Open Collections

UBC Theses and Dissertations

UBC Theses Logo

UBC Theses and Dissertations

Vegetation - environment relationships of forest communities on central eastern Vancouver Island Beese, William John 1981

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

Item Metadata

Download

Media
831-UBC_1981_A6 B44.pdf [ 21.98MB ]
Metadata
JSON: 831-1.0095194.json
JSON-LD: 831-1.0095194-ld.json
RDF/XML (Pretty): 831-1.0095194-rdf.xml
RDF/JSON: 831-1.0095194-rdf.json
Turtle: 831-1.0095194-turtle.txt
N-Triples: 831-1.0095194-rdf-ntriples.txt
Original Record: 831-1.0095194-source.json
Full Text
831-1.0095194-fulltext.txt
Citation
831-1.0095194.ris

Full Text

VEGETATION - ENVIRONMENT RELATIONSHIPS OF FOREST COMMUNITIES ON CENTRAL EASTERN VANCOUVER ISLAND by William John Beese B.S., Southern I l l i n o i s University, 1978 THESIS SUBMITTED IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF FORESTRY in THE FACULTY OF GRADUATE STUDIES (Faculty of Forestry) We accept this thesis as conforming to the required standard THE UNIVERSITY OF BRITISH COLUMBIA May, 1981 ©William John Beese, 1981 In p r e s e n t i n g t h i s t h e s i s i n p a r t i a l f u l f i l m e n t o f t h e r e q u i r e m e n t s f o r an a d v a n c e d d e g r e e a t t h e U n i v e r s i t y o f B r i t i s h C o l u m b i a , I a g r e e t h a t t h e L i b r a r y s h a l l make i t f r e e l y a v a i l a b l e f o r r e f e r e n c e and s t u d y . I f u r t h e r a g r e e t h a t p e r m i s s i o n f o r e x t e n s i v e c o p y i n g o f t h i s t h e s i s f o r s c h o l a r l y p u r p o s e s may be g r a n t e d by t h e Head o f my D e p a r t m e n t o r by h i s r e p r e s e n t a t i v e s . I t i s u n d e r s t o o d t h a t c o p y i n g o r p u b l i c a t i o n o f t h i s t h e s i s f o r f i n a n c i a l g a i n s h a l l n o t be a l l o w e d w i t h o u t my w r i t t e n p e r m i s s i o n . D e p a r t m e n t o f F o r e s t r y  The U n i v e r s i t y o f B r i t i s h C o l u m b i a 2075 Wesbrook P l a c e V a n c o u v e r , C a n a d a V6T 1W5 D a t e A p r i l 9, 1981 D E - 6 B P 7 5 - 5 1 1 E i "OveA the. changing Zand AuA^aceA of, the Eauth, tkzn.iL ib> a mantle ofa vegetation, a Living tapestsiy o{j plant communities, that JJ> subtly beautiful, inteLtectually challenging, and economically uubz^ut." Flank E. EgleA, 7977 To my wife Beverley, whose constant help, encouragement and patience throughout t h i s project w i l l always be remembered. and To Dr. Ph i l Robertson, Southern I l l i n o i s University, Department of Botany, whose enthusiasm for plant ecology and excellent teaching were an i n s p i r a t i o n . i i ABSTRACT The Habitat Type c l a s s i f i c a t i o n system, as introduced in coastal B r i t i s h Columbia by MacMillan Bloedel Limited, was used for c l a s s i f i c a t i o n of old-growth plant communities in four vegetation zones on central eastern Vancouver Island. A t o t a l of 14 forest habitat types were i d e n t i f i e d . A combination of ordination and t r a d i t i o n a l association table synthesis was used in defining types. Polar ordination and an improved reciprocal averaging technique named DEtrended CORrespondence ANAlysis (DECORANA) were found to be useful for both the c l a s s i f i c a t i o n and evaluation of environmental relationships among the types. Findings substantiate claims that DECORANA i s the best general purpose, indirect ordination technique currently available. In most cases, DECORANA produced superior results to polar ordination with the same data input. The most successful ordinations of individual vegetation zones were obtained for understory percent coverage data transformed to an 8-point scale. Ordinations of tree species alone, or tree and understory species combined were less useful. The d i s t r i b u t i o n of old-growth vegetation in response to environmental gradients was interpreted. A moisture gradient accounted for most of the var i a t i o n between habitat types within any single vegetation zone. Interpretation of the moisture gradient was tested using a Water Stress Index (WSI), which predicts potential s o i l moisture stress from s i t e and climatic data. Climatic data used for input in the WSI model were extrapolated from the nearest available Resource Analysis Branch (RAB) or Atmospheric Environment Service (AES) cli m a t o l o g i c a l stations. S o i l texture, rooting depth, coarse fragment content and depth of the surface organic horizons were used to calculate Available Water Storage Capacity (AWSC) for use in the model. The WSI proved to be a useful integrator for general s i t e comparison of potential moisture stress. In most cases, the WSI prediction showed a closer relationship to ordination axes interpreted as moisture gradients than any other single factor, except for s i t e s with primary moisture inputs from seepage, high water table, or those with complex orographic r a i n f a l l influences. Lack of accurate climatic data, d i f f i c u l t y in quantifying s o i l moisture relationships and the influence of complex microclimatic effects reduced the u t i l i t y of the index. The temperature data available were suitable for quantifying some of the broad macroclimatic differences between zones, and for evaluating differences between samples at the elevational extremes within the Tsuga heterophylla and Abies  amabi1is - Tsuga heterophylla vegetation zones. They were not adequate for a comparison of many of the plots on a s i t e -s p e c i f i c basis because of problems in extrapolation from a limited data base and because of the exclusion of s o i l temperature and cold a i r drainage relationships. Potential solar radiation was used as an integrator of slope, aspect and l a t i t u d e . Some habitat types were found to occur more frequently in certain radiation environments. The elevational location of vegetation zones varied by as much as 80 metres as a result of solar radiation e f f e c t s . Detailed transect sampling of continuous old-growth forests i v in leave-strips in the Cameron and Nanaimo River valleys revealed substantial differences in species composition and d i s t r i b u t i o n between the two valleys that were accounted for by climate and overall s o i l texture. The wetter climate and f i n e r -textured s o i l s of the Cameron valley supported habitat types dominated by