moss - WH - WRC, 212.89 l e s s than expected. I t should be noted that the p r e f e r e n t i a l o r d e r i n g s of the ecosystems are approximate, as some of the ecosystems were present i n r e l a t i v e l y low q u a n t i t i e s . Most but not a l l female home ranges on the study p l o t s were i d e n t i f i e d d u r i n g the breeding p e r i o d census, 1980. Although the data are l e s s complete than those f o r males, the observed u t i l i z a t i o n of ecosystems may i n d i c a t e t hat females are l e s s s e l e c t i v e of h a b i t a t than are males (Table 19). C o n s i s t e n t p r e f e r e n c e by male winter wrens was evident among the ecosystems grouped by f o r e s t f l o o r h a b i t a t s f o r the census of the study area and study p l o t s (Table 20). T e r r i t o r i e s occupied c o n s i s t e n t d i s p r o p o r t i o n a t e l y l a r g e r areas of the ecosystems grouped by sword f e r n f o r e s t f l o o r h a b i t a t , frequent d i s p r o p o r t i o n a t e l y l a r g e r areas of ecosystems grouped by salmonberry f o r e s t f l o o r h a b i t a t , but showed a r e l a t i v e avoidance of the ecosystems grouped 169 by m o s s / l i t t e r and s a l a l f o r e s t f l o o r h a b i t a t s . The data tor female home ranges were again l e s s complete than those f o r males. D i s t r i b u t i o n of winter wren o b s e r v a t i o n s among the ecosystems and ecosystems grouped by f o r e s t f l o o r h a b i t a t s The areas (m 2) of ecosystems and f o r e s t f l o o r h a b i t a t s occupied by i n d i v i d u a l male t e r r i t o r i e s and female home ranges were not u t i l i z e d e q u a l l y by winter wrens. Some areas were u t i l i z e d i n t e n s i v e l y while o t h e r s were s c a r c e l y u t i l i z e d . For i n s t a n c e , the ecosystems Mahonia -G a u l t h e r i a - WH - DF, 132.9 and POLYPODIUM - GAULTHERIA -DF - WRC, 62.90 of the study area g e n e r a l l y l a y adjacent to ecosystems p r e f e r r e d by winter wrens ( F i g u r e 18), and, while i n c l u d e d w i t h i n t e r r i t o r i e s and home ranges (Table 19), were u t i l i z e d r e l a t i v e l y l e s s i n t e n s i v e l y by the b i r d s . In a d d i t i o n , w i t h i n some ecosystems and ecosystems grouped by f o r e s t f l o o r h a b i t a t s , t e r r i t o r i e s and home ranges were more ephemeral than o t h e r s , o f t e n l a s t i n g only p a r t of a census p e r i o d . Such p a t t e r n s of u t i l i z a t i o n are b e t t e r shown by a c o n s i d e r a t i o n of winter wren o b s e r v a t i o n s than by areas (m 2) of t e r r i t o r i e s and home ranges. 170 Here I compare the d i s t r i b u t i o n s of w i n t e r wren o b s e r v a t i o n s w i t h the d i s t r i b u t i o n s of K l i n k a ' s ecosystems and ecosystems grouped by f o r e s t f l o o r h a b i t a t s . As was the case f o r the d i s t r i b u t i o n s of the a r e a s (m 2) of male t e r r i t o r i e s , w i n t e r wrens a r e shown t o u t i l i z e the ecosystems and grouped ecosystems d i f f e r e n t i a l l y , and t o p r e f e r c o n s i s t e n t l y some ecosystems and f o r e s t f l o o r h a b i t a t s t o o t h e r s . S i n g i n g and c h i p p i n g w i n t e r wrens were o b s e r v e d w i t h r e l a t i v e l y e q u a l e f f i c i e n c y i n a l l ecosystems and f o r e s t f l o o r h a b i t a t s . S i l e n t b i r d s p r o b a b l y were found more e f f i c i e n t l y i n ecosystems and f o r e s t f l o o r h a b i t a t s w i t h o u t heavy u n d e r s t o r y c o v e r a g e . These a r e a s i n c l u d e d the m o s s / l i t t e r and, t o a l e s s e r e x t e n t , the s a l a l f o r e s t f l o o r h a b i t a t s throughout the y e a r , and the salmonberry f o r e s t f l o o r h a b i t a t d u r i n g t h e w i n t e r and e a r l y b r e e d i n g p e r i o d s when the salmonberry was l e a f l e s s . Such e f f i c i e n c i e s a l s o a p p l y t o the ecosystems grouped by t h e s e f o r e s t f l o o r h a b i t a t s ( T a b l e s 11 and 15). Because th e s e e f f i c i e n c i e s g e n e r a l l y ran c o u n t e r t o the o b s e r v e d d i s t r i b u t i o n of w i n t e r wren o b s e r v a t i o n s , they were i g n o r e d i n the comparisons and any apparent d i f f e r e n c e s a r e c o n s e r v a t i v e . 171 The observed d i s t r i b u t i o n s of the three c a t e g o r i e s of winter wren o b s e r v a t i o n s on the v a r i o u s ecosystems and ecosystems grouped by f o r e s t f l o o r h a b i t a t s were compared, as were the areas (m 2) of male t e r r i t o r i e s and female home ranges, w i t h the corresponding expected v a l u e s of the v a r i o u s ecosystems u s i n g X 2 t e s t s (Tables 21 to 23). The s t a t i s t i c a l comparisons of the expected and observed d i s t r i b u t i o n s of the numbers of o b s e r v a t i o n s were g e n e r a l l y h i g h l y s i g n i f i c a n t (P < 0.001) f o r the study area, study p l o t s , and removal p l o t E. F a i l u r e t o depart from the expected d i s t r i b u t i o n at t h i s l e v e l of s i g n i f i c a n c e only o c c u r r e d when the number of o b s e r v a t i o n s was £ 100. These data a l s o i n d i c a t e a d i f f e r e n t i a l u t i l i z a t i o n of the v a r i o u s ecosystems and f o r e s t f l o o r h a b i t a t s by winter.wrens. Expected numbers of o b s e r v a t i o n s of winter wrens w i t h i n the v a r i o u s ecosystems and ecosystems grouped by f o r e s t f l o o r h a b i t a t s were c a l c u l a t e d from t o t a l areas of the v a r i o u s ecosystems and f o r e s t f l o o r h a b i t a t s based on p r o p o r t i o n a l random d i s t r i b u t i o n of o b s e r v a t i o n s . The numbers of o b s e r v a t i o n s found d u r i n g the v a r i o u s census p e r i o d s of the study areas d i f f e r e d from the expected values (Tables 24 and 25). The p a t t e r n s of the d i f f e r e n c e s from the expected v a l u e s f o r the v a r i o u s study TABLE 21 STATISTICAL COMPARISONS (X 2) OF THE EXPECTED AND OBSERVED DISTRIBUTIONS OF THE NUMBERS OF WINTER WREN OBSERVATIONS AMONG THE ECOSYSTEMS AND ECOSYSTEMS GROUPED BY FOREST FLOOR HABITATS DURING THE VARIOUS CENSUS PERIODS OF THE STUDY AREA, 1978-81 Census Period Breeding /78 Breeding /80 Breeding /81 Breeding /82 (400; 171 (494; 199 (525; 310 (977; 100 571) a 693) 835) 1,097) Ecosystems (9df) 49.7; 23.8 d; 77.5; 35.1; 93.7; 43.8; 57.1; 10.7 C; 63.6 b 99.2 119.7 50.2 Forest Floor Habitats from 43.1; 15.7; 58.6; 17.8; 88.3; 22.6; 37.4; i . i c ; Grouped Ecosystems (3df) 58.3 74.4 106.0 32.4 Numbers of observations; ( t e r r i t o r i a l males; other birds; t o t a l winter wren observations). b 2 X values; t e r r i t o r i a l males; other birds; t o t a l winter wren observations. c not s i g n i f i c a n t . d P < 0.01.; P for a l l other x 2 values ( 0.001. TABLE 22 STATISTICAL COMPARISONS (X 2) OF THE EXPECTED AND OBSERVED DISTRIBUTIONS OF THE NUMBERS OF WINTER WREN OBSERVATIONS AMONG THE ECOSYSTEMS AND ECOSYSTEMS GROUPED BY FOREST FLOOR HABITATS DURING THE VARIOUS CENSUS PERIODS OF THE STUDY PLOTS, 1979-80 Units Ecosystems (9df) 8 2 ) a 37.4 C Census Period 13.3 C; 2 5 . l e ; 261.3; 269.8 484.4 148.5 155.2 r% J r l ! ! d 1 ^ / 7 9 F a l 1 / 7 9 W i n t e r / ? 9 Breeding /80 (23; 59; (866; 603; (104; 214; (146; 162; (1,212; 784; M 6 9 ) 318) 308) 1,996) 53.1; 130.4; 80.2; 87.3; 428.6; 249.6; 645.8 Forest Floor Habitats from Grouped Ecosystems (3df) 10.9 ; 19.6; 254.0; 188.5; 49.7; 64.3; 57.6; 57.4; 402.9; 203.4; 2 9 - 8 439.6 112.1 114.2 599.4 a Numbers o f observations; ( t e r r i t o r i a l males; other b i r d s ; t o t a l winter wren observations). X values; t e r r i t o r i a l males; other b i r d s ; t o t a l winter wren observations. c Not s i g n i f i c a n t . d P < 0 . 0 5 . e P < 0.01. P f o r a l l other x 2 values < 0.001. -J U) TABLE 23 STATISTICAL COMPARISONS (X2) OF THE EXPECTED AND OBSERVED DISTRIBUTIONS OF THE NUMBERS OF WINTER WREN OBSERVATIONS AMONG THE ECOSYSTEMS DURING ALL CENSUSES OF REMOVAL PLOT E, 1979-80 Census P e r i o d U n i t s Breeding /79 Breeding /80 (28; 8; (31; 6; 3 6 ) a 37) Ecosystems (2df) 21.5 d; 29.0; 51.8; 1.9 C; 42.3 b 44.6 Numbers o f observations ( t e r r i t o r i a l males; other b i r d s ; t o t a l w i n t e r wren o b s e r v a t i o n s ) . 2 X v a l u e s ; t e r r i t o r i a l males; other b i r d s ; t o t a l w i n t e r wren o b s e r v a t i o n s , not s i g n i f i c a n t . P f o r a l l other x 2 values < 0.001. TABLE; 24 NUMBERS OF CENSU p m m s J f J m m { ^ OBSERVED NUMBERS OF OBSERVATIONS OF WINTER WRENS WITHIN THE VARIOUS ECOSYSTEMS WERE GREATER THAN T11E EXPECTED VALUES Ecosystems o Locations o Q i < OS •o r-l O u, Q ro Q 10 (si to Q to Q 00 Study Area (4 census periods) 0 ; l ; 0 a 0;0;0 0;0;0 0;1;0 Study P l o t s (5 census periods) 0;0;0 0;0;0 0;2;1 0;0;0 Removal P l o t E (2 census periods) 4;2;4 1;1;1 4;4;4 4 0;0;0 0;0;0 5;5;5 3 0;1;0 0;0;0 2;2;2 T e r r i t o r i a l males; other b i r d s ; t o t a l winter wren observati TABLE 25 NUMBERS OF CENSUS PERIODS IN WHICH THE OBSERVED NUMBERS OF OBSERVATIONS OF WINTER WRENS WITHIN THE VARIOUS ECOSYSTEMS GROUPED BY FOREST FLOOR HABITATS WERE GREATER THAN THE EXPECTED VALUES Locations Forest F l o o r H a b i t a t s M o s s / l i t t e r S a l a l Sword f e r n Salmonberry Study Area (4 census periods) 0 ; l ; 0 a 0;1;0 4;3;4 2;0;2 Study P l o t s (5 census periods) 0;0;0 0;0;0 5;5;5. 3;2;1 T e r r i t o r i a l males; other b i r d s ; t o t a l w i n t e r wren observations. 177 areas were again g e n e r a l l y s i m i l a r . In a d d i t i o n , the general p a t t e r n of d i f f e r e n c e s i n the d i s t r i b u t i o n of winter wren o b s e r v a t i o n s was s i m i l a r to that found f o r d i f f e r e n c e s f o r areas (m 2) of t e r r i t o r i e s and home ranges. Numbers of winter wren o b s e r v a t i o n s on the study area and study p l o t s were d i s p r o p o r t i o n a t e l y g r e a t e r than expected, i n descending order of rank, w i t h i n the ecosystems TIARELLA - POLYSTICHUM - WRC, 71.4/71.89 and RUBUS - POLYSTICHUM - WRC, 72.4/72.47 suggesting a pre f e r e n c e f o r these ecosystems by winter wrens. Greater than expected numbers of o b s e r v a t i o n s a l s o o c c u r r e d w i t h i n the ecosystem MOSS - (POLYSTICHUM) - WH - WRC, 31.412 on the study area but not on the study p l o t s . T h i s ecosystem oc c u r r e d on the study p l o t s adjacent to the ecosystem TIARELLA - POLYSTICHUM - WRC, 71.4/71.89 which a l s o o c c u r r e d with i t on the i n d i v i d u a l t e r r i t o r i e s and home ranges. Under these circumstances, TIARELLA - POLYSTICHUM - WRC, 71.4/71.89 was p r e f e r r e d by the b i r d s and accounted f o r the lower numbers of o b s e r v a t i o n s on MOSS -POLYSTICHUM) - WH - WRC, 31.412. Elsewhere over the study area, and on removal p l o t E, the ecosystems MOSS -(POLYSTICHUM) - WH - WRC, 31.412 and 31.12 oc c u r r e d on t e r r i t o r i e s and home ranges with l e s s p r e f e r r e d ecosystems and was i t s e l f p r e f e r r e d . 