Polyst ichum muniturn (swordfern) and Achlys  t r i p h y l l a ( v a n i l l a leaf) with an absence of Gaultheria shallon (salal) dominated habitat types. The drier climate and coarser-textured s o i l s in the Nanaimo valley supported salal-dominated habitat types along the entire south-facing slope and lower north-facing slope. S i g n i f i c a n t differences in tree species d i s t r i b u t i o n were also observed. Habitat types were related to units in the exis t i n g plant community c l a s s i f i c a t i o n s in B. C. and the P a c i f i c Northwestern U. S. Most of the habitat types could be related to types previously described in B r i t i s h Columbia, and many were similar to those found in coastal Oregon and Washington. Some differences in the f l o r a of Vancouver Island and the mainland were noted. There i s a need for a compilation of the descriptions of a l l of the plant communities on Vancouver Island in a single volume. V TABLE OF CONTENTS ABSTRACT i i LIST OF TABLES x LIST OF FIGURES xv LIST OF APPENDICES xx ACKNOWLEDGEMENTS ".' xxi CHAPTER 1 - INTRODUCTION 1 1.1. Thesis objectives 3 1.2. Ecological c l a s s i f i c a t i o n in B r i t i s h Columbia 4 1.3. Gradient analysis approach 14 CHAPTER 2 - LITERATURE REVIEW . . 17 2.1. Studies of vegetation - environment in B. C 17 2.2. Studies of vegetation - environment in the P a c i f i c Northwest U. S. .' . 19 CHAPTER 3 - STUDY AREA ' ... . 22 3.1. Location and physiography 22 3.2. Bedrock geology 24 3.3. Surf ic i a l geology 24 3.4. S o i l s 27 3.5. Climate 29 3.6. Vegetation 32 CHAPTER 4 - METHODS OF STUDY 33 4.1. F i e l d studies .... 33 4.1.1. Study strategy and reconnaissance 33 4.1.2. Plot selection and sampling 35 4.2. Data analysis 38 4.2.1. Ordination techniques 38 4.2.1.1. Choice of methods 38 4.2.1.2. Polar ordination 39 4.2.1.3. Reciprocal averaging 41 4.2.1.4. DECORANA 42 4.2.1.5. Application of ordination methods 43 4.2.2. Tabular analysis technique 45 4.2.3. Correlation with environmental data 46 4.2.3.1. Climatic data 46 4.2.3.2. Water stress index 50 CHAPTER 5 - RESULTS AND INTERPRETATION 56 5.1. Environmental calculations 56 5.2. Ordination of the entire data set 61 5.3. Description and analysis of the Pseudotsuga  menziesi i (PSME) and Pseudotsuga menziesi i - Tsuga  heterophylla (PSME-TSHE) vegetation zones 66 5.3.1. Description of the vegetation series 66 5.3.2. Description of habitat types 67 5.3.2.1. Pseudotsuga menziesi i - Arbutus menziesi i / Gaultheria shallon (PSME-ARME/GASH) habitat type 69 5.3.2.2. Pseudotsuga menziesi i / Gaultheria  shallon - Berberis nervosa (PSME/GASH-BENE) habitat type 72 5.3.2.3. Pseudotsuga menziesi i / Holodiscus  discolor / Polystichum muni turn (PSME/HODI/POMU) habitat type 73 5.3.2.4. Abies grandis / Polystichum muni turn v i i (ABGR/POMU) habitat type 76 5.3.2.5. Thuja p l i c a t a / Lysichitum americanum (THPL/LYAM) habitat type 78 5.3.3. Ordination results 79 5.3.3.1. DECORANA 79 5.3.3.2. Polar ordination 87 5.3.3.3. Relationship of ordination axes to environmental gradients 90 5.4. Description and analysis of the Tsuga heterophylla (TSHE) vegetation zone 93 5.4.1. Description of the vegetation series 93 5.4.2. Description of habitat types 94 5.4.2.1. Tsuga heterophylla / Gaultheria shallon (TSHE/GASH) habitat type 94 5.4.2.2. Tsuga heterophylla / Gaultheria shallon -Berberis nervosa / Achlys t r i p h y l l a (TSHE/GASH-BENE/ACTR) habitat type 98 5.4.2.3. Tsuga heterophylla / Polystichum muni turn (TSHE/POMU) habitat type 100 5.4.2.4. Unc l a s s i f i e d samples 101 5.4.3. Ordination results 104 5.4.3.1. DECORANA 104 5.4.3.2. Polar ordination 110 5.4.3.3. Relationship of ordination axes to environmental gradients 113 5.5. Description and analysis of the Abies amabilis -Tsuga heterophylla (ABAM-TSHE) vegetation zone 118 5.5.1. Description of the vegetation series 118 v i i i 5.5.2. Description of habitat types 119 5.5.2.1. Chamaecyparis nootkatensis / Gaultheria  shallon (CHNO/GASH) habitat type 120 5.5.2.2. Abies amabilis / Vaccinium alaskaense -Vaccinium parvi folium (ABAM/VAAL-VAPA) habitat type 124 5.5.2.3. Abies amabilis / Achlys t r i p h y l l a -T i a r e l l a t r i f o l i a t a (ABAM/ACTR-TITR) habitat type 126 5.5.2.4. Abies amabilis / Vaccinium alaskaense / Streptopus spp. (ABAM/VAAL/STREPTOPUS) habitat type 129 5.5.2.5. Abies amabilis / Vaccinium alaskaense -Vacc inium ovalifolium / Rubus pedatus (ABAM/VAAL(OV)/RUPE) habitat type 132 5.5.2.6. ABAM / Oplopanax horridum (ABAM/OPHO) habitat type 135 5.5.3. Ordination results 137 5.5.3.1. DECORANA 137 5.5.3.2. Polar ordination 139 5.5.4. Relationship of ordination axes to environmental gradients 139 5.6. Transect comparison 142 5.6.1. Environmental c h a r a c t e r i s t i c s of the S. Nanaimo and Cameron ri v e r valleys 145 5.6.2. D i s t r i b u t i o n of vegetation zones and series ..148 5.6.3. Di s t r i b u t i o n of habitat types 149 5.6.4. Ordination results from grouped transect data 151 5.6.5. Interpretation of environmental gradients ....159 CHAPTER 6 - DISCUSSION 163 6.1. Comparison of habitat types to plant communities in previous c l a s s i f i c a t i o n s 163 6.2. Methodology 168 6.3. Environmental gradients 170 6.4. Ordination as an aid in c l a s s i f i c a t i o n 176 CONCLUSIONS 178 LITERATURE CITED 179 Appendix I 191 Appendix II 204 Appendix III 206 Appendix IV 210 Appendix V 216 Appendix VI 223 X LIST OF TABLES Table I. Hierarchy in the habitat type c l a s s i f i c a t i o n system (from Layser and Schubert 1979). 11 Table II. Selected climatic data for the study area 30 Table I I I . P r e c i p i t a t i o n estimates compared to measured pr e c i p i t a t i o n for selected RAB c1imatological stations. 58 Table IV. Summary of major species for habitat types in the PSME and PSME-TSHE vegetation zones 68 Table V. Summary of major species for habitat types in the TSHE vegetation zone 95 Table VI. Summary of major species for habitat types in the ABAM-TSHE vegetation zone 121 Table VII. Habitat types present in the Cameron and Nanaimo transects 146 Table VIII. Relationship of habitat types to other plant community c l a s s i f i c a t i o n s in B. C. and the P a c i f i c Northwestern U. S 164 Table A l . Tree canopy layers used in f i e l d t a l l y (after Smith 1962) 204 Table A2. Understory and tree canopy layers used in f i e l d t a l l y and data analysis (Brooke et a l . 