178 The observed u t i l i z a t i o n of the ecosystems Ribes -Oplopanax - WRC, 812.89 and Vaccinium - Lysichichum - WRC, 911.0 i n d i c a t e d no marked p r e f e r e n c e by winter wrens f o r these ecosystems, whereas the ecosystems POLYPODIUM -GAULTHERIA - DF - WRC, 62.90, Mahonia - G a u l t h e r i a - WH -DF, 132.9, Moss - WH, 211.4, (LICHEN) - GAULTHERIA - DF, 11.30 and G a u l t h e r i a - WH - DF, 131.30/131.4 were u t i l i z e d , i n d e c r e a s i n g order, by wrens much l e s s than expected than i f the ecosystems had been u t i l i z e d randomly. The ecosystems Mahonia - G a u l t h e r i a - WH - DF, 132.9 and POLYPODIUM - GAULTHERIA - DF - WRC, 62.90 d i d not show g r e a t e r than expected numbers of winter wren o b s e r v a t i o n s . These f i n d i n g s support the o b s e r v a t i o n s made e a r l i e r t h a t these ecosystems showed g r e a t e r than expected areas (m 2) of male t e r r i t o r i e s on the study area and of female home ranges on the study p l o t s because of t h e i r p r o x i m i t y to p e f e r r e d ecosystems (Table 19). C o n s i s t e n t p r e f e r e n c e s were again shown by winter wrens, based on numbers of o b s e r v a t i o n s , among the ecosystems grouped to f o r e s t f l o o r h a b i t a t s f o r the census p e r i o d s of a l l the study areas (Table 25). D i s p r o p o r t i o n a t e l y higher numbers of o b s e r v a t i o n s c o n s i s t e n t l y o c c u r r e d w i t h i n the ecosystems grouped to sword f e r n f o r e s t f l o o r h a b i t a t . Frequent 179 d i s p r o p o r t i o n a t e l y higher numbers of o b s e r v a t i o n s o c c u r r e d i n the ecosystems grouped to areas of salmonberry f o r e s t f l o o r h a b i t a t but there was r e l a t i v e avoidance of the ecosystems grouped to m o s s / l i t t e r and s a l a l f o r e s t f l o o r h a b i t a t s . S c o u l l a r (1980), who a l s o worked at the Research F o r e s t , found a d i f f e r e n t i a l d i s t r i b u t i o n of numbers of o b s e r v a t i o n s of winter wrens among v e g e t a t i o n types ( p l a n t a s s o c i a t i o n s and s u b - a s s o c i a t i o n s of a v a r i e t y of s e r a i stages of f o r e s t v e g e t a t i o n grouped at the l e v e l of p l a n t a l l i a n c e - see Appendices 1 and 2 ) . He a l s o found a strong p r e f e r e n c e by winter wrens f o r o l d e r s e r a i stages of f o r e s t v e g e t a t i o n , as, f o r example, the f o r e s t v e g e t a t i o n i n v e s t i g a t e d i n t h i s study. Because he observed r e l a t i v e l y more winter wrens i n these s e r a i stages, he thought the s i g n i f i c a n t d i f f e r e n c e s i n the use of v e g e t a t i o n types by winter wrens to be i n a d v e r t e n t . As a consequence, and i n c o n t r a s t to the f i n d i n g s of t h i s study, S c o u l l a r thought that winter wrens used most v e g e t a t i o n types at s i m i l a r l e v e l s although moist v e g e t a t i o n types were p r e f e r r e d . Winter wren p r e f e r e n c e s found i n t h i s study f o r ecosystems and ecosystems grouped by f o r e s t f l o o r h a b i t a t s were g e n e r a l l y c o n s i s t e n t f o r the study area and study 1 8 0 p l o t s as i n d i c a t e d by the areas (m 2) of male t e r r i t o r i e s and the numbers of o b s e r v a t i o n s (Table 26). The ecosystems grouped by the m o s s / l i t t e r and s a l a l f o r e s t f l o o r h a b i t a t s were g e n e r a l l y avoided by winter wrens both as i n d i v i d u a l ecosystems and as grouped ecosystems. The ecosystems grouped by the salmonberry f o r e s t f l o o r h a b i t a t , both as i n d i v i d u a l ecosystems and as grouped ecosystems, were f r e q u e n t l y p r e f e r r e d by male winter wrens f o r t h e i r t e r r i t o r i e s , but, as i n d i c a t e d by numbers of o b s e r v a t i o n s , were u t i l i z e d by the b i r d s without marked p r e f e r e n c e . The ecosystems grouped to the sword f e r n f o r e s t f l o o r h a b i t a t were g e n e r a l l y , both as i n d i v i d u a l ecosystems and as grouped ecosystems, f r e q u e n t l y to s t r o n g l y p r e f e r r e d by winter wrens. The exception was the ecosystem MOSS - (POLYSTICHUM - WRC - WH, 31.412 which was s t r o n g l y p r e f e r r e d by males f o r t h e i r t e r r i t o r i e s but which was u t i l i z e d without p r e f e r e n c e as i n d i c a t e d by numbers of o b s e r v a t i o n s . The d i s t r i b u t i o n s of winter wrens on the removal p l o t s D and E and on the study area were not compared s t a t i s t i c a l l y due to the l a c k of exact e q u i v a l e n c y of ecosystems. However, study p l o t B c o n s i s t e d l a r g e l y of the ecosystems TIARELLA- POLYSTICHUM - WRC, 71.4/71.89 which were very s i m i l a r to the ecosystems TIARELLA -TABLI-: 26 SUN WARY OF WINTER WREN PREFERENCES FOR ECOSYSTEMS AND ECOSYSTEMS GROUPED BY FOREST FLOOR HABITATS Ecosystems Ecosystems Grouped by Forest Floor Habitats 3 Winter Wren Preference^ Study Area (Including Study Plots) Mahonia-Gaultheria - WH-DF, 132.9 Moss - WH, 211.4 (LICHEN)-GAWLTHERIA - DF, 11.30 Gaultheria - WH-DF, 131.30/131.4 POLYPODIUM-GAULTHERIA - DF-WRC, 62.90 MOSS-(POLYSTICHUM) - WRC-WH, 31.412 TIARELI^POLYSTImiM - WRC, 71.4/71.89 RUBUS-POLYSTICHUM - WRC, 72.4/72.47 Ribes-Oplopanax - WRC, 812.89 Vaccinium-Lysichitum - WRC, 911.0 Removal Plot D Areas- (m2) of Male Territories Moss/litter Salal Sword fern Salmonberry TIARELLA-POLYSTICHUM - WRC, 71.12/71.4/71.89 ) Sword fern Removal Plot E Moss - WH, 211.23 Mahonia-Moss - WRC-WH, 212.89 MOSS^TPOLYSTICHUM) - WH-WRC, 31.12 Moss/litter - ) -;}-:}• (+)j l N/Ae N/A Numbers of Observations N/A N/A Average Total Vegetation Cover (%) e Understory Middlestory (C) 41 62 35 52 }' 2} 26 60 ) 60 6 1 21 (B2) 20 8 16 11 26 68 62 \ 1 | 48 } 1 '4 37 M7 ) 10 J 39 J 4 )4 Ranked Total Volumes of Invertebrates from the Forest Floor Habitats (l-highest)d N/A N/A Groupings of the ecosystems by forest floor habitats are shown for the study area and removal plots in Tables 11 and 15 respectively. i_ Winter wren preference for ecosystems and ecosystems grouped by forest floor habitats are derived from Tables 19 and 29 (areas -m2) and Tables 24 and 25 (numbers of observations) as follows -, avoidance. + , strong preference; (+), frequent preference; 0, no marked preference; 'Vegetation cover values are taken for ecosystems from Table 9 and for ecosystems grouped by forest floor habitats from Table 13. dRankings of total volumes of invertebrates from the forest floor habitats are taken from Table 33. Not applicable. 182 POLYSTICHUM - WRC, 71.12/71.4/71.89 of removal p l o t D. The ecosystems of study p l o t B and removal p l o t D were a l s o g e n e r a l l y grouped to the same f o r e s t f l o o r h a b i t a t -sword f e r n ( F i g u r e s 18'and 44; Tables 9 and 13). The numbers and p a t t e r n s of d i s t r i b u t i o n of male t e r r i t o r i e s d u r i n g the breeding p e r i o d on removal p l o t D were very s i m i l a r t o , and c o n s i s t e n t with, those of study p l o t B (F i g u r e s 9, 12, and 13). The ecosystems of removal p l o t E were u t i l i z e d i n the same gen e r a l p a t t e r n as were the s i m i l a r ecosystems of the study area (Table 26) with the exceptions of the ecosystem MOSS - WH, 211.23 which the males d i d not a v o i d f o r t h e i r t e r r i t o r i e s , and the ecosystem MOSS - POLYSTICHUM) - WH -WRC, 31.12 which was s t r o n g l y p r e f e r r e d . In a d d i t i o n , the numbers and d i s t r i b u t i o n s of male t e r r i t o r i e s d u r i n g the breeding p e r i o d on removal p l o t E were s i m i l a r t o , and c o n s i s t e n t with, those of s i m i l a r ecosystems on study p l o t s A and B of the study area ( F i g u r e s 9, 12, and 13). The c o n s i s t e n t p a t t e r n s of d i f f e r e n t i a l u t i l i z a t i o n of ecosystems and ecosystems grouped by f o r e s t f l o o r h a b i t a t s by winter wrens i n the o l d e r s e r a i stages of f o r e s t v e g e t a t i o n documented i n t h i s study suggest t h a t the d i s t r i b u t i o n of winter wrens on the study area and study p l o t s were clumped by s u i t a b l e h a b i t a t . The c o n s i s t e n c y 183 of the p a t t e r n s of d i f f e r e n t i a l u t i l i z a t i o n of ecosystems and ecosystems grouped by f o r e s t f l o o r h a b i t a t s by winter wrens on the study a r e a , study p l o t s , and removal p l o t s throughout the breeding, f a l l and winter p e r i o d s s t r o n g l y suggest that K l i n k a ' s ecosystems and ecosystems grouped by f o r e s t f l o o r h a b i t a t s are i n d i c a t i v e of winter wren d i s t r i b u t i o n . F u r t h e r , I propose that K l i n k a ' s ecosystems and ecosystems grouped by f o r e s t f l o o r h a b i t a t s are as i n d i c a t i v e of winter wren d i s t r i b u t i o n as they are (grouped to s u i t a b l e c r i t e r i a ) of the a t t r i b u t e s which were o r i g i n a l l y used to d e f i n e the ecosystems ( v e g e t a t i o n , parent m a t e r i a l s , s o i l s , and h y d r o l o g i c and n u t r i e n t regimes). However, the a b i l i t y of K l i n k a ' s ecosystems to i n d i c a t e winter wren d i s p e r s i o n elsewhere would r e q u i r e f u r t h e r t e s t i n g . The d i s t r i b u t i o n of f o r e s t b i r d s has o f t e n been shown to be a f u n c t i o n of v e g e t a t i o n (e.g., L a u r e n z i et a l . 1982; P i e t z and T e s t e r 1982; S c o u l l a r 1980; Morse 1979; Franzreb 1978; Wishart and Bi d e r 1976; Anderson and Shugart 1974; P a r t r i d g e 1974; Sturman 1968). However, very l i t t l e i s known about why b i r d s p r e f e r one v e g e t a t i o n to another, or how they choose a p p r o p r i a t e v e g e t a t i o n . In t h i s study, given the d i f f e r e n c e s i n cover, and s p e c i e s composition of the v e g e t a t i o n of the ecosystems and 184 grouped ecosystems (Tables 9, and 13, and 26), i t i s apparent that the v e g e t a t i o n may i n f l u e n c e the h a b i t a t s e l e c t i o n and d i s p e r s i o n of winter wrens. The combined v e g e t a t i o n cover of the understory and B 2 - m i d d l e s t o r y v e g e t a t i o n l a y e r s , i n which the winter wrens l i v e , was l e a s t i n the ecosystems grouped by the m o s s / l i t t e r f o r e s t f l o o r h a b i t a t , both as i n d i v i d u a l ecosystems and as a group (Table 26). The low a v a i l a b i l i t y of cover may have made these ecosystems and grouped ecosystems l e s s s u i t a b l e f o r winter wrens due t o the l a c k of s h e l t e r from adverse weather and enemies, and i t may h e l p e x p l a i n the apparent avoidance (Table 26) of these areas by winter wrens. The i n d i v i d u a l ecosystems grouped by the s a l a l f o r e s t f l o o r h a b i t a t , both as i n d i v i d u a l ecosystems and as grouped ecosystems were avoided by winter wrens (Table 26), although they had o n l y s l i g h t l y l e s s combined understory and B 2 - m i d d l e s t o r y v e g e t a t i o n cover than the i n d i v i d u a l and grouped ecosystems of the sword f e r n and salmonberry f o r e s t f l o o r h a b i t a t s . In the s a l a l , winter wrens moved awkwardly and with d i f f i c u l t y , both on the f o r e s t f l o o r and from branch to branch, and f l i g h t u s u a l l y o c c u r r e d above the f o r e s t f l o o r v e g e t a t i o n . These o b s e r v a t i o n s were i n d i r e c t c o n t r a s t to the r e l a t i v e ease 185 of movement i n the other f o r e s t f l o o r h a b i t a t s . The l i m i t a t i o n s on winter wren movement imposed by the s t r u c t u r e of s a l a l , and by the f u n c t i o n a l and s t r u c t u r a l c h a r a c t e r i s t i c s of winter wrens may have r e s u l t e d i n a lower s u i t a b i l i t y f o r winter wrens of the i n d i v i d u a l ecosystems and grouped ecosystems of the s a l a l f o r e s t f l o o r h a b i t a t , and may h e l p account f o r the avoidance of these areas by winter wrens. The deciduous salmonberry which dominated the B 2 -m i d d l e s t o r y v e g e t a t i o n of the salmonberry f o r e s t f l o o r h a b i t a t i s without s h e l t e r i n g f o l i a g e from l a t e f a l l to e a r l y s p r i n g , as opposed to the f o l i a g e s of s a l a l and swordfern which p r o v i d e year-round s h e l t e r . As a r e s u l t , the ecosystems grouped by the salmonberry f o r e s t f l o o r h a b i t a t , both as i n d i v i d u a l ecosystems and as grouped ecosystems, may a l s o have been s l i g h t l y l e s s s u i t a b l e f o r winter wrens because of lack of s h e l t e r d u r i n g the winter p e r i o d . Lack of winter s h e l t e r may h e l p e x p l a i n the lower p r e f e r e n c e (frequent p r e f e r e n c e to no marked p r e f e r e n c e ) e x h i b i t e d by winter wrens f o r these areas i n comparison to the strong p r e f e r e n c e shown f o r the sword f e r n f o r e s t f l o o r h a b i t a t . 186 On the other hand, the s t r u c t u r e of the sword f e r n of the ecosystems grouped to the sword f e r n f o r e s t f l o o r h a b i t a t , both as i n d i v i d u a l ecosystems and as grouped ecosystems, appeared w e l l s u i t e d as s h e l t e r and to both winter wren f l i g h t through the clumps of swordfern, and to the s k i p p i n g movement of winter wrens on the f o r e s t f l o o r i n v o l v i n g both legs and wings ( G r i n n e l l and S t o r e r 1924), and may h e l p account f o r the st r o n g p r e f e r e n c e shown by winter wrens f o r these areas (Table 26). Winter Wren D i s p e r s i o n As A F u n c t i o n Of Food I n t r o d u c t i o n Food has long been thought to be a f a c t o r i n the d i s p e r s i o n of b i r d s . In p a r t i c u l a r , food has been shown to be c o r r e l a t e d with t e r r i t o r y s i z e i n ovenbirds ( S e i u r u s  a u r o c a p i l l u s ) (Stenger 1958), d u n l i n ( C a l i d r i s a l p i n a ) (Holmes 1970), red grouse (Lagopus lagopus) ( M i l l e r et a l . 1970; Lance 1978), European and North A t l a n t i c I s l a n d subspecies of wrens (Cody and Cody 1972a, b) and rufous hummingbirds (Selasphorus r u f u s ) (Gass et a l . 1976; Gass 1978, 1979). More r e c e n t l y however, Zach and F a l l s (1979) have shown that the l a r g e s c a l e n e g a t i v e c o r r e l a t i o n between food supply and t e r r i t o r y s i z e found by Stenger 187 (1958) a c r o s s four h a b i t a t types d i d not apply at a s m a l l e r s c a l e where there was no c l o s e c o r r e l a t i o n between the s i z e of f o r a g i n g areas or song t e r r i t o r i e s and food supply. In t h i s study, due to the complexity of the mosaic d i s t r i b u t i o n of f o r e s t f l o o r h a b i t a t s w i t h i n i n d i v i d u a l t e r r i t o r i e s I was unable to compare t e r r i t o r y s i z e and food supply. However, here I do demonstrate that the t o t a l volumes of i n v e r t e b r a t e s a v a i l a b l e to winter wrens were g r e a t e r i n the f o r e s t f l o o r h a b i t a t s p r e f e r r e d by winter wrens. Food i s proposed as a f a c t o r i n winter wren d i s p e r s i o n . Methods I n d i v i d u a l stomach contents of 122 winter wrens (91, 25, and 6 from breeding, f a l l , and winter p e r i o d s r e s p e c t i v e l y ) c o l l e c t e d d u r i n g the study were preserved i n a l c o h o l f o r l a t e r a n a l y s i s . The i d e n t i f i a b l e contents were composed almost e n t i r e l y of i n v e r t e b r a t e s p l u s a few seeds. I i d e n t i f i e d the i n v e r t e b r a t e s to f a m i l y wherever p o s s i b l e , although i n some cases i d e n t i f i c a t i o n was only p r a c t i c a b l e to order or c l a s s (Appendix 4). The numbers of i n d i v i d u a l s and the percentage volumes f o r the v a r i o u s taxa were estimated f o r the contents of each winter wren 188 stomach. However, the q u a n t i t i e s of i n v e r t e b r a t e s i n the stomachs were too small ( u s u a l l y about 0.05 ml) t o permit measurement of a c t u a l volumes. I sampled the i n v e r t e b r a t e s from the f o r e s t f l o o r v e g e t a t i o n and upper l i t t e r combined, w i t h i n the four f o r e s t f l o o r h a b i t a t s of the study p l o t s , over the winter p e r i o d , 1979-80, and the breeding p e r i o d , 1980. At approximately two week i n t e r v a l s two, one-m 2 i n v e r t e b r a t e sample p l o t s were sampled on each f o r e s t f l o o r h a b i t a t f o r a t o t a l of 10 p l o t s per f o r e s t f l o o r h a b i t a t per p e r i o d . The p l o t s were chosen to represent the f o r e s t f l o o r h a b i t a t s and the v a r i o u s ecosystems w i t h i n the f o r e s t f l o o r h a b i t a t s . I d e s c r i b e d the v e g e t a t i o n cover of the m i d d l e s t o r y (B), understory (C), and the bryophyte (D) l a y e r s and the ground cover of each i n v e r t e b r a t e p l o t (p. 101). The f o r e s t f l o o r v e g e t a t i o n i n c l u d i n g the shrubs was then c u t , and the upper l i t t e r was scraped up to a depth of about 1 cm (the depth to which a winter wren c o u l d forage w i t h i t s b i l l ) . The v e g e t a t i o n and l i t t e r of each i n v e r t e b r a t e p l o t were p l a c e d together i n a c l o t h bag as shown i n the foreground of F i g u r e 50. The bags were u s u a l l y hung f o r a day or two to a l l o w excess moisture to d r i p away and then the c o n t e n t s of each bag was s p l i t to equal volumes df FIGURE 50 MODIFIED BERLESE TYPE FUNNELS USED TO EXTRACT INVERTEBRATES FROM THE SAMPLES OF FOREST FLOOR VEGETATION AND SURFACE LITTER 191 v e g e t a t i o n and l i t t e r ( s p l i t - s a m p l e s ) of as s i m i l a r compositions as p o s s i b l e . I e x t r a c t e d the i n v e r t e b r a t e s from the s p l i t samples us i n g m o d i f i e d B e r l e s e type f u n n e l s (Peterson 1953; Beirne 1962; M a r t i n 1977) as shown i n F i g u r e 50. The funnel chambers were c o n s t r u c t e d from black p l a s t i c p a i l s and had volumes of approximately 25:1. W i t h i n the chambers, the samples were supported by two spaced l a y e r s of 1.27 cm and one l a y e r of 0.64 cm mesh s c r e e n s . An example of a f u n n e l chamber i s shown on top of the f r o n t funnel stand. The o u t s i d e f u n n e l was c o n s t r u c t e d of black sheet p l a s t i c u s i n g the mould shown l e a n i n g a g a i n s t the f u n n e l stand. The l i g h t and heat f o r the f u n n e l s were s u p p l i e d by 150 watt b u l b s . Ten days to 2 weeks were r e q u i r e d to dry the s p l i t samples completely. The e x t r a c t e d i n v e r t e b r a t e s and a l c o h o l from the funnel b o t t l e s were s t o r e d i n p l a s t i c b o t t l e s as shown on the top of the funnel stand f o r l a t e r a n a l y s i s . I s o r t e d the e x t r a c t e d i n v e r t e b r a t e s from the d e b r i s which came through the e x t r a c t o r with them. Only i n v e r t e b r a t e s £ 1 mm o v e r a l l l e n g t h were c o n s i d e r e d as f i e l d o b s e r v a t i o n s and a n a l y s e s of stomach co n t e n t s suggested that winter wrens seldom took i n v e r t e b r a t e s below t h i s s i z e l i m i t . Any which they d i d take must have 1 92 c o n t r i b u t e d v e r y l i t t l e t o t h e i r f o o d . I i d e n t i f i e d t h e i n v e r t e b r a t e s f r o m t h e e x t r a c t o r s a s f o r t h e w i n t e r w r e n s t o m a c h c o n t e n t s ( A p p e n d i x 4). T h e t o t a l v o l u m e ( m l ) o f i n v e r t e b r a t e s f r o m e a c h s p l i t - s a m p l e was m e a s u r e d . T h e n u m b e r s o f i n d i v i d u a l s a n d t h e v o l u m e s f o r t h e v a r i o u s t a x a w e r e c a l c u l a t e d ( p e r c e n t a g e v o l u m e x t o t a l v o l u m e ) . The n u m b e r s o f i n d i v i d u a l s a n d v o l u m e s o f t h e i n d i v i d u a l i n v e r t e b r a t e t a x a i n t h e p a i r s o f s p l i t - s a m p l e s w e r e c o n s i s t e n t l y v e r y s i m i l a r . The p a i r s o f s p l i t - s a m p l e s w e r e t h e r e f o r e summed t o g i v e t h e t o t a l number a n d v o l u m e o f i n v e r t e b r a t e s o f e a c h t a x o n f o r e a c h i n v e r t e b r a t e s a m p l e p l o t . T h e v a r i o u s i n v e r t e b r a t e t a x a p r o b a b l y w e r e n o t e x t r a c t e d e q u a l l y by t h e f u n n e l s . I n a d d i t i o n , p r e d a t i o n , n a t u r a l m o r t a l i t y , a n d r e p r o d u c t i o n may h a v e o c c u r r e d i n t h e f u n n e l s o v e r t h e e x t r a c t i o n p e r i o d . Some o f t h e l e s s m o b i l e t a x a may h a v e r e m a i n e d a n d b e e n d e s s i c a t e d i n t h e c h a m b e r s w h i l e some o f t h e more p h o t o t r o p h i c a r t h r o p o d s o f t h e f o r e s t f l o o r v e g e t a t i o n may n o t h a v e b e e n r e p e l l e d by t h e h e a t a n d l i g h t t o t h e same d e g r e e a s t h e i n v e r t e b r a t e s o f t h e l i t t e r a n d may a l s o h a v e r e m a i n e d a n d b e e n d e s s i c a t e d i n t h e c h a m b e r s . 