1970) 204 Table A3. Vigor, s o c i a b i l i t y and abundance scales used in f i e l d observations 205 Table A4. AEA and RAB climatological stations used for data extrapolat ion 210 Table A5. Radiation index table for Water Stress Index (WSI) calculations (Ballard 1974) 211 Table A6. Available water storage in s o i l layers making up the root zone (from Ballard 1974) 212 Table A7. Environmental data for individual sample p l o t s . .213 Table A8. Explanation of terms and abbreviations used in s i t e c h a r a c t e r i s t i c tables ; 216 Table A9. Pseudotsuqa menziesi i - Arbutus menziesi i / Gaultheria shallon (PSME-ARME/GASH) habitat type - s i t e character i s t i e s 217 Table A10. Pseudotsuga menziesi i / Gaultheria shallon -Berberis nervosa (PSME/GASH-BENE) habitat type - s i t e c h a r a c t e r i s t i c s .217 Table A l l . Pseudotsuga menziesi i / Holodiscus discolor / Polystichum munitum (PSME/HODI/POMU) habitat type - s i t e c h a r a c t e r i s t i c s 217 Table A12. Abies grandis / Polystichum munitum (ABGR/POMU) habitat type - s i t e c h a r a c t e r i s t i c s 218 Table A13. Thuja p i i c a t a / Lysichitum amer icanum (THPL/LYAM) habitat type - s i t e c h a r a c t e r i s t i c s 218 Table A14. Tsuga heterophylla / Gaultheria shallon (TSHE/GASH) habitat type - s i t e c h a r a c t e r i s t i c s 219 Table A15. Tsuga heterophylla / Gaultheria shallon -Berberis nervosa / Achlys t r i p h y l l a (TSHE/GASH-BENE/ACTR) habitat type - s i t e c h a r a c t e r i s t i c s 219 Table A16. Tsuga heterophylla / Polyst ichum munitum (TSHE/POMU) habitat type - s i t e c h a r a c t e r i s t i c s 220 Table A17. Chamaecyparis nootkatensis / Gaultheria shallon (CHNO/GASH) habitat type - s i t e c h a r a c t e r i s t i c s 220 Table A18. Abies amabilis / Vaccinium alaskaense  Vaccinium parvifolium (ABAM/VAAL-VAPA) habitat type s i t e c h a r a c t e r i s t i c s 221 Table A19. Abies amabilis / Achlys t r i p h y l l a - T i a r e l l a  t r i f o l i a t a (ABAM/ACTR-TITR) habitat type - s i t e c h a r a c t e r i s t i c s .221 Table A20. Abies amabilis / Vaccinium alaskaense / Streptopus spp. (ABAM/VAAL/STREPTOPUS) habitat type -si t e c h a r a c t e r i s t i c s 222 Table A21. Abies amabi1i s / Vaccinium alaskaense  Vacc inium ovali folium / Rubus pedatus (ABAM/VAAL(OV)/RUPE) habitat type - s i t e character i st i c s 222 Table A22. Abies amabilis / Oplopanax horridum (ABAM/OPHO) habitat type - s i t e c h a r a c t e r i s t i c s 222 Table A23. Explanation of terms and abbreviations used in the association tables 223 Table A24. Association tables for the Pseudotsuga menziesii  Arbutus menziesi i / Gaulther ia shallon (PSME-ARME/GASH) habitat type 224 Table A25. Association tables for the Pseudotsuga menziesi i / Gaultheria shallon - Berberis nervosa (PSME/GASH-BENE) habitat type 225 Table A26. Association tables for the Pseudotsuga menziesii / Holodiscus discolor / Polystichum muni turn (PSME/HODI/POMU) habitat type 226 Table A27. Association tables for the Abies grandis / Polystichum muni turn (ABGR/POMU) habitat type 227 Table A28. Association tables for the Thuja p l i c a t a / Lysichitum americanum (THPL/LYAM) habitat type 228 Table A29. Association tables for the Tsuga heterophylla / Gaultheria shallon (TSHE/GASH) habitat type 229 Table A30. Association tables for the Tsuga heterophylla / Gaultheria shallon - Berberis nervosa / Achlys t r i p h y l l a (TSHE/GASH-BENE/ACTR) habitat type 230 Table A31. Association tables for the Tsuga heterophylla / Polystichum muni turn (TSHE/POMU) habitat type 231 Table A32. Association tables for the u n c l a s s i f i e d samples in the TSHE zone 232 Table A33. Association tables for the Chamaecyparis  nootkatensis / Gaultheria shallon (CHNO/GASH) habitat type 233 Table A34. Association tables for the Abies amabilis / Vaccinium alaskaense - Vaccinium parvifolium (ABAM/VAAL-VAPA) habitat type '..234 Table A35. Association tables for the Abies amabilis / Achlys t r i p h y l l a - T i a r e l l a t r i f o l i a t a (ABAM/ACTR-TITR) habitat type 235 Table A36. Association tables for the Abies amabilis / Vacc inium alaskaense / Streptopus spp. (ABAM/VAAL/STREPTOPUS) habitat type 236 Table A37. Association tables for the Abies amabilis / Vacc inium alaskaense - Vacc inium ovali folium / Rubus  pedatus (ABAM/VAAL(OV)/RUPE) habitat type 237 Table A38. Association tables for the Abies amabilis / Oplopanax horridum (ABAM/OPHO) habitat type 238 xiv Table A39. Association tables for the samples in the ABAM-TSME vegetation zone 239 X V LIST OF FIGURES Figure 1. Study area and location of cl i m a t b l o g i c a l stations. 23 Figure 2. South Nanaimo River v a l l e y looking northwest. ... 34 Figure 3.-Cameron River valley looking southeast •••• 34 Figure 4. P r e c i p i t a t i o n versus temperature for sample pl o t s . 59 Figure 5. Temperature versus potential solar radiation for sample p l o t s . . . . 60 Figure 6. P r e c i p i t a t i o n versus AWSC for sample p l o t s . ..... 62 Figure 7. DECORANA ordination of a l l samples showing the d i s t r i b u t i o n of vegetation zones. 63 Figure 8. Pseudotsuga menziesi i - Arbutus menziesi i / Gaultheria 1shallon (PSME-ARME/GASH) habitat type 70 Figure 9. Pseudotsuga menziesi i / Gaultheria shallon Berberis nervosa (PSME/GASH-BENE) habitat type 70 Figure 10. Pseudotsuga menz i esi i / Holodi scus d i s c o l o r / Polystichum munitum (PSME/HODI/POMU) habitat type 75 Figure 11. Abies grandi s / Polyst ichum mun i turn (ABGR/POMU) habitat type . . 75 Figure 12. Thuja p i i c a t a / Lysi c h i turn americanum (THPL/LYAM) habitat type 80 Figure 13. DECORANA ordination of samples in the PSME and PSME-TSHE zones - vegetation s e r i e s . 82. Figure 14. DECORANA ordination of samples in the PSME and PSME-TSHE zones - habitat t y p e s . ' 82 xvi Figure 15. DECORANA ordination of species in the PSME and PSME-TSHE zones 84 Figure 16. Reciprocal averaging ordination of samples in the PSME and PSME-TSHE zones - habitat types 85 Figure 17. DECORANA ordination of samples in the PSME and PSME-TSHE zones - habitat types (understory only) 85 Figure 18. DECORANA ordination of samples in the PSME and PSME-TSHE zones (overstory only) using basal area 88 Figure 19. Polar ordination of samples in the PSME and PSME-TSHE zones - habitat types (understory only) 88 Figure 20a. DECORANA ordination of samples in the PSME and PSME-TSHE zones - vegetation series, with WSI plotted. . 