193 Winter wren food The r e s u l t s of the stomach co n t e n t s analyses are intended o n l y as a general i n d i c a t i o n of winter wren food. Many of the b i r d s were taken from young f o r e s t s and p l a n t a t i o n s composed of ecosystems and f o r e s t f l o o r h a b i t a t s d i f f e r e n t from those of the study area. The stomach c o n t e n t s of these b i r d s were not n e c e s s a r i l y r e p r e s e n t a t i v e of the food of the winter wrens of the study area. Only 31 b i r d s were a v a i l a b l e from sword f e r n f o r e s t f l o o r h a b i t a t , a l l taken d u r i n g the breeding p e r i o d on and around removal p l o t D. No b i r d s were taken from salmonberry f o r e s t f l o o r h a b i t a t . The winter wren stomachs c o n t a i n e d a wide d i v e r s i t y of arthropods and a few molluscs (Table 27). Due to the almost complete fragmentation of the stomach c o n t e n t s , the fr e q u e n c i e s (% occurrences) of the v a r i o u s taxa are probably b e t t e r estimates of winter wren food than numbers of i n d i v i d u a l s or percentage volumes. Frequencies are l a t e r used to rank the r e l a t i v e importance of the v a r i o u s i n v e r t e b r a t e taxa i n the food of winter wrens. Seasonal changes i n frequency of i n v e r t e b r a t e s found i n stomachs o c c u r r e d among v a r i o u s p e r i o d s p a r t i c u l a r l y among the taxa Diplopoda, Hemiptera, C o l e o p t e r a , and Hymenoptera. However, winter wren food remained d i v e r s e TABLE 27 SUMMARY OF THE WINTER WREN STOMACH CONTENTS Mean number of i n d i v i d u a l s ; frequency (I occurrence); mean percentage volume (range of numbers of i n d i v i d u a l s ) (range of percentage volumes) Invertebrate Taxa Arthropoda Diplopoda (Millipedes) Breeding(91) Period (Number of stomaches) F a l l (25) Winter(6) Totals(122) Chordeumida J u l i d a Unidentified Diplopoda Chilopoda (Centipedes) Unid e n t i f i e d Chilopoda Arachnida Pseudoscorpionida (Pseudoscorpions) Phalangida (Harvestmen) A c a r i (Mites and Ticks) Araneida (Spiders) Agelenidae *1;1;<1 (0-4)(0-15) 0;2;<1 (0-1) (0-15) < i ; i ; < i (0 1)(0- 1) <1;3;<1 (0-6)(0-100) <1;12;<1 (0-1)(0-5) <1;11;3 (0-2)(0-100) <1;21;<1 (0-5)(0-12) <1;4;<1 (0-1)(0-20) <1;8;<1 (0-1)(0-1) <1;28;5 (0-1) (0-40) 0;0;0 0;0;0 <1;8;<1 (0-1)(0-1) <1;8;5 (0-2)(0-85) 3;28;<1 (0-21)(0-8) 0;0;0 0;0;0 <1;50;8 (0-1)(0-30) <1;17;1 (0-1)(0-10) 0;0;0 0;0;0 0;0;0 <1;33;<1 (0-3)(0-2) 0;0;0 <1;3;<1 (0-4)(0-15) <1;10;4 (0-1)(0-40) *1;2;<1 (0-6)(1-10) <1;2;<1 (0-6)(0-100) <1;11;<1 (0-1)(0-5) <1;10;2 (0-2)(0-100) 1;23K1 (0-21)(0-12) <1;3;<1 (0-1)(0-20) Table 27 (continued) Invertebrate Taxa S a l t i c i d a e U n i d e n t i f i e d Araneida Insecta Collembola ( S p r i n g t a i l s ) Hypogastruridae Sminthuridae Thysanura ( B r i s t l e t a i l s ) Malachilidae Hemiptera (Bugs) Pentatomidae Tingidae Uni d e n t i f i e d Hemiptera Neuroptera (Lacewings, etc.) Mean number of i n d i v i d u a l s ; frequency (% occurrence); mean percentage volume (range of numbers of i n d i v i d u a l s ) (range o f percentage volumes) Period (Number o f stomaches) BreedingOl) F a l l (25) Winter(6) Totals (122) <1;2;<1 0;0;0 0;0;0 <1;<1;<1 (0-1) (0-5) (0-1)(0-5) 1;57;12 <1;32;3 <1;17;<1 1;51;5 (0-5) (0-90) (0-3)(0-24) (0-1)(0-3) (0-5)(0-90) <1;3;<1 (0-4)(0-10) 0;0;0 0;0;0 <1;4;<1 (0-3)(0-1) 0;0;0 <1;17;<1 (0-1)(0-2) <1;2;<1 (0-4)(0-10) <1;2;<1 (0-3)(0-2) 0;0;0 <1;4;<1 (0-1)(0-1) <1;S;1 (0-1)(0-55) 0;0;0 <1;7;1 (0-3)(0-50) *1;2;<1 (0-2)(0-25) <1;16;<1 (0-1)(0-10) <1;4;<1 (0-2)(0-3) <1;4;<1 (0-1)(0-1) 0;0;0 0;0;0 <1;<1;<1 (0-1)(0-1) 0;0;0 <1;17;<1 (0-2)(0-3) <1;17;6 (0-1)(0-35) 1;17;1 (0-5)(0-5) <1;7;<1 (0-1)(0-55) <1;2;<1 (0-2)(0-3) 1 • 7*2 (0-3)(0-50) 1;3;<1 (0-5)(0-25) Table 27 (continued) Invertebrate Taxa Coleoptera (Beetles) Carabidae Staphylinidae Byrrhidae Elateridae Cantharidae Phalacridae C o c c i n e l l i d a e Cerambycidae Curculionidae U n i d e n t i f i e d Coleoptera Diptera ( F l i e s ) Tipulidae Mean number of i n d i v i d u a l s ; frequency (% occurrence); mean percentage volume (range o f numbers of i n d i v i d u a l s ) (range o f percentage volumes) Period (Number o f stomaches) Breeding(92) F a l l ( 2 5 ) Winter(6) Totals(122) 1;30;4 (0-5)(0-100) <1;7;<1 (0-2)(0-<l) 1;31;1 (0-30(0-56) *1;1;<1 (0-1)(0-<l) <1;W;<1 (0-3)(0-<l) <1;1;<1 (0-1)(0-^1) <1;2;<1 (0-1)(0-<l) (0-1)(0-40) 1;28;5 (0-6)(0-95) 1;57;6 (0-6)(0-95) <1;44;<1 (0-2)(0-1) 41;8;<1 (0-1)(0-<d) <1;40;<1 (0-5)(0- 1) 0;0;0 <1;4;<1 (0-1)(0-<l) 0;0;0 0;0;0 0;0;0 <1;52;<1 (0-4)(0-<l) 1;92;5 (0-11) (0-75) 1;50;<1 (0-4)(0- 1) *1;17;<1 (0-1)(0-<l) <1;33;<1 (0-1)(0- 1) 0;0;0 0;0;0 0;0;0 0;0;0 0;0;0 <l ; 3 3 K l (0-2)(0-<l) 3;100;<1 (1-S)(0-<1) 2;34;1 (0-5)(0-100) l ; 7 x : i (0-2)(0- 1) 1;33;<1 (0-30)(0-56) <1;<1;<1 (0-1)(0-<l) <1;8;<1 (0-3) (0-<l) <1;<1;<1 (0-1)(0-C1) <U2;<1 (0-1)(0-<l) «fl;<l;<l (0-1)(0-40) 1;33;2 (0-6)(0-95) 5;66;4 (0-11)(0-95) <1;14;2 (0-4)(0-70) <1;4;<1 (0-1)(0-1) 0;0;0 <1;12;1 (0-4)(0-70) Table 27 (continued) Mean number of i n d i v i d u a l s ; frequency (4 occurrence); mean percentage volume (range of numbers of i n d i v i d u a l s ) (range of percentage volumes) Invertebrate Taxa Phoridae Un i d e n t i f i e d Diptera Lepidoptera (Moths and B u t t e r f l i e s ) Noctuidae Uni d e n t i f i e d Lepidoptera Hymenoptera (Bees, Wasps, Ants, etc.) Ichneumonidae Scelionidae Formicidae Unidentified Hymenoptera Unid e n t i f i e d Arthropod Fragments Mollusca Gastropoda (Snails) U n i d e n t i f i e d Gastropods U n i d e n t i f i e d Seeds Breeding(92) < i ; i ; < i (0-1) (0-4) U;14;3 (0-6) (0-70) <1;3;2 (0-1)(0-98) 1;28;10 (0-12) (0-85) 0;0;0 <1;3;<1 (0-1) (0-<l) <1;23;1 (0-2)(0-50) *1;2;<1 (0-1)(0-2) -;91;29 (0-100) <1;2;<1 (0-1)(0-5) <1;4;<1 (0-10)(0-8) Period (Number of stomaches) F a l l (25) Winter(6) 0;0;0 <1;16;<1 (0-3)(0-4) 0;0;0 <1;40;8 (0-5)(0-80) <1;4;<1 (0-1)(0-2) <1;4;<1 (0-1)(0-<l) 1;6;<1 (0-2)(0-3) 0;0;0 0;96;20 (0-90) 0;0;0 2;52;4 (0-12)(0-25) 0;0;0 0;0;0 0;0;0 <1;33;10 (0-1)(0-50) 0;0;0 <1;17;<1 (0-1)(0-<l) 0;0;0 0;0;0 -;100;9 (5-15) 0;0;0 0;0;0 Totals(122) <1;<1;<1 (0-1)(0-4) <1;14;1 (0-6)(0-70) <1;2;1 (0-1)(0-98) 1;30;9 (0-12)(0-85) <1;<1;<1 (0-1)(0-2) 1;4;<1 (0-1)(0-<l) <1;22;<1 (0-2)(0-50) <1;2;<1 (0-1)(0-2) -;92;19 (0-100) <1;2;<1 (0-1)(0-5) 2;16;1 (0-12)(0-25) 198 throughout a l l p e r i o d s . The a r t h r o p o d orders which were observed to occur at ^ 10% frequency i n winter wren stomachs were, i n descending frequency: C o l e o p t e r a , Araneida, L e p i d o p t e r a , Hymenoptera, D i p t e r a , A c a r i , Hemiptera, Pseudoscorpionida, J u l i d a , and Phalangida. Almost a l l winter wren stomachs appeared f u l l or n e a r l y so. Many stomachs c o n t a i n e d a r e s i d u e of arthropod hard p a r t s . S o f t - b o d i e d i n v e r t e b r a t e s ( A n n e l i d a , Phalangida, A c a r i , Araneida, and D i p t e r a and L e p i d o p t e r a i n c l u d i n g l a r v a e , and some Hempitera) were observed taken by winter wrens f a r more commonly than they o c c u r r e d i n the stomach c o n t e n t s . These taxa, and probably o t h e r s as w e l l , were l i k e l y underrepresented i n the stomach conte n t s , not only i n terms of numbers of i n d i v i d u a l s , f r e q u e n c i e s , and percentage volumes, but a l s o i n numbers of f a m i l i e s i d e n t i f i e d . The f i n d i n g s of t h i s study t h a t winter wrens use a wide v a r i e t y of i n v e r t e b r a t e s as food are supported g e n e r a l l y by o b s e r v a t i o n s elsewhere i n North America (e.g., P r e b l e and McAtee 1923; Bent 1948) and a l s o i n the Old World (e.g., H a r r i s s o n and Buchan 1934; Armstrong and Thorpe 1952; Armstrong 1955). 199 P o t e n t i a l w i n t e r wren food w i t h i n the ecosystems grouped by f o r e s t f l o o r h a b i t a t s Here I compare the volumes of i n v e r t e b r a t e s a v a i l a b l e as winter wren food among the f o r e s t f l o o r h a b i t a t s and compare these d i s t r i b u t i o n s with the d i s t r i b u t i o n s of winter wrens i n the ecosystems grouped by f o r e s t f l o o r h a b i t a t s . The v e g e t a t i o n of the f o r e s t f l o o r h a b i t a t s of the i n v e r t e b r a t e p l o t s was g e n e r a l l y s i m i l a r to t h a t d e s c r i b e d f o r the c o r r e s p o n d i n g f o r e s t f l o o r h a b i t a t s of the study area (Tables 28 and 13) and was t h e r e f o r e c o n s i d e r e d r e p r e s e n t a t i v e of the f o r e s t f l o o r h a b i t a t s . Summaries of the i n v e r t e b r a t e s e x t r a c t e d from the i n v e r t e b r a t e p l o t samples are shown f o r the breeding p e r i o d i n Table 29 and f o r the winter p e r i o d i n Table 30. Subordinate taxa (Appendix 4) were grouped to order and sometimes c l a s s t o f a c i l i t a t e the c a l c u l a t i o n of volumes. Because of the v a r i a t i o n i n the s i z e of the i n d i v i d u a l i n v e r t e b r a t e s both w i t h i n and among the grouped i n v e r t e b r a t e taxa, the f r e q u e n c i e s (% occurrences) and volumes are probably more r e p r e s e n t a t i v e of the v a r i o u s i n v e r t e b r a t e taxa than simple numbers of i n d i v i d u a l s . The mean f r e q u e n c i e s (% occurrences) of s e l e c t e d i n v e r t e b r a t e taxa from the winter wren stomachs (Table 27) are compared with the products of the f r e q u e n c i e s X the TABLE 28 ABRIDGED VEGETATION DATA OF THE FOREST FLOOR HABITATS OF THE INVERTEBRATE PLOTS (means) C l a s s i f i c a t i o n F o r e s t F l o o r Habitat L o c a t i o n (Study P l o t ) Number o f Invertebrate P l o t s Physiography Slope (I) Vegetation Coverage (!) B ( B i ; B 2 ; T ) a D u l l Oregon-grape Red a l d e r Red huckleberry S a l a l Salmonberry Vi n e maple Western hemlock M o s s / l i t t e r A 20 18 0;<1;<1£ 0;<1;<1 0;2;2 S a l a l A 20 22 0;<1;<1 0;69;69 0;<1;<1 Sword f e r n B 20 18 0;2;2 0;<1;<1 24;0;24 Salmonberry C 20 14 24;0;24 0;<1;<1 0;70;70 19;<1;20 1;0;1 B - (middlestory l a y e r (B 2 - small trees and shrubs between 2 and 10m i n h e i g h t ; B? - shrubs and woody p l a n t s between 0.15 and 2.0m i n height; T - t o t a l f o r B l a y e r ) . to o o Table 28 (continued) Forest F l o o r Habitat M o s s / l i t t e r Western red cedar T o t a l s 0;3;3 C T r i f o l i a t e - l e a v e d foamflower Western sword f e r n T o t a l s < 1 D Hylocomium splendens 11 Isothecium s t o l o n i f e r u m  Leucolepis m e n z i e s i i Plagiothecium