92 Figure 20b. DECORANA ordination of samples in the PSME and PSME-TSHE zones, with AWSC plotted 92 Figure 21. Tsuga heterophylla / Gaultheria shallon (TSHE/GASH) habitat type 97 Figure 22. Tsuga heterophylla / Gaultheria shallon -Berberis nervosa / Achlys t r i p h y l l a (TSHE/GASH-BENE/ACTR) habitat type 97 Figure 23. Tsuga heterophylla / Polyst ichum muniturn (TSHE/POMU) habitat type 102 Figure 24. Tsuga heterophylla zone - u n c l a s s i f i e d sample with depauperate understory 102 Figure 25. DECORANA ordination of samples in the TSHE zone - habitat types and series ( a l l species) 105 Figure 26. DECORANA ordination of samples in the TSHE zone - habitat types and series (understory only) 105 Figure 27. Di s t r i b u t i o n of major understory species on a xvi i DECORANA understory ordination 108 Figure 28. Dis t r i b u t i o n of species and habitat types along an RA ordination axis - TSHE zone 109 Figure 29. DECORANA ordination of samples in the TSHE zone, with temperature plotted (overstory only) I l l Figure 30. Polar ordination of samples in the TSHE zone with TSHE/GASH endpoints chosen for high WSI, radiation. 112 Figure 31. Polar ordination of samples in the TSHE zone with TSHE/GASH-BENE/ACTR endpoints chosen for high WSI, radiat ion 112 Figure 32. Environmental variables plotted on the DECORANA understory ordination of Figure 26: 114 a. Water Stress Index (WSI) 114 b. Potential Annual Solar Radiation 114 c. Average P r e c i p i t a t i o n (May - September) 114 Figure 33. Chamaecyparis nootkatensis / Gaultheria shallon (CHNO/GASH) habitat type 123 Figure 34. Abies amabilis / Vaccinium alaskaense Vaccinium parvi folium (ABAM/VAAL-VAPA) habitat type ....123 Figure 35. Abies amabilis / Achlys t r i p h y l l a - T i a r e l l a t r i f o l i a t a (ABAM/ACTR-TITR) habitat type 128 Figure 36. Abies amabi1i s / Vaccinium alaskaense / Streptopus spp. (ABAM/VAAL/STREPTOPUS) habitat type 128 Figure 37. Abies amabilis / Vacc inium alaskaense Vacc inium ovalifolium / Rubus pedatus (ABAM/VAAL(OV)/RUPE) habitat type 133 Figure 38. Abies amabilis / Oplopanax horridum (ABAM/OPHO) xvi i i habitat type 133 Figure 39. DECORANA ordination of samples in the ABAM-TSHE zone - habitat types 138 Figure 40. DECORANA ordination of samples in the ABAM-TSHE , zone (overstory only) - vegetation s e r i e s . 138 Figure 41. Polar ordination of samples in the ABAM-TSHE zone - habitat types 140 Figure 42. Environmental variables plotted on a DECORANA ordination of samples in the ABAM-TSHE zone: 141 a. Water Stress Index (WSI) 141 b. Elevation 141 Figure 43. Di s t r i b u t i o n of vegetation in the South Nanaimo River valley 143 Figure 44. Di s t r i b u t i o n of vegetation in the Cameron River valley 144 Figure 45. DECORANA ordination of samples in the Cameron and Nanaimo transects zone - vegetation series 152 Figure 46. DECORANA ordination of samples in the Cameron and Nanaimo transects - habitat types (understory only). 152 Figure 47. Species-by-sample matrix in RA axis 1 ordination order for samples in the Cameron and Nanaimo transects 155 Figure 48. Polar ordination of samples in the Cameron and Nanaimo transects -- habitat types (understory only). ..157 Figure 49. DECORANA ordination of samples in the Cameron and Nanaimo transects using overstory basal area - ' series 157 xix Figure 50. DECORANA ordination of overstory species in the Cameron and Nanaimo transects (accompanying Figure 49). 158 Figure 51. Species-by-sample . matrix in RA axis 1 ordination order for an ordination of tree species, Cameron and Nanaimo transects 160 Figure 52. Environmental variables plotted on a polar ordination for the Cameron and Nanaimo transects: 161 a. WSI and Potential Solar Radiation 161 b. Average Growing Season Temperature ( °C) 161 0 X X LIST OF APPENDICES Appendix I. Keys to the Vegetation Zones and Vegetation Series of south coastal B r i t i s h Columbia (Packee 1979). 191 Appendix II. Canopy layers and vigor, abundance and s o c i a b i l i t y classes used in f i e l d observations and data analysi s 204 Appendix I I I . Latin and common names of species found on sample plots 206 Appendix IV. Climatic stations, radiation tables, available water storage tables and environmental data used for input in the Water Stress Index (WSI) model 210 Appendix V. Site c h a r a c t e r i s t i c s for individual sample plots by habitat type 216 Appendix VI. Association tables for the habitat types 223 xxi ACKNOWLEDGEMENTS I would l i k e to express my sincere gratitude to my advisor, Dr. J. P. Kimmins, for his counselling and encouragement throughout the study; to my committee, Dr. Gary Bradfield and Dr. Roy Strang, for their help and guidance; and to Dr. Ed Packee of MacMillan Bloedel Limited for his generous support and enthusiasm for the project. Special thanks are extended to Kirsteen Laing, Margaret Symon and A l i s t a i r Handley for their cheerful, ambitious f i e l d assistance. Susan Phelps also deserves a special thank-you for her constant help with computer challenges. I would also l i k e to thank Carol Kennedy and the staff of the MB Woodlands Services laboratory for analysis of s o i l samples; Ms. B. Atkey and Mrs. B. Beese for assistance in typing the manuscript; M. C. Coligado of the RAB for his help with compiling climatic data; Dr. Andy Black, for his assistance with questions on climatology; Dr. T. M. Ballard, for use of the Water Stress Index and his counsel during the analysis; John Pinder-Moss, for his assistance in using the UBC Botany Herbarium; and to the UBC MTS system, for providing the "tools" for the analysis and manuscript preparation. F i n a l l y , I am grateful for the opportunity and f i n a n c i a l support made available for thi s study by MacMillan Bloedel Limited; for the support, f a c i l i t i e s , faculty, technicians and o f f i c e staff of the Faculty of Forestry; and for f i n a n c i a l assistance by the Vancouver Foundation. 