undulatum 12 Pleurozium s c h r e b e r i Rhyt i d iadelphus loreus 2 S t o k e s i e l l a oregana 16 T o t a l s 44 Ground Coverage ($) Humus 55 Decayed wood 4 S a l a l Sword f e r n Salmonberry 3;0;3 3;71;72 24;2;27 44;61;82 <1 4 72 6 < 1 75 27 25 6 <1 <1 1 9 11 2 2 4 <1 1 1 <1 27 2 6 59 15 42 41 76 54 TABLE 29 SUMMARY (MEANS AND RANGES) OF THE INVERTEBRATES EXTRACTED FROM THE INVERTEBRATE PLOT SAMPLES FROM THE VARIOUS FOREST FLOOR HABITATS DURING THE BREEDING PERIOD (Only taxa f o r which the sample si z e s were s u f f i c i e n t to permit c a l c u l a t i o n o f volumes are shown) Mean number of i n d i v i d u a l s ; frequency (% occurrence); mean volume (ml) (range of numbers of i n d i v i d u a l s ) (range of volumes) Invertebrate Taxa Annelida Oligochaeta Arthropoda Crustacea M o s s / l i t t e r 0.6;10;0.01 (0-6)(0-0.13) Forest Floor Habitat S a l a l Sword f e r n 0;0;0 2.8;60;0.02 (0-9)(0-0.10) Salmonberry 18.0;100;0.06 (1-51) «0.01-0.25) Isopoda (Sow bugs) Diplopoda (Millipedes) Chilopoda (Centipedes) Arachnida Pseudoscorpionida Phalangida (Harvestmen) A c a r i (Mites and Ticks Araneida (Spiders) 0;0;0 6.7;90;0;06 (1-27) (0-0.46) 15.5;100;0.04 (3-46) «0.01-0.20) 34.7;100;0.01 (1-166) «0.01-0.04) 1.4;40;<0.01 (0-8)(0-0.02) 24.4;100;0.01 (9-56) «0.01-0.01) 73.1;100;0.04 (15-123)(0.01-0.11) 0;0;0 8.7;100;0.01 (1-37) «0.01-0.03) 9.5;100;0.02 (1-27) «0.01-0.08) 27.8;100;0.01 (17-43) «0.01-0.02) 10.3;80;0.02 (0-29) (0-0.07) 40.7;100;0.01 (17-88)(0.01-0.02) 97.2;100;0.08 (19-190)(0.03-0.16) 6.8;60;0.02 (0-47)(0-0.07) 45.9;100;0.10 (14-120)(0.02-0.23) 32.9;100;0.12 (8-65) (0.03-0.21) 70.0;100;0.02 (10-215) «0.01-0.05) 16.9;100;0.04 (4-36)(0.01-0.08) 133.2;100;0.04 (42-364)(0.01-0.08) 70.1;100;0.06 (35-117)(0.02-0.08) 23.2;100;0.06 (5-53)^0.01-0.14) 79.5;100;0.38 (12-172)(0.20-0.60) 24.8;100;0.13 (8-43)(0.06-0.22) 26.6;100;0.02 (11-75) «0.01-0.02) 9.5;100;0.02 (2-17) K0.01-0.04) 90.7;100;0.04 (35-175)(0.02-0.06) 92.4;100;0.17 (48-186)(0.10-0.34) O Table 29 (continued) Mean number of i n d i v i d u a l s ; frequency (% occurrence); mean volume (ml) (range of numbers of i n d i v i d u a l s ) (range o f volumes) Invertebrate Taxa Insecta Collembola ( S p r i n g t a i l s ) Orthoptera (Grasshoppers and Crickets) Psocoptera (Psocids) Hemiptera (True Bugs, Aphids, etc.) Coleoptera (Beetles) Diptera ( F l i e s ) Lepidoptera (Moths and B u t t e r f l i e s ) Tricoptera ( C a d d i s f l i e s ) Hymenoptera (Ants, Bees, Wasps, etc . Mollusca Gastropoda (Snails) M o s s / l i t t e r 66.3;100;0.02 (15-136) «0.01-0.05) 0.3;10;<0.01 (0-3) (0-0.01) 2.8;70;<0.01 (0-8)(0-<0.01) 0.5;40;<0.01 (0-2)(0-0.01) 56.4;100;0.07 (29-112)(0.03-0.16) 14.5;90;0.01 (0-34)(0-0.04) 0.1;10;<0.01 (0-1)(0-<0.01) 0;0;0 23.7;100;0.02 (7-52)(0.01-0.03) 0.6;30;<0.01 (0-3)(0-0.01) Forest Floor Habitat S a l a l Sword fern 258.6;100;0.07 (117-499)(0.03-0.20) 0;0;0 27.i;ioo;0.oi (5-61) «0.01-0.02) 3.2;70;<0.01 (0-9)(0-0.02) 77.9;100;0.09 (39-118)(0.03-0.23) 22.3;100;0.01 (6-73)^0.01-0.03) 0.5;20;<0.01 (0-3)(0-0.01) 0.1;10;<0.01 (0-l)(0-<0.01) 26.1;100;0.02 (3-81) «0.01-0.07) 0.3;20;<0.01 (0-2)(0-0.01) 241.4;100;0.08 (118-393)(0.02-0.14) 0.1;10;(0.01 (0-1)(0-0.04) 11.0;100 ;<0.01 (1-55)(0-0.01) S.4;80;<0.01 (0-25)«0.01-0.01) m.O;ioo;0.09 (53-183)(0.02-0.17) 60.7;100;0.04 (22-103)(0.01-0.11) 0.4;30;<0.01 (0-2)(0-0.01) 0.1;10;<0.01 (0-l)(0-<0.01) 11.1;100;0.02 (3-26) «0.01-0.02) 0.9;60;<0.01 (0-3)(0-0.01) Salmonberry 215.4;100;0.09 (97-509)(0.04-0.23) 0.1;10;<0.01 (0-1)(0- 0.01) 9.8;90;<0.01 (0-33)(0-0.01) 3.5;60;<0.01 (0-10)(0-0.01) 227.8;100;0.18 (80-399)(0.10-0.27) 190.8;100;0.10 (17-626)(<0.01-0.32) 1.4;80;0.01 (0-4)(0-0.04) 0;0;0 13.1;100;0.01 (2-43) KO. 01-0.03) 1.8;50;<0.01 (0-7)(0-0.01) TABLE 30 SUMMARY (MEANS AND RANGES) OF THE INVERTEBRATES EXTRACTED FROM THE INVERTEBRATE PLOT SAMPLES FROM THE VARIOUS FOREST FLOOR HABITATS DURING THE WINTER PERIOD (Only taxa f o r which the sample si z e s were s u f f i c i e n t to permit c a l c u l a t i o n o f volumes are shown) Mean number of i n d i v i d u a l s ; frequency (I occurrence); mean volume (ml) (range of numbers of in d i v i d u a l s ) (range of volumes) Invertebrate Taxa Annelida Oligochaeta Arthropoda Crustacea Isopoda (Sow bugs) Diplopoda (Millipedes) Chilopoda (Centipedes) Symphyla (Symphylans) Arachnida Pseudoscorpionida Phalangida (Harvestmen) A c a r i (Mites and Ticks M o s s / l i t t e r 1;30;0.01 (0-4)(0-0.03) Forest Floor Habitat S a l a l Sword f e r n 0;0;0 6;90;0.01 (0-14)(0-0.06) 3;70;0.01 (0-13)(0-0.05) 3;S0;<0.01 (0-12)(0-<0.01) 25;100;0.01 (1-124) «0.01-0.02) 1;60;<0.01 (0-3)(0-0.01) 24;100;<0.01 (4.49) «0.01-0.01) 1;30;<0.01 (0-3) (0-0.02) 0;0;0 7;90;0.02 (0-31) (0-0.06) 2;70;0.02 (0-7) (0-0.09) 11;20;<0.01 (0-1) (0-<0.01) 38;100;0.02 (2-101) «0.01-0.04) 6;100;<0.01 (1-19) «0.01-0.01) 26;100;<0.01 (6-50) «0.01-0.01) 3;90;<0.01 (0-8)(0-0.02) 3;60;0.01 (0-9)(0-0.05) 30;100;0.06 (12-55)(0.02-0.24) 7;90;0.05 (0-19)(0-0.14) 0;0;0 51;100;0.02 (9-127) «0.01-0.06) 9;100;0.01 (2-25) «0.01-0.02) 75;100;0.02 (19-173)(0.01-0.05) Salmonberry 5;90;0.06 (0-20)(0-0.54) 8;90;0.06 (0-19) (0-0.13) 54;100;0.27 (13-215)(0.01-1.02) 6;90;0.05 (0-13)(0-0.12) 0;0;0 58;100;0.02 (10-121)(0.01-0.08) 5;90;0.01 (0-9)(0-0.02) 33;100;0.02 (3-72) «0.01-0.04) Table 30 (continued) Invertebrate Taxa Araneida (Spiders) Insecta Collembola ( S p r i n g t a i l s ) Thysanura ( B r i s t l e t a i l s ) Orthoptera (Grasshoppers and Cri c k e t s ) Hemiptera (True Bugs, Coleoptera (Beetles) Diptera ( F l i e s ) Lepidoptera (Ants, Bees, Wasps, etc.) Hymenoptera Mollusca Gastropoda (Snails) Mean number of i n d i v i d u a l s ; frequency (% occurrence); mean volume (ml) (range of numbers of i n d i v i d u a l s ) (range of volumes) Forest Floor Habitat M o s s / l i t t e r S a l a l Sword f e r n Salmonberry 34;100;0.03 (12-76)(0.01-0.05) 64;100;0.07 (33-124)(0.04-0.20) 45;100;0.06 (16-70)(0.01-0.11) 39;100;0.07 (19-74)(0.02-0.12) 65;100;0.04 (34-121)(0.02-0.10) 114;100;0.06 (40-172)(0.03-0.08) 126;100;0.06 (43-165)(0.03-0.19) 107;100;0.06 (43-229)(0.03-0.11) 0;0;0 <1;20;0.01 (0-3)(0-0.08) 0;0;0 0;0;0 0;0;0 <1;10;0.01 (0-1)(0-0.01) 0;0;0 0;0;0 <1;20;<0.01 (0-1) (0-0.01) 1;60;<0.01 (0-4)(0-0.03) 2;90;0.01 (0-8)(0-0.03) 2;80;0.01 (0-4)(0-0.03) 42;100;0.09 (17-67)(0.03-0.30) 49;100;0.05 (8-70)(0.01-0.09) 88;100;0.11 (20-212)(0.03-0.31) 58;100;0.10 (27-118)(0.05-0.15) 13;100;<0.01 (0-33) «0.01-0.02) 9;100;<0.01 (1-31) «0.01-0.01) 38;100;0.02 (8-78) «0.01-0.06) 71;100;0.03 (11-263) (0.01-0.10) 0;0;0 <1;40;<0.01 (0-2) (0-0.01) <1;70;0.01 (0-2) (0-0.02) <1;60;0.04 (0-2)(0-0.13) 2;50;<0.01 (0-10)(0-0.02) 1;80;<0.01 (0-3)(0-<0.01) 4;70;<0.01 (0-17)(0-0.01) 4;100;0.01 (1-10) «0.01-0.03) <1;40;<0.01 (0-2)(0-<0.01) 2;50;<0.01 (0-10)(0-0.01) 5;90;0.01 (0-10)(0-0.05) 11;90;0.01 (0-30) (0-0.02) 206 mean volumes of the i n v e r t e b r a t e taxa from the v a r i o u s f o r e s t f l o o r h a b i t a t s (Tables 29 and 30) us i n g Spearman's c o e f f i c i e n t of rank c o r r e l a t i o n ( r s ) d u r i n g the breeding and winter p e r i o d s (Tables 31 and 32). C o n s i s t e n t l y h i g h or s t a t i s t i c a l l y s i g n i f i c a n t c o r r e l a t i o n s would i n d i c a t e an o p p o r t u n i s t i c and unbiased use of the v a r i o u s i n v e r t e b r a t e taxa of the f o r e s t f l o o r as food by winter wrens. A l t e r n a t i v e l y , c o n s i s t e n t l y low c o r r e l a t i o n s would i n d i c a t e a d i f f e r e n t i a l s e l e c t i o n and use of the i n v e r t e b r a t e taxa. However, the c a l c u l a t e d c o r r e l a t i o n s (Tables 31 and 32) are n e i t h e r s t a t i s t i c a l l y s i g n i f i c a n t or c o n s i s t e n t l y high, nor are they c o n s i s t e n t l y low. As a r e s u l t , I was unable to d i s t i n g u i s h between the a l t e r n a t i v e s of o p p o r t u n i s t i c as opposed to weak s e l e c t i v e use of the v a r i o u s i n v e r t e b r a t e taxa of the f o r e s t f l o o r f o r food by winter wrens. Under these circumstances, I t h e r e f o r e c o n s i d e r e d the t o t a l volumes of i n v e r t e b r a t e s e x t r a c t e d from the i n d i v i d u a l i n v e r t e b r a t e p l o t samples as the best p o s s i b l e , although a d m i t t e d l y crude, index of a v a i l a b l e food f o r winter wrens i n the v a r i o u s f o r e s t f l o o r h a b i t a t s . The means of the t o t a l volumes of i n v e r t e b r a t e s e x t r a c t e d from the i n v e r t e b r a t e p l o t samples from the v a r i o u s f o r e s t f l o o r h a b i t a t s o c c u r r e d i n the f o l l o w i n g 207 TABLE 31 RANKINGS AND STATISTICAL COMPARISONS (SPEARMAN'S COEFFICIENT OF RANK CORRELATION-r ) OF THE MEAN FREQUENCIES (t OCCURRENCES) OF SELECTED INVERTEBRATE TAXA FROM s THE WINTER WREN STOMACHS, WITH THE PRODUCTS OF THE MEAN FREQUENCIES X THE MEAN VOLUMES OF THE INVERTEBRATE TAXA AS EXTRACTED FROM THE INVERTEBRATE PLOT SAMPLES FROM THE VARIOUS FOREST FLOOR HABITATS DURING THE BREEDING PERIOD Invertebrate Taxa a From the Winter Wren Stomachs During the Breeding Period . (Totals A l l P e r i o d s ) 0 M o s s / l i t t e r From the S a l a l ! Extractors Sword fern Salmonl Coleoptera 1(1) 1 1 3 2 Araneida 2(2) 3.5 2 5 3 Lepidoptera 3(3) 13.5 13 IS 13 Hymenoptera 4(4) 5.5 4.5 9.5 12 Diptera 5(5) 9 9 7 5 A c a r i 6(6) 7.5 9 7 9 Hemiptera 7(7) 11.5 12 14 IS Pseudoscorpionda 8(8) 7.5 9 9.5 10.S Diplopoda 9(9.5) 2 9 2 1 Phalangida 10(9.5) 11.5 6 7 10.5 Chilopoda 11.5(12) 3.5 4.5 1 4 Collembola 11.5(11) 5.5 3 4 6 Oligochaeta 14(14) 13.5 14.5 11.5 7.5 Isopoda 14(14) 15 14.5 11.5 7.5 Psocoptera 14(14) 10 9 13 14 r s ( P ) c 0.425 (CO.06) 0.439 K0.06) 0.140 «0.31) 0.138 K0.32) a \ Only the invertebrate taxa which occurred i n winter WTen stomachs at mean frequencies of - 10% and/or occurred i n any f o r e s t f l o o r h a b i t a t at mean frequencies of ^  50% and mean volumes of £0.01 ml were included. D Ranking f o r t o t a l winter wren stomachs i n a l l periods are shown i n brackets, Numbers of stomachs: breeding period - 91; t o t a l a l l periods - 122. C A l l r s values 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 (P } 0.05). 