1 CHAPTER 1 INTRODUCTION Forest management in North America faces many challenges in the decades ahead, most of which depend upon a t r a n s i t i o n from harvesting old-growth forests to applying intensive forest management practices to younger stands. An essential element in this process is the c l a s s i f i c a t i o n of land units to f a c i l i t a t e s i t e - s p e c i f i c management. The need for c l a s s i f i c a t i o n of forest ecosystems has been recognized throughout the world. This has resulted in numerous c l a s s i f i c a t i o n schemes, each with a pa r t i c u l a r methodology and conceptual model of the nature of forest vegetation. As a result of th i s d i v e r s i t y , there s t i l l e xists today some disagreement about how to c l a s s i f y forests. There i s no need to defend ecological s i t e c l a s s i f i c a t i o n : the advantages for communication, predicting responses to management treatment, and extrapolation of research results are well-documented (Daubenmire 1952, Krajina 1960, Kimmins 1977, Jones 1978). The need to c l a s s i f y was recognized by early phytosociologists in Europe and North America. From five major h i s t o r i c a l t r a d i t i o n s , twelve schools of vegetation c l a s s i f i c a t i o n have developed (Kessell 1979), as reviewed by Whittaker (1962, 1973). One might l i k e to believe that a single system would be desirable; however, d i f f e r i n g regional vegetation c h a r a c t e r i s t i c s and c l a s s i f i c a t i o n objectives make dif f e r e n t approaches advantageous. No single approach i s suitable for a l l possible uses in a l l areas of the world. Unfortunately, a problem arises when several c l a s s i f i c a t i o n systems are applied to the same set of ecosystems within a 2 region. Competition for the adoption of a p a r t i c u l a r approach can threaten the usefulness of c l a s s i f i c a t i o n for communication purposes and often leads to unnecessary duplication of e f f o r t . Several important steps are necessary to increase the usefulness and r e l i a b i l i t y of t r a d i t i o n a l forest c l a s s i f i c a t i o n systems and to evaluate which of several alternative systems is the best for a given area. F i r s t , there i s a need to apply quantitative, objective numerical techniques to substantiate subjective c l a s s i f i c a t i o n schemes. While there i s no mechanical substitute for human interpretation, the objective testing of subjective c l a s s i f i c a t i o n e f f o r t s using numerical methods can help i d e n t i f y subtle relationships and avoid the danger of f i t t i n g samples into a pre-conceived mental model. Secondly, there is a need to compare d i f f e r e n t c l a s s i f i c a t i o n s of the same landscape. This i s p a r t i c u l a r l y relevant in coastal B r i t i s h Columbia where some controversy exists over which of several c l a s s i f i c a t i o n systems is the most useful for forest management. Thirdly, there is a need to e s t a b l i s h the significance of s t r u c t u r a l l y — defined ecosystem units. Environmental and physiological features of p r a c t i c a l concern must be related to readily i d e n t i f i a b l e c l a s s i f i c a t i o n units i f a system i s to function as a predictive tool in forest management decisions. This thesis addresses steps 1 and 3 in this process. Step 2 requires a detailed application of several c l a s s i f i c a t i o n systems to a s p e c i f i c landscape, which i s not the intent of t h i s invest igat ion. 3 1.1. Thesis objectives The o v e r a l l objectives of thi s thesis are: 1. To investigate community-environment and species-environment relationships by comparing vegetation composition and species d i s t r i b u t i o n s to measurable gradients of environmental features, and 2. To assess the usefulness of ordination as an aid in iden t i f y i n g vegetation units at various h i e r a r c h i c a l l e v e l s , and in v e r i f y i n g pre-existing c l a s s i f i c a t i o n units. The s p e c i f i c objectives of thi s thesis are: 1. To investigate the effectiveness of an improved reciprocal averaging technique (DECORANA) and Wisconsin Polar ordination in i d e n t i f y i n g vegetation zones, series and habitat types from ordinations of overstory and understory species, separately and combined. 2. To ide n t i f y the r e l a t i v e positioning of vegetation communities along ordination axes and interpret these relationships with regard to gradients of moisture, temperature, and solar radiation. 3. To assess the u t i l i t y of a Water Stress Index (WSI) model for predicting s o i l moisture stress using existing c l i m a t i c data and information c o l l e c t e d from a single v i s i t to a s i t e , and to evaluate i t s usefulness in interpreting a moisture gradient. 4. To use Braun-Blanquet association table analysis, assisted by the B.C. Forest Service table preparation program and the results of ordination, to characterize units of the habitat type c l a s s i f i c a t i o n system, and to assess the value of using these methods in combination for ecosystem c l a s s i f i c a t i o n in B r i t i s h Columbia. The usefulness of a forest s i t e c l a s s i f i c a t i o n system depends, to a large degree, upon the forester's a b i l i t y to identify units in the f i e l d and make q u a l i t a t i v e judgements 4 about a s i t e for management purposes. This study attempts to correlate e a s i l y measured s o i l and topographic features and information from pre-existing climatic stations, geologic surveys and landform maps with semi-quantitative vegetation measurements to determine what the forester may or may not be able to infer from vegetation "indicators". A j u s t i f i c a t i o n for th i s approach is that laboratory analysis of s o i l data and complex climatic measurements are both time-consuming and cos t l y ; consequently, they are not always available to the f i e l d forester in situations that require a quick decision. This study uses the habitat type c l a s s i f i c a t i o n system which has been applied throughout the western U.S. and has been introduced in Coastal B r i t i s h Columbia by MacMillan Bloedel Limited. The only previous applications in Canada were in south central B.C. (McLean 1970) and southern Alberta (Ogilvie 1963). Following the c l a s s i f i c a t i o n of vegetation units, the analysis of their relationship to environment w i l l be attempted using several computer-assisted ordination techniques which have proven useful elsewhere, but have not been applied in any published vegetation studies in B.C. It i s hoped that t h i s investigation w i l l contribute to the ultimate goal of l i n k i n g readily-recognizable vegetation c l a s s i f i c a t i o n units with other s i t e c h a r a c t e r i s t i c s in order to f a c i l i t a t e intensive, s i t e -s p e c i f i c forest management practices. 1.2. Ecological C l a s s i f i c a t i o n in B r i t i s h Columbia Since the early 1950's, Dr. V.J. Krajina of the University of B r i t i s h Columbia Department of Botany and many of his 5 students have undertaken the important task of characterizing the d i v e r s i t y of ecosystems in B.C. Ideally, a l l b i o t i c and ab i o t i c attributes of an area should be evaluated and form the basis for the c l a s s i f i c a t i o n of ecosystems. This "ecosystematic" approach is advocated by Krajina, and is the foundation of the Biogeoclimatic c l a s s i f i c a t i o n system (Krajina 1959, 1965, 1972). Biogeoclimatic c l a s s i f i c a t i o n i s an h i e r a r c h i c a l system based on two levels of c l a s s i f i c a t i o n : biogeoclimatic and biogeocoenotic (Klinka e_t a_l. 1979). At the biogeoclimatic l e v e l , five categories are recognized: 1) formation, 2) region, 3) zone, 4) subzone, and 5) variant. The formation and region are broad areas of similar macroclimate, dominant s o i l processes and vegetation physiognomy of zonal ecosystems. Zonal ecosystems are those in which climate is expressed most strongly, moisture and nutrient conditions are intermediate in rel a t i o n to other ecosystems in the zone, and vegetation development, when free from major disturbance, leads to the "climatic climax" state (Klinka et a l . 1979). E a r l i e r publications used the term "mesic" in the same context as zonal (Klinka 1977). A climax ecosystem is one in which species composition of the vegetation is s e l f -perpetuating over time. 1 Biogeoclimatic zones are defined by Krajina (1969) as "geographical segments of the earth's climate, s o i l and biota that are characterized by the same general macroclimate and the same zonal s o i l and biota (phytocoenosis 1There i s some controversy in ecological l i t e r a t u r e concerning the v a l i d i t y of the concept of "climax". In this thesis, climax vegetation refers to old-growth forests thought to represent most closely the potential self-regenerating plant community that developes in the absence of disturbance. 6 and zoocoenosis)". Descriptions of zones include the ranges of macroclimatic variables and the c h a r a c t e r i s t i c s of zonal ecosystems. Accessory features include a c h a r a c t e r i s t i c combination of species and pattern of plant orders occurring in the zone. Zones are named for the climatic climax tree species found in zonal ecosystems, and may be preceded by a broad location description (e.g. Coastal Douglas-fir zone). The subzone is defined by s p e c i f i c ranges in climatic variables and by zonal ecosystems c l a s s i f i e d at the biogeocoenotic association l e v e l . Subzone names carry additional descriptive terms (e.g. Drier Maritime Coastal Douglas-fir subzone). Krajina's (1959, 1965) descriptions of zones and subzones were accompanied by descriptions of predominant pedogenic processes. The most recent use of the biogeoclimatic approach has not used s o i l as a d i f f e r e n t i a t i n g or accessory c h a r a c t e r i s t i c defining zone, subzone or variant (Klinka et a l . 1979). Subzones are subdivided into" variants on the basis of climate and associated vegetation. The biogeocoenotic type i s used to c l a s s i f y zonal ecosystems in t h i s category. Names of variants include a physiographic region modifier (e.g. Nanaimo and Georgia Drier Montane Coastal Douglas-fir variant). Occasionally, a biogeoclimatic phase i s recognized where l o c a l r e l i e f a f f e c t s regional climate, as in the case of aspect differences or cold a i r drainage. This designation i s not considered part of biogeoclimatic c l a s s i f i c a t i o n . Categories in the biogeocoenot ic l e v e l are distinguished on the basis of vegetation and s o i l . The basic unit at t h i s l e v e l 7 is the biogeocoenotic association. Associations are d i f f e r e n t i a t e d by c h a r a c t e r i s t i c combinations of species and s o i l moisture and nutrient regime. Related associations are grouped into plant orders. 2 Associations are subdivided into biogeocoenotic types 3 primarily on the basis of edaphic features, though there may be s l i g h t f l o r i s t i c differences evident. Types are further subdivided into biogeocoenotic variants on the basis of v a r i a b i l i t y in successional stage due to disturbance. Plant orders and associations are named with c h a r a c t e r i s t i c understory species preceding the dominant tree species. Type names include the s o i l order, great group or subgroup and a humus form d e s c r i p t i o n . 4 The preceding description of the biogeoclimatic approach is based on Klinka et a l . (1979). E a r l i e r descriptions include those of Krajina (1959, 1965, 1969, 1972), Klinka (1977), Jones and Annas (1978), Mueller-Dombois and Ellenberg (1974) and B e i l et a l . (1976). The biogeoclimatic system has been adopted by the B r i t i s h Columbia Ministry of Forests for applying s i t e - s p e c i f i c management prescriptions. Guides for tree species selection and prescribed burning have been prepared on t h i s basis (Klinka 1977, Utzig et a l . 1978). 2"Associations which are related in f l o r i s t i c composition, successional trends, broad environment-vegetation relationships, and p r e v a i l i n g pedogenic processes." (Klinka et a l . 1979) 3 The biogeocoenosis type of Sukachev (1944), Sukachev and Dy l i s (1964). 4A new, comprehensive humus form c l a s s i f i c a t i o n has recently been developed in B. C. (Klinka e_t a l . 1980a) based largely upon work by Bernier (1968) in an attempt to standardize a previously unclear aspect of s o i l description. 