2 0 8 TABLE 32 RANKINGS AND STATISTICAL COMPARISONS (SPEARMAN'S COEFFICIENT OF RANK CORRELATION-r=) OF THE MEAN FREQUENCIES (4 OCCURRENCES) OF SELECTED INVERTEBRATE TAXA FROM THE WINTER WREN STOMACHS, WITH THE PRODUCTS OF THE MEAN FREQUENCIES X THE MEAN VOLUMES OF THE INVERTEBRATE TAXA AS EXTRACTED FROM THE INVERTEBRATE PLOT SAMPLES FROM THE VARIOUS FOREST FLOOR HABITATS DURING THE WINTER PERIOD Invertebrate Taxa a From the Winter Wren Stomachs During the Winter Period , (Totals A l l P e r i o d s ) " M o s s / l i t t e r From the S a l a l Extractors Sword fern Salmonbc-Coleoptera 1(1) 1 3 1 2 Diplopoda 2(9.5) 5 5 3 1 Hemiptera 3(7) 11.5 11 10 14 A c a r i 4(6) 8.5 8 7 10.5 Lepidoptera 5(3) 14 12 12 9 Araneida 7(2) 3 1 3 3 Collembola 7(11) 2 2 3 4 Hymenoptera 7(4) 11.5 10 14 12 Neuroptera 9(12) 14 14.5 15 IS Oligochaeta 12.5(14.5) 7 13 13 5.5 Isopoda 12.5(14.5 14 14.5 11 5.5 Chilopoda 12.5(13) 6 6 5 7 Pseudoscorpionda 12.5(8) 4 4 7 10.5 Phalangida 12.5(9.5) 10 8 9 13 Diptera 12.5(5) 8.5 8 7 8 r s ( P ) c 0.218 «0.22) 0.283 «0.16) 0.250 «0.19) 0.363 «0.10) Only the invertebrate taxa which occurred i n winter wren stomachs at mean frequencies of £ 10% and/or occurred i n any f o r e s t f l o o r h a b i t a t at mean frequencies of i- 504 and mean volumes of i O . O l ml were included. k Ranking f o r t o t a l winter wren stomachs i n a l l periods are shown i n brackets. Numbers of stomachs: winter period - 6; t o t a l a l l periods - 122. c A l l r g values 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 (P > 0.05). 209 d e c r e a s i n g rank i n both the breeding and winter p e r i o d s : salmonberry, sword f e r n , s a l a l , and m o s s / l i t t e r (Table 33). The mean volumes of i n v e r t e b r a t e s from the sword f e r n samples were n e a r l y twice those of the m o s s / l i t t e r and the s a l a l samples while the means f o r the salmonberry samples were n e a r l y twice those of the sword f e r n samples. The i n v e r t e b r a t e volumes from the salmonberry p l o t samples were g r e a t e r (P < 0.05) than the samples from the other f o r e s t f l o o r h a b i t a t s d u r i n g both the breeding and winter p e r i o d s while the i n v e r t e b r a t e volumes from the sword f e r n p l o t samples were g r e a t e r (P < 0.05) than the i n v e r t e b r a t e volumes from the m o s s / l i t t e r and s a l a l samples d u r i n g the breeding p e r i o d . De Catanzaro (1979) working on the study area a l s o found d i f f e r e n c e s i n l i t t e r fauna among the f o r e s t f l o o r h a b i t a t s . The abundance of major groups of i n v e r t e b r a t e s was g r e a t e r on h y g r i c s i t e s (sword f e r n f o r e s t f l o o r h a b i t a t ) than on mesic and x e r i c s i t e s ( m o s s / l i t t e r and s a l a l f o r e s t f l o o r h a b i t a t s r e s p e c t i v e l y ) . My f i n d i n g s and those of de Catanzaro on the r e l a t i v e abundance of i n v e r t e b r a t e s roughly p a r a l l e l the observed d i s t r i b u t i o n s of the areas (m 2) of winter wren t e r r i t o r i e s (Table 19) and the numbers of winter wren o b s e r v a t i o n s (Table 24) i n which the ecosystems grouped t o f o r e s t f l o o r h a b i t a t s were TABLE 33 STATISTICAL COMPARISONS (DUNCAN'S MULTIPLE RANGE TEST) OF THE MEANS OF THE TOTAL VOLUMES (ml) OF INVERTEBRATES FROM THE INVERTEBRATE PLOT SAMPLES OF THE VARIOUS FOREST FLOOR HABITATS DURING THE BREEDING AND WINTER PERIODS Salmonberry Forest F l o o r H a b i t a t s Sword f e r n S a l a l M o s s / L i t t e r Breeding P e r i o d 1.25 ± 0.21* (0.83 - 1.60) 0.63 (0.20 ± 0.21 - 0.95) 0.36 ± 0.13 (0.15 - 0.60) 0.29 ± 0.19 (0.15 - 0.70) w i n t e r 0.80-0.48 0.43 ± 0.22 0.23 ± 0.15 0.20 ± 0.09 P e r i o d (0.30 - 1.85) (0.19 - 0.80) (0.10 - 0.60) (0.08 - 0.40) a ml; X ± SD (range); a l l n = 10 b Means arranged i n decreasing magnitude; means not underscored by the same l i n e are s i g n i f i c a n t l y d i f f e r e n t a t P < 0.05. 21 1 u t i l i z e d i n the f o l l o w i n g order of p r e f e r e n c e (Table 26): s t r o n g p r e f e r e n c e - sword f e r n ; frequent t o no marked p r e f e r e n c e - salmonberry; avoided - s a l a l and m o s s / l i t t e r . I t h e r e f o r e propose t h a t the q u a n t i t y of a v a i l a b l e food was a f a c t o r i n the h a b i t a t s e l e c t i o n and d i s p e r s i o n of winter wrens, although i n t h i s case i t i s not c l e a r whether food i s an u l t i m a t e or proximate f a c t o r , or both. 2 1 2 CONCLUSIONS Male w i n t e r wrens i n t h i s s t u d y were t e r r i t o r i a l on n o n - o v e r l a p p i n g t e r r i t o r i e s ( F i g u r e s 4 t o 13) a t an average d e n s i t y o f about 60 p e r km 2 ( T a b l e 6 ) . Females o c c u p i e d g e n e r a l l y n o n - o v e r l a p p i n g home ranges ( F i g u r e s 14 t o 17), but were not shown t o be t e r r i t o r i a l , a l t h o u g h t h i s p o s s i b i l i t y c o u l d not be e x c l u d e d . I propose t h a t w i n t e r wrens a r e spaced by t e r r i t o r i a l i t y ( Q u e s t i o n 1, p . 2 ) . W i n t e r wrens were c o n s i s t e n t l y b u t n o t u n i f o r m l y d i s t r i b u t e d o v e r t h e s t u d y a r e a ( F i g u r e s 4 t o 13, and 14 t o 17) ( Q u e s t i o n 2 ) . S u r p l u s , p o t e n t i a l l y t e r r i t o r i a l males were a v a i l a b l e d u r i n g t h e b r e e d i n g p e r i o d , w h i c h c o u l d have o c c u p i e d t h e empty o r s p a r s e l y o c c u p i e d a r e a s ( F i g u r e 13; T a b l e 7) ( Q u e s t i o n 3 ) . Wi n t e r wrens showed d i f f e r e n t i a l use of the v a r i o u s i n d i v i d u a l ecosystems a t the l e v e l of s u b - a s s o c i a t i o n and ecosystems grouped by the dominant f o r e s t f l o o r v e g e t a t i o n ( T a b l e s 16 t o 18, and 21 t o 2 3 ) . W i n t e r wrens p r e f e r r e d , b o t h as i n d i v i d u a l ecosystems and as grouped ecosystems, t h e ecosystems grouped t o the sword f e r n f o r e s t f l o o r h a b i t a t . I n t h e same way, w i n t e r wrens showed f r e q u e n t t o no marked p r e f e r e n c e f o r the ecosystems grouped t o t h e salmo n b e r r y f o r e s t f l o o r h a b i t a t , but a v o i d e d t h e 213 ecosystems grouped to the s a l a l and m o s s / l i t t e r f o r e s t f l o o r h a b i t a t s (Tables 19 and 20, and 24 to 26). I propose that winter wrens are clumped by s u i t a b l e h a b i t a t (Question 4 ) . D i f f e r e n c e s i n cover and s p e c i e s composition of the f o r e s t f l o o r v e g e t a t i o n (Tables 9, 13, and 26) are proposed as p o s s i b l e f a c t o r s i n f l u e n c i n g winter wren d i s p e r s i o n . I suggest that these f a c t o r s r e l a t e to requirements of winter wrens f o r s h e l t e r from adverse weather and enemies, and to the f u n c t i o n a l and s t r u c t u r a l c h a r a c t e r i s t i c s of the b i r d s . The t o t a l volumes of i n v e r t e b r a t e s a v a i l a b l e to winter wrens were g r e a t e r i n f o r e s t f l o o r h a b i t a t s p r e f e r r e d by winter wrens (Tables 26 and 33). Food i s a l s o proposed as a f a c t o r i n winter wren d i s p e r s i o n (Question 5 ). I f winter wrens are spaced by t e r r i t o r i a l i t y and clumped by s u i t a b l e h a b i t a t , then I suggest that these two f a c t o r s i n f l u e n c e the p a t t e r n s of d i s p e r s i o n of winter wrens i n the c o a s t a l western hemlock f o r e s t of the U n i v e r s i t y of B r i t i s h Columbia Research F o r e s t i n south-western B r i t i s h Columbia, and perhaps elsewhere as w e l l . E c o l o g i c a l ( b i o p h y s i c a l ) approaches to f o r e s t land c l a s s i f i c a t i o n , i n which both b i o l o g i c a l and p h y s i c a l f a c t o r s are c o n s i d e r e d , are w e l l e s t a b l i s h e d i n North 214 America (e.g., K r a j i n a 1960, 1972, 1978; Mueller-Dombois 1965; Thomas et a l . 1979; W i l k i n 1980; Leak 1982). E c o l o g i c a l c l a s s i f i c a t i o n and mapping have been used i n the study of w i l d l i f e (e.g., Luckhurst et a l . 1973; Luckhurst 1975; V o i d 1977; Resource A n a l y s i s Branch 1979a, b; Thomas et a l . 1979; V o i d et a l . 1980; Ho l r o y d 1980; P r e s c o t t 1980). However, most work done thus f a r on w i l d l i f e has been at the reconnaissance l e v e l (1:50,000 s c a l e and s m a l l e r ) . W i l d l i f e has not been s t u d i e d at the l e v e l of ecosystem s u b - a s s o c i a t i o n and a s s o c i a t i o n , which i n most landscapes can only be shown at l a r g e s c a l e mapping (1:10,000 s c a l e and l a r g e r ) , and which f o r the most p a r t remains as yet undefined and unmapped. In the Province of B r i t i s h Columbia, the mapping of the e n t i r e p r o v i n c e to the l e v e l of b i o g e o c l i m a t i c subzone and v a r i a n t (1:500,000) s c a l e i s i n p r o g r e s s (e.g., Kimmins 1979; K l i n k a et a l . 1979). Ecosystem s u b - a s s o c i a t i o n s and a s s o c i a t i o n s i n s e l e c t e d areas throughout the p r o v i n c e are being d e s c r i b e d and mapped ( K l i n k a 1976; K l i n k a et a l . 1980; M i t c h e l l and Green 1981; I n s e l b e r g et a l . 1982). The B r i t i s h Columbia M i n i s t r y of F o r e s t s has adopted the ecosystem s p e c i f i c approach to the s i l v i c u l t u r a l management of i t s f o r e s t l a n d base (Schmidt 1978; Kimmins 1979). 215 In t h i s study, I used K l i n k a ' s (1976) pioneer s y n e c o l o g i c a l d e s c r i p t i o n and mapping of the Research F o r e s t i n a study of winter wren d i s p e r s i o n . The f i n d i n g s of t h i s study suggest that K l i n k a ecosystems, and ecosystems grouped by f o r e s t f l o o r v e g e t a t i o n of c o a s t a l western hemlock (dry subzone) are i n d i c a t i v e of the d i s t r i b u t i o n of winter wrens, although f u r t h e r work w i l l be r e q u i r e d to t e s t t h i s h y p o t h e s i s f u l l y . Should t h i s h y p o t h e s i s be c o r r e c t , then ecosystems and ecosystems grouped by a p p r o p r i a t e c r i t e r i a may be i n d i c a t i v e of the d i s t r i b u t i o n s of other s p e c i e s of w i l d l i f e as w e l l . T h i s view i s supported g e n e r a l l y f o r b i r d s by S c o u l l a r ' s (1980) f i n d i n g s t h a t the d i s t r i b u t i o n s of s i x of the ten b i r d s p e c i e s he s t u d i e d were r e l a t e d to v e g e t a t i o n types (p. 179). However, he a l s o found s u c c e s s i o n (not s t u d i e d here) to be more important than v e g e t a t i o n type i n d etermining the d i s t r i b u t i o n of most b i r d s p e c i e s he s t u d i e d . Should the p r o v i n c e of B r i t i s h Columbia p e r s i s t i n the s i l v i c u l t u r a l management of i t s f o r e s t lands on the b a s i s of ecosystems, and should ecosystems, i n c l u d i n g a c o n s i d e r a t i o n of s u c c e s s i o n , prove i n d i c a t i v e of the d i s t r i b u t i o n of w i l d l i f e s p e c i e s g e n e r a l l y , then ecosystems can and should a l s o be used as the base f o r the 216 study and management of w i l d l i f e i n the p r o v i n c e , and perhaps elsewhere as w e l l . 