8 A d i f f e r e n t approach to forest ecosystem c l a s s i f i c a t i o n has been introduced in B r i t i s h Columbia by Dr. E. C. Packee, MacMillan Bloedel Limited. This approach i s known as the habitat type c l a s s i f i c a t i o n system. It was developed by Daubenmire (1952, 1968) in eastern Washington and northern Idaho, and has since been applied by the U.S. Forest Service throughout the western United States ( P f i s t e r 1977). Its main difference from the biogeoclimatic system i s that c l a s s i f i c a t i o n units are defined by vegetation c h a r a c t e r i s t i c s alone. Though relationships between climatic and edaphic factors and vegetation units are recognized, these features are not used as an integral part of defining units, as i s the case in the biogeoclimatic approach. There are two sets of terms that have been used to characterize the habitat type system. One relates to vegetation taxonomy, the other to geographic mapping l e v e l s . These terms have caused some confusion, which has been largely c l a r i f i e d by Bailey et a l . (1978) and by Layser and Schubert (1979). The habitat type system as outlined by Daubenmire (1968) recognizes four levels in a landscape hierarchy: habitat type, vegetation zone, vegetation province and vegetation region. The basic unit in the system i s the habitat type, defined as: " a l l parts of the landscape that support, or are capable of s u p p o r t i n g t h e same kind of r e l a t i v e l y stable phytocoenosis (homogeneous as to dominants in a l l layers) in the absence of disturbance" (Daubenmire 1968). Consequently, the potential climax plant association i s the basis for c l a s s i f i c a t i o n of a given s i t e , even though a serai community may be present. 9 Habitat types are named by the climax tree species (most shade tole r a n t ) , followed by the dominant understory species (e.g. Pseudotsuga menziesii / Gaultheria shallon habitat type). The habitat type i s e s s e n t i a l l y the same as the climax plant association of Krajina (1960, 1965). A habitat type may be subdivided into phases where differences in environmental c h a r a c t e r i s t i c s result in minor variation in species composition. Such variation may be the result of aspect, topography, s o i l or other environmental features. The next l e v e l in the hierarchy i s the vegetation zone. A vegetation zone is a geographical area of generally uniform macroclimate supporting the same climatic climax association in zonal ecosystems (Daubenmire 1968). Zonal i s defined as in the biogeoclimatic system. Zones are named for the dominant climatic climax tree species (e.g. Pseudotsuga menziesii vegetation zone). Macroclimate for vegetation zones i s e s s e n t i a l l y similar to that of biogeoclimatic subzones (Packee 1976). Zones are grouped into vegetation provinces occupying a d i s t i n c t i v e geographical area and sharing a "somewhat common and d i s t i n c t i v e geologic history" and "strong threads of taxonomic homogeneity" (Daubenmire 1968). Vegetation provinces are grouped into vegetation regions representing broad geographical areas having a c h a r a c t e r i s t i c physiognomy and climatic regime. 5 The habitat type system has also been described as a taxonomic vegetation hierarchy with the following l e v e l s : 5Daubenmire's "region" emphasizes geographic area but i s at the same level as the "formation" of Clements. Region has been used d i f f e r e n t l y by Arno (1979) and Bailey et a l . (1979). 10 association, series, subformation and formation (Layser and Schubert 1979). The association is the climax plant community that occupies the physical environment or landscape unit c a l l e d the habitat type. The vegetation series i s a grouping of a l l associations with the same dominant tree species at climax (Daubenmire and Daubenmire 1968). The series i s recognized by evaluating the reproductive success of tree species. Climax communities at the series (and association) l e v e l may be the result of c l i m a t i c , edaphic, topographic or topo-edaphic conditions; hence, the polyclimax concept of Tansley (1935) i s recognized. The two highest levels are based on physiognomy. The formation includes vegetation with the same potential physiognomy at climax and may be divided into subformations on the basis of " d i s t i n c t i v e physiognomy within a formation" (Daubenmire 1968). Table I summarizes these h i e r a r c h i c a l l e v e l s . Most applications of the habitat type system have used "habitat type" and "association" interchangeably ( i . e . i t is of l i t t l e p r a c t i c a l value to dis t i n g u i s h between the landscape unit and the plant community occupying i t ) and have grouped habitat types (or associations) into series (Hoffman and Alexander 1976, P f i s t e r e_t a_l. 1977, Henderson et a l . 1977, Layser and Schubert 1979, Moir and Ludwig 1979). The term "community type" has also been used at the same l e v e l as the habitat type to designate areas of serai and/or uncertain successional status (Dyrness et a l . 1974, P f i s t e r et a l . 1977, Franklin et a l . 1979). Some workers in Oregon and Washington have not used the Table I. Hierarchy i n the habitat type c l a s s i f i c a t i o n system (from Layser and Schubert 1979). TAXONOMIC TERMS GEOGRAPHIC TERMS C l a s s i f i c a t i o n l e v e l s in the taxonomic hierarchy Mapping l e v e l s in the physical environment FORMATION SUBFORMATION SERIES ASSOCIATION ''community type." •PHASE REGION PROl/INCE ZONE HABITAT TYPE PHASE 12 series l e v e l , but rather have used associations and zones (Franklin and Dyrness 1973, Dyrness et a l . 1974) or habitat types and zones (Franklin e_t a l . 1979); The forested zones of Franklin and Dyrness (1973) are not defined according to Daubenmire (1968). Instead, "...forested zones are not based on climatic climax communities, but rather on areas in which a single tree species i s the major climax dominant." This difference i s greater in an academic sense than in. actual practice. In applying the habitat type system in B. C , Packee (1976) has used the zone (of Franklin), series and habitat type. He i s the only author who has used both the zone and series in a single c l a s s i f i c a t i o n . This may be c r i t i c i z e d in one sense because i t combines both geographic and taxonomic terms in the same c l a s s i f i c a t i o n , thus deviating from a true hierarchy. He has also used the Biome and Bi o t i c Province at higher l e v e l s , which are ecosystem classes rather than vegetation classes. This sheds some confusion over the units being c l a s s i f i e d . Nevertheless, there may be some j u s t i f i c a t i o n for using both zones and se r i e s . Zones e s s e n t i a l l y id e n t i f y a l l areas with the same climatic climax series on modal s i t e s . Other series exist within a zone as edaphic climaxes. The series l e v e l