2 1 7 REFERENCES 218 REFERENCES Anderson, S.H.; Shugart, H.H., J r . H a b i t a t s e l e c t i o n of bree d i n g b i r d s i n an east Tennessee deciduous f o r e s t . Ecology 55: 828-837; 1974. Armstrong, E.A. Some notes on the song of the wren. B r i t i s h B i r d s 38: 70-72; 1944. Armstrong, E.A. The behaviour and breeding b i o l o g y of the I c e l a n d wren. I b i s 92: 384-401; 1950. Armstrong, E.A. 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Zoologicheskii Zhurnal 51: 770-772; 1972. 253 A P P E N D I C E S APPENDIX 1 SYNOPSIS OF THE SYNECOLOGICAL CLASSIFICATION OF THE COASTAL WESTERN HEMLOCK FOREST OF THE UNIVERSITY OF BRITISH COLUMBIA RESEARCH FOREST ( s o i l parent m a t e r i a l components omitted) This appendix i s summarized from Table 8 o f K l i n k a ' s d i s s e r t a t i o n ( K l i n k a 1976) and Mapsheet 3 o f the Ecosystems U n i t s Maps o f the U.B.C. Research Forest ( U n i v e r s i t y o f B r i t i s h Columbia Research Forest 1976). The c l a s s i f i c a t i o n system has been m o d i f i e d t o in c l u d e recent changes (K. K l i n k a , p e r s o n a l communi-ca t i o n ) . 255 Appendix 1 Order 1 ALLIANCE ECOSYSTEM ASSOCIATION Ecosystem P s e u d o t s u g e t a l i a m e n z i e s i i , I GAULTHERIA - DOUGLAS-FIR, 1 a * 2 ' 3 (LICHEN) -GAULTHERIA - D F 4 , 11 b LICHEN - GAULTHERIA - LP - DF , 12 a $ b GAULTHERIA - DF, 13 a* G a u l t h e r i a - WH - DF, 131 a § b* Mahonia - G a u l t h e r i a - WH - DF, 132 T s u g e t a l i a h e t e r o p h y l l a e , I I MOSS - WESTERN HEMLOCK, 2 a § b MOSS - WH, 21 a* Moss - WH, 211 a § b* Mahonia - Moss - WRC - WH, 212 MOSS - (POLYSTICHUM) - WESTERN HEMLOCK, 3 a § b* MOSS - (POLYSTICHUM) - WRC - WH, 31 VACCINIUM - WESTERN HEMLOCK, 4 b VACCINIUM - GAULTHERIA - DF - WH, 41 b VACCINIUM - MOSS - WH, 42 BLECHNUM - AMABILIS FIR - WESTERN HEMLOCK, 5 b BLECHNUM - AF - WH, 51 Appendix 1 (continued) b BLECHNUM - STREPTOPUS - AF - WH, 52 b BLECHNUM - WH - WRC, 53 Thujetalia plicatae, III MOSS - POLYSTICHUM - DOUGLAS-FIR - WESTERN REDCEDAR, b RIBES - VM, 61 a $ b* POLYPODIUM - GAULTHERIA - DF - WRC, 62 a & b* POLYPODIUM - POLYSTICHUM - DF - WRC, 63 a § b MAHONIA - POLYSTICHUM - DF - WRC, 64 TIARELLA - POLYSTICHUM - WESTERN REDCEDAR, 7 a 5 b* TIARELLA - POLYSTICHUM - WRC, 71 a § b* RUBUS - POLYSTICHUM - WRC, 72 a § b ADIANTUM - POLYSTICHUM - WRC, 73 OPLOPANAX - WESTERN REDCEDAR, 8 OPLOPANAX - WRC, 81 a § b Polystichum - Oplopanax - WRC, 811 a § b* Ribes - Oplopanax - WRC, 812 LYSICHITUM - WESTERN REDCEDAR, 9 VACCINIUM - LYSICHITUM - WRC, 91 a § b* Vaccinium - Lysichitum - WRC, 911 b Vaccinium - Lysichitum - YC - WRC, 912 Populetalia balsamiferae, IV RED ALDER - SITKA AIDER, 10 b ATHYRIUM - ARUNCUS - RA - SA, 257 Appendix 1 (continued) *See Appendix 2 for an explanation of the conventions used in writing the synecological units. 2 Coastal western hemlock biogeoclimatic zone: a - dry sub-zone; b - wet sub-zone. 3* - present on the study areas (ecosystems only). 4 See Appendix 2 for an explanation of the abbreviations. 258 APPENDIX 2 SUMMARY OF THE KLINKA SYNECOLOGICAL UNITS OF THE STUDY AREA, THEIR CLASSIFICATION, PARENT MATERIALS, NUTRIENT AND MOISTURE REGIMES, AND ASSOCIATED SOILS ~ This appendix like Appendix 1 is summarized from Table 8 of Klinka's dissertation (Klinka 1976) and Mapsheet 3 of the Ecosystems Units Maps of the U.B.C. Research Forest (University of British Columbia Research Forest 1976). The classification system has been modified to include recent changes (K. Klinka, personal communication). APPENDIX 2 Classification Soil Parent Materials; Soil Nutrient Regimes; Soil Moisture Regimes Order ALLIANCE ECOSYSTEM ASSOCIATION (at the level of plant association) Ecosystem (at the level of plant community) Pseudotsugetalia menziesii, i*»2»3>4 GAULTHERIA - DOUGLAS-FIR, 1 (LICHEN) - GAULTHERIA - DF, 11 (LICHEN) - GAULTHERIA - DF, 11.30 Veneer (moraine materials), organic deposits; Submesotrophic; Very xeric. GAULTHERIA - DF, 13 Gaultheria - WH - DF, 131.30 Gaultheria - WH - DF, 131.23 Gaultheria - WH - DF, 131.4 Veneer (moraine materials), organic deposits; Submesotrophic; Xeric. Blanket, veneer (moraine materials); Submeso-trophic; Subxeric. Glaciofluvial deposits; Mesotrophic; Xeric Mahonia - Gaultheria - WH-DF, 132.9 Veneer (colluvial materials); Subeutrophic; Xeric. Tsugetalia heterophyllae, II MOSS - WESTERN HEMLOCK, 2 MOSS - WH, 21 Moss - WH, 211.23 Moss - WH, 211.4 Mahonia - Moss - WRC - WH, 212.89 Blanket, veneer (moraine materials); Submeso-trophic; Submesic. Glaciofluvial deposits; Mesotrophic; Submesic. Blanket, veneer (colluvial materials); Sub-eutrophic; Submesic Associated Soils at the Level of So i l Series Loamy sand Lithic Mini Humo-Ferric and Lithic Podzols with mor humus; Lithic F o l i s o l and Proto-ranker with mor humus. Loamy sand Li t h i c Mini Humo-Ferric and Lit h i c Podzols with mor humus; Lithic F o l i s o l with mor humus. Sandy loam Mini Humo-Ferric Podzol with mor humus. Sandy loam Mini and Orthic Humo-Ferric Podzols with mor humus. Sandy loam Mini and Orthic Humo-Ferric Podzols with mor humus. Loamy sand Mini Humo-Ferric Podzol with mor humus. Sandy loam Mini and Orthic Humo-Ferric Podzols with mor humus; loamy sand Sombric Humo-Ferric Podzol with moder humus. Appendix 2 (continued) Classification Soil Parent Materials; Soil Nutrient Regimes; Soil Moisture Regimes Associated Soils at the Level of S o i l Series Tsugetalia heterophyllae, II (continued) MOSS - (POLYSTICHUM) - WESTERN HEMLOCK, 3 MOSS - (POLYSTICHUM) - WH - WRC, 31 MOSS - (POLYSTICHUM) - WH - WRC, 31.12 Deep deposits, blanket (moraine materials); Subeutrophic; Subhygric. MOSS - (POLYSTICHUM) - WH - WRC, 31.412 Glaciofluvial deposits, deep deposits and blanket (moraine materials); Subeutrophic; Subhygric. Sandy loam Orthic Humo-Ferric Podzol with mor humus. Loamy sand (Gleyed) Mini and Orthic Humo-Ferric Podzols with moder humus. Thujetalia plicatae, III MOSS - POLYSTICHUM - DOUGLAS-FIR - WESTERN REDCEDAR, 6 POLYPODIUM - GAULTHERIA - DF - WRC, 62 POLYPODIUM - GAULTHERIA - DF-WRC, 62.90 Veneer (colluvial materials), organic deposits; Subeutrophic; Subhygric. POLYPODIUM - POLYSTICHUM-DF-WRC, 63 POLYPODIUM - POLYSTICHUM - DF-WRC, 63.89 Blanket, veneer (colluvial materials); Sub-eutrophic; Subhygric. TIARELLA - POLYSTICHUM - WESTERN REDCEDAR, 7 TIARELLA - POLYSTICHUM - WRC, 71 Deep deposits, blanket (moraine materials); Subeutrophic; Hygric. Glaciofluvial deposits; Subeutrophic; Hygric. Glaciofluvial deposits; glaciomoraine deposits; Subeutrophic; Hygric. TIARELLA - POLYSTICHUM - WRC, 71.4 TIARELLA - POLYSTICHUM - WRC, 71.47 Lit h i c F o l i s o l and Protoranker with mor and moder humus; sandy loam Lithic Podzol with mor and moder humus. Loamy sand and sandy loam skeletal Mini and Orthic Humo-Ferric and Mini and Sombric Ferro-Humic Podzols with moder humus. Loamy (Gleyed) Mini Humo-Ferric and Ferro-Humic Podzols with moder humus. Sandy loam (Gleyed) Mini and Sombric Humo-Ferric and Ferro-Humic Podzols with moder humus. Appendix 2 (continued) Classification Thujetalia plicatae, III (continued) RUBUS - POLYSTICHUM - WRC, 72 RUBUS - POLYSTICHUM - WRC, 72.4 RUBUS - POLYSTICHUM - WRC, 72.47 OPLOPANAX - WESTERN REDCEDAR, 8 OPLOPANAX - WRC, 81 Ribes - Oplopanax - WRC, 812.89 LYSICHITUM - WESTERN REDCEDAR, 9 VACCINIUM - LYSICHITUM - WRC, 91 Vaccinium - Lysichitum - WRC, 911.0 Other Map Units R (Rock) C (Clearcut) Soil Parent Materials; Soil Nutrient Regimes; Soil Moisture Regimes Associated Soils at the Level of S o i l Series Glaciofluvial deposits; eutrophic; Hygric. Glaciofluvial deposits, glaciomoraine deposits; Eutrophic; Hygric. Sandy loam Gleyed Mini Humo-Ferric Podzol with moder humus; sandy loam Gleyed Mini Humo-Ferric Podzol with mull humus; loamy sand Orthic and Cumulic Regosol with mull and hydromull humus; sandy Orthic Sombric Brunisol with mull humus; sandy loam Gleyed Sombric Humo Ferric Podzol with mull humus; Terric Humisol with hydromull humus; s i l t y clay Orthic Humic Gleysol with, hydromull humus; clay loam Gleyed Sombric Humo-Ferric Podzol with mull humus. Blanket, veneer (colluvial materials); Sandy loam (Gleyed) Sombric and Mini Humo-Ferric Eutrophic; Hygric. Podzol with moder humus; loamy and s i l t y clay Gleyed Sombric and Mini Humo-Ferric Podzols with mull humus. Organic deposits; Subeutrophic; Subhydric. Sandy loam Orthic Humic Gleysol with hydromoder humus; Terric and Typic Humisols with hydromoder humus. Non-forested ecosystems on rocks Area clearcut in the late winter of 1977 to CTl Appendix 2 (continued) * The conventions for writing Klinka's four synecologic taxa are as follows: Order - Roman numerals; f i r s t letter of the f i r s t word capitalized. ALLIANCE - Arabic numerals; f i r s t integer of the map unit number; a l l capital letters; tree species spelled out. ECOSYSTEM ASSOCIATION - Arabic numerals; second integer of the map unit number; a l l capital l e t t e r s ; tree species abbreviated; at the level of plant association. Ecosystem - Arabic numerals; third integer of the map unit number; f i r s t letters of a l l words capitalized; tree species abbreviated i n capitals; at the level of plant community. ECOSYSTEM ASSOCIATIONS which have subordinate ecosystems at the plant community level do not have s o i l parent material modifiers and are not used as map units. Key to the numerical map symbols: Vegetation Soil Parent Material PLANT ALLIANCE PLANT ASSOCIATION Plant Community 131.4 41 L Genetic category of unconsolidated deposits So i l parent materials: R - bedrock .1 - Deep deposits) .2 - Blanket ) materials .3 - Veneer ) .4 - Glaciofluvial deposits .5 - A l l u v i a l deposits .6 - Glaciolacustrine .7 - Glaciomarine .8 - Blanket .9 - Veneer 0 - Organic Colluvial materials Tree species abbreviations: AF - amabilis f i r DF - Douglas f i r LP - lodgepole pine RA - red alder SA - Sitka alder VM - vine maple WH - western hemlock WRC - western redcedar YC - yellow cedar 263 APPENDIX 3 LIST OF PLANT SPECIES FOUND ON THE VEGETATION AND FOREST FLOOR SAMPLE PLOTS Vascular and non-vascular plant species are listed separately. The vascular plant species are arranged alphabetically by common name while the non-vascular plant species are listed by scientific name. The nomenclature of the vascular plants follows Taylor and MacBryde (1977) while that of the non-vascular plants generally follows Appendix X of Klinka's dissertation (1976). The authorities given by Klinka for bryophytes were Crum, Steere and Anderson (1973), Frye and Clark (1937-1947), Lawton (1971), Nyholm (1969), Schofield (1968a, b) and Schuster (1966-1974) while the authorities given for lichens were Hale and Culberson (1960) and Otto and Ahti (1967). Dr. W.B. Schofield, Professor of Botany, University of British Columbia, kindly identified some of the more difficult bryophytes. Ms. C. Jones identified some of the grasses, sedges and composites. A more complete listing of the plant species of the University of British Columbia Research Forest was given in Klinka (1976). 264 Appendix 3 Vascular Plant Species Alpine enchanter's-nightshade American red raspberry American skunk-cabbage American speedwell American vanilla leaf Baldhip rose Bigleaf maple Black cottonwood Black raspberry Broad-leaved starflower Canadian bunchberry Coastal American red elder Coast Douglas f i r Common cattail Common lady fern Common paper birch Common pearly everlasting Cucumberroot twistedstalk Circaea alpina Linnaeus subsp. alpina Rubus idaeus Linnaeus subsp. melanolasius (Dieck) Focke Lysichiton americanum Hulten and St. John Veronica americana ex Bentham in A.P. de Candolle Achlys triphylla (J.E. Smith) A.P. de Candolle subsp. triphylla Rosa gymnocarpa Nuttall in Torrey and Gray Acer macrophyllum Pursh Populus balsamifera Linnaeus subsp. trichocarpa (Torrey and Gray ex W.J. Hooker Brayshaw Rubus leucodermis D. Douglas ex Torrey and Gray var. leucodermis Trientalis latifolia W.J. Hooker Cornus canadensis Linnaeus Sambucus racemosa Linnaeus subsp. pubens (A. Machaux) House var. arborescens (Torrey and Gray) A- Gray Pseudosuga menziesii (Mirbel) Franco var. menziesii Typha latifolia Linnaeus Athyrium felix-femina (Linnaeus) Roth subsp. cyclosorum (Ruprecht) Christensen in Hulten Betula papyrifera Marshal var. papyrifera Anaphalis margaritacea (Linnaeus) Bentham and Hooker Streptopus amplexifolius (Linnaeus) A.P. de Candolle in Lamark and Candolle var. americanus J.A. Schulter 2 6 5 Appendix 3 (continued) Vascular Plant Species (continued) Deer fern Devil's-club Douglas' spirea Dull Oregon-grape English holly Fireweed Five-leaved creeping raspberry Fowl blue grass Fringed pinesap Green speenwort High bush cranberry Indian-pipe Indian-plum Kentucky blue grass Large-leaved rattlesnake orchid Licorice fern Northern maidenhair fern Northern twinflower Oval-leaved blueberry Pacific bleeding heart Blechnum spicant (Linnaeus) Roth Oplopanax horridus (J.E. Smith) Miquel Spiraea douglasii W.J. Hooker subsp. douglasii Mahonia nervosa (Pursh) Nuttal Ilex aquifolium Linnaeus Epilobium angustifolium Linnaeus subsp. circumvagum Mosquin Rubus pedatus J.E. Smith Poa palustris Linnaeus Hypopitys monotropa Crantz Asplenium viride Hudson Viburnum edule (A. Michaux) Rafinesque Monotropa uniflora Linnaeus Oemleria cerasiformis (Hooker and Arnoth) Landon Poa pratensis Linnaeus subsp. pratensis  Goodyera oblongifolia Rafinesque Polypodium glycyrrhiza D.C. Eaton Adianturn pedatum Linnaeus subsp. aleuticum (Ruprecht) Calder and Taylor Linnaea borealis Linnaeus subsp. longiflora (Torrey) Hulten Vaccinium ovalifolium J.E. Smith in Rees Dicentra formosa (Haworth) Walpers subsp. formosa Pacific crab apple Malus fusca (Rafinesque) C.K. Schneider Appendix 3 (continued) Vascular Plant Species (continued) Pacific oenanthe Pacific Saskatoon Pacific trailing blackberry Piggy-back plant Red alder Red baneberry Red huckleberry Rusty Pacific menziesia Salal Salmonberry Sheep sorrell Siberian spring beauty Sitka mountain-ash Sitka spruce Small-flowered wood rush Spear thistle Spike bent grass Shiny shield fern Sticky chickweed Oenanthe sarmentosa K.B. Presl ex A.P. de Candolie Amelanchier alnifolia (Nuttall) Nuttall var. semimtegrifolia (W.J. Hooker) C.L. Hitchcock in Hitchcock et al Rubus ursinus Chamisso and Schlechtendal subsp. macropetalus (D. Douglas ex W.J. Hooker) Taylor and MacBryde Tolmiea menziesii (Pursh) Torrey and Gray Alnus rubra Bongard Actaea rubra (W. Acton) Willdenow subsp. arguta (Nuttall ex Torrey and Gray) Hulten Vaccinium parviflorum J.E. Smith in Rees Menziesia ferruginea J.E. Smith subsp. feriaiginea Gaultheria shallon Pursh Rubus spectabilis Pursh Rumex acetosella Linnaeus subsp. acetosella Claytonia siberica Linnaeus var. siberica Sorbus sitchensis M.J. Roemer subsp. sitchensis Picea sitchensis (Bongard) Carriere Lazula parviflora (Ehrhart) N.A. Desvaux subsq. fastigiata (E.H.F. Meyer) Hamet-Ahti Cirsium vulgare (E. Sari) Tenore Agrostis exarata Trinius subsp. exarata var. exarata Dryopteris assimilis S. Walker Cerastium glomeratum Thuillier 267 Appendix 3 (continued) Vascular Plant Species (continued) Sticky willowherb Stiff club-moss Stink currant Sweet-scented bedstraw Sword-leaved rush Sylvan goat's-beard Trailing evergreen yellow violet Trifoliate-leaved foamflower Vine maple Wall-lettuce Western bracken Western fescue Western flowering dogwood Western hemlock Western red cedar Western sword fern Western thimbleberry Western white trillium White clover Wood strawberry Epilobium glandulosum Lehmann Lycopodium annotinum Linnaeus subsp. annotinum Ribes bracteosum D. Douglas ex W.J. Hooker Galium triflorum A. Michaux Juncus ensifolius Wikstrom var. ensifolius Aruncus dioicus Walter Fernald Viola sempervirens Green Tiarella trifoliata Linnaeus Acer circinatum Pursh Mycelis muralis (Linnaeus) Dumortier Pteridium aquilinum (Linnaeus) Kuhn in Decken subsp. aquilinum var. pubescens L.M. Underwood Festuca occidentalis W.J. Hooker Cornus nuttallii Audubon ex Torrey and Gray Tsuga heterophylla (Rafinesque) Sargent Thuja plicata Don ex D. Don in Lambert Polystichum munitum (Kaulfuss) K.B. Presl f. rounitum Rubus parviflorus Nuttall subsp. parviflorus Trillium ovatum Pursh f. ovatum Trifolium repens Linnaeus Fragaria vesca Linnaeus subsp. bracteata (A.A. Heller) Staudt 268 Appendix 3 (continued) Vascular Plant Species (continued) Wool-grass bulrush Scirpus cyperinus (Linnaeus) Kunth var. brachypodus (Fernald) Gilly Yorkshire fog Holcus lanatus Linnaeus Appendix 3 (continued) Non-vascular Plant Species Bazzania denudata (Torr.) Trevis Calypogeia muelleriana (Schiffn.) C. Muell. Calypogeia trichomanis (L.) Corda Cladina rangiferina (L.) Harm. Claopodium crispifolium (Hook.) Ren. and Card. Conocephalum conicum (L.) Dum. Dendroalsia abietina (Hook.) Britt. Dicranella palustris (Dicks.) Crundw. ex Warb. Dicranoweisia cirrata (Hedw.) Lindb. ex Milde Dicranum fuseescens Turn. Dicranum howellii Ren. and Card. Ditrichum heteromallum (Hedw.) Britt. Heterocladium macounii Best Hylocomium splendens (Hedw.) B.S.G. Hypnum circinale Hook. Hypnum dieckii Ren. and Card, ex Roell Isopterygium elegans (Brid.) Lindb. Isothecium stoloniferum (Hook) Brid. Lepidozia reptans (L.) Dum. Leucolepis menziesii (Hook.) Steere ex L. Koch Metzgeria conjugata Lindb. Neckera douglasii Hook. Neckera menziesii Hook. Pellia neesiana (Gottsche) Limpr. Appendix 3 (continued) Non-vascular Plant Species (continued) Plagiochila asplenioides (L.) Dim. Plagiomnium insigne (Mitt.) Koponen Plagiomnium insigne (Mitt.) Koponen Plagiothecium undulatum (Hedw.) B.S.G. Pleurozium schreberi (Brid.) Mitt. Pogonotum contortum (Brid.) Lesq. Porella navicularis (Lehm. and Lindenb.) Lindb. Polytrichum juniperinum Hedw. Porella r o e l l i i Steph. Rhacomitrium aciculare (Hedw.) Brid. Rhacomitrium canescens (Hedw.) Brid. Rhacomitrium lanuginosum (Hedw.) Brid. Rhizomnium glabrescens (Kindb.) Koponen Rhytidiadelphus lbreus (Hedw.) Warnst. Scapania americana C. Muell. Scapania bolanderi Aust. Sphagnum palustre L. Stokesiella oregana (Sull.) Robins. Tetraphis pe Hue Ida Hedw. 271 APPENDIX 4 LIST OF INVERTEBRATE TAXA IDENTIFIED FROM THE INVERTEBRATE SAMPLE PLOTS AND THE WINTER WREN STOMACHES The invertebrates from the Berlese funnels and the winter wren stomaches were identified to family where possible although some were identified only to order or class. The general references used in the identifications were Chu (1949), Jaques (1951), Borror et al (1976), Bland and Jaques (1978), Kaston (1978), Danks (1979), and McDaniel (1979). Mr. S.G. Cannings, of the Spenser Entomological Museum, University of British Columbia helped in many insect identifications while Dr. W. Lazorko identified many of the minute beetles including some to the level of genus. Dr. A.L. Turnbull, Department of Biology, Simon Fraser University helped in identifying many of the more difficult spiders. APPENDIX 4 INVERTEBRATE TAXA (*-found in winter wren stomaches) Phylum Annelida Class Oligochaeta Order Prosopora Family Lumbriculidae (incl. Lumbricus sp.) Phy. Arthropoda* CI. Crustacea* Ord. Isopoda CI. Diplopoda* Ord. Polyxenida Polydesmida Chordeumida* Julida* Platydesmida Unid. Diplopoda* CI. Chilopoda* Ord. Geophilomorpha Lithob iomorpha Unid. Chilopoda* CI. Symphyla CI. Arachnida* Ord. Pseudoscorpionida* Phalangida* Acari* Araneida* Fam. Amaurobiidae Theridiidae Linyphiidae Micryphant idae Agelenidae* Clubionidae Thomisidae Salticidae* Unid. Araneida* [Sow bugs] [Millipedes] [Centipedes] [Symphylans] [Pseudoscorpions] [Harvestmen] [Mites and Ticks] [Spiders] Appendix 4 (continued) CI. Insecta* Ord. Collembola* Fam. Hypogastruridae* Onychiuridae Isotomidae Entomobryidae Sminthuridae* Ord. Thysanura* Fam. Machilidae* Ord. Orthoptera Fam. Tetrigidae Raphidiophoridae Ord. Psocoptera Fam. Polypsocidae Ord. Hemiptera* Subord. Heteroptera* Fam. Pentatomidae* Lygaeidae Tingidae* Nabidae Miridae Dipsocoridae Subord. Homoptera* Fam. Delphacidae Cixiidae Aphididae Unid. Hemiptera* Ord. Thysanoptera Fam. Phlaeothripidae Ord. Neuroptera* Fam. Hemerobiidae Unid. Neuroptera* [Springtails] [Bristletails] [Grasshoppers and Crickets] [Psocids] [True Bugs, Aphids, others] [Thrips] [Lacewings and others] 274 Appendix 4 (continued) Ord. Coleoptera* [Beetles] Fam. Carabidae* (incl. Clivina sp.) Dytiscidae Hydrophilidae (Sphaeridium sp.) Ptiliidae Micropeplidae (incl. Micropeplus sp.) Leptinidae Leiodidae (incl. Empelus brunhipennis, Agathidium sp., Catops sp.) Scydmaenidae (incl. Lophioderis sp.) Silphidae (incl. Agyrtes sp.) Scaphidiidae Staphylinidae* (incl. Omaliinae sp., Boletobiini sp.) Pselaphidae Helodidae (incl. Helodes sp.) Byrrhidae* (incl. Lioligus sp., Lioon sp.) Elateridae* Throscidae Lampyridae Cantharidae* Bostrichidae Nitidulidae (incl. Epuraea sp.) Cryptophagidae (incl. Henoticus sp., Atomaria sp. Phalacridae* Coccinellidae* Lathridiidae (incl. Corticarina sp., Eniemus sp.) Mordellidae (incl. Anaspis sp.) Meloidae Cerambycidae* Chrysomelidae Brentidae Scolytidae Curculionidae* Unid. Coleoptera* Ord. Diptera* [Flies] Subord. Nematocera* Fam. Trichoceridae Tipulidae Chaoboridae Culicidae Ceratopogonidae Chironomidae Mycetophilidae Sciaridae Cecidomyidae 275 Appendix 4 (continued) Subord. Brachycera* Fam. Stratiomyidae Rhagionidae Tabanidae Phoridae* Syrphidae Lauxaniidae Heleomyzidae Drosophilidae Anthomyiidae Museidae Tachinidae Hippoboscidae Unid. Diptera* Ord. S iphonaptera Fam. Pulicidae Ord. Lepidoptera* Fam. Tortricidae Pterophoridae Geometridae Noctuidae* Unid. Lepidoptera* Ord. Trichoptera Fam. Polycentropodidae Ord. Hymenoptera* Subord. Apocrita* Fam. Braconidae Ichneumonidae* Aphelinidae Aulacidae Scelionidae* Platygastridae Eurytomidae Formicidae* Unid. Hymenoptera* Phy. Mollusca CI. Gastropoda [Fleas] [Moths and Butterflies] [Caddisflies] [Ants, Bees, Wasps and allies] [Snails and Slugs] 

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