can, therefore, be a useful subdivision of the zone above the l e v e l of the habitat type to distinguish dominant tree species. Even though the same series may occur in more than one zone (as can associations), a series (or association) i s only a cli m a t i c climax in a single zone. This approach has been c r i t i c i z e d for adding an unnecessary 13 l e v e l that i s accounted for at the habitat type l e v e l . It has been j u s t i f i e d as useful for interpretation of species selection for reforestation without the need for i d e n t i f i c a t i o n of habitat types. This thesis considers both the zone and series as already applied in B. C , while recognizing the h i e r a r c h i c a l problems with t h i s approach. The habitat type c l a s s i f i c a t i o n method has several desirable features. One i s the use of dichotomous "keys" to aid i d e n t i f i c a t i o n of communities in the f i e l d . Such a key has been developed for zones and series on Vancouver Island and the south coastal mainland of B.C. (Packee 1979). Another feature i s the use of standardized abbreviations for the names of plant species, derived from the f i r s t two l e t t e r s of the l a t i n generic name and s p e c i f i c epithet (e.g. Pseudotsuga menziesii = PSME). A number i s added in cases where these are the same for more than one species (Garrison e_t a l . 1976, Packee 1981). Abbreviations are used in the naming of zones, series and habitat types (e.g. Pseudotsuga menziesi i / Gaultheria shallon habitat type PSME/GASH). A slash (/) i s used to separate species of d i f f e r e n t growth form, and a dash (-) for species of the same growth form. This nomenclature i s convenient for computer analysis and f i e l d notes where f u l l species names are impractical. A preliminary guide for tree species selection and prescribed burning has been prepared on the basis of d i f f e r e n t moisture regimes in tentative vegetation series on Vancouver Island (Packee 1979). B r i t i s h Columbia has also been c l a s s i f i e d according to the Forest Regions of Rowe (1972) and Forest Cover Types of the 14 s Society of American Foresters (1967). These systems are intended to cover broad geographical areas; consequently, they are not appropriate for s i t e - s p e c i f i c ecosystem c l a s s i f i c a t i o n and w i l l not be discussed here. For a review of these schemes for B r i t i s h Columbia see Packee (1976). Other c l a s s i f i c a t i o n s that have been applied in B.C. are reviewed by Jones (1978). 1.3. Gradient analysis approach Gradient analysis is an approach to vegetation description that relates gradients of environmental factors to vegetation composition and structure. There are two main types of gradient analysis: d i r e c t and i n d i r e c t . In d i r e c t gradient analysis, vegetation samples are arranged and studied in r e l a t i o n to measured environmental gradients. Whittaker used th i s technique in several studies (Whittaker 1956, 1960; Whittaker and Niering 1964, 1965) and has described the methodology in d e t a i l (Whittaker 1967, 1973, 1975). With the indirect method, environmental gradients are inferred from an ordering of vegetation samples through calculations of sample s i m i l i a r i t y . Some of the f i r s t applications of indirect techniques were by Bray and Curtis (1957), Bray (1956, 1960, 1961) and Goodall (1954). Both methods u t i l i z e the technique of ordination, in which samples are arranged with respect to one or more axes of v a r i a t i o n . These axes may represent either d i r e c t environmental measurements or compositional gradients derived solely from analyzing the vegetation data. Several mathematical techniques are available for 15 ordination, which have been presented and compared in recent l i t e r a t u r e (Gauch and Whittaker 1972, Kessell and Whittaker 1976, Gauch et a l . 1977, Noy-Meir and Whittaker 1977, Fasham 1977). Four of the more common techniques are: Wisconsin polar ordination (Bray and Curtis 1957); weighted averaging (Ellenberg 1948, Whittaker 1948, Curtis and Mcintosh 1951); p r i n c i p a l component analysis (Goodall 1954); and reciprocal averaging ( H i l l 1973). B r i e f l y , weighted averaging assigns species scores based upon known, or assumed, ecological preference of species to some environmental factor ( i . e . moisture) and uses these values to derive sample ordination scores. Polar ordination arranges samples in r e l a t i o n to two chosen reference samples by comparing each sample to these "endpoints" using some index of sample s i m i l a r i t y . P r i n c i p a l component analysis (PCA) uses eigenvector analysis to extract axes of va r i a t i o n from a species-by-sample matrix viewed as points in a multidimensional space. Reciprocal averaging i s an eigenanalysis method similar to PCA but uses a d i f f e r e n t type of standardization. Polar ordination and reciprocal averaging were chosen for th i s study because investigations have shown that they produce superior results to other methods (Gauch e_t a l . 1977, Kessell 1979). They w i l l be described more f u l l y in Chapter 4. For a more complete explanation of p r i n c i p a l component analysis, the reader may consult G i t t i n s (1969), Pielou (1977) and Orloci (1978). The weighted averages method is explained f u l l y by Whittaker (1967). Ordination is a useful technique for data reduction, for analysis of relationships between samples or between species, 16 and for comparison of variation in a body of vegetation data to environmental gradients. Moreover, ordination may a s s i s t in c l a s s i f i c a t i o n by grouping together samples that are similar and segregating samples that have l i t t l e in common. It should be noted, however, that the delineation of groups i s done subjectively by the user and is not part of the ordination process. Ordination has proven useful in recent work in c l a s s i f i c a t i o n (Pfister et a l . 1977) and in . p r a c t i c a l application to forest fuels management when combined with computer modeling in a technique named "gradient modeling" (Kessell 1979). 17 CHAPTER 2 LITERATURE REVIEW There have been numerous studies of vegetation-environment relationships in coastal B r i t i s h Columbia, Washington and Oregon. Some of the more recent of these w i l l be reviewed b r i e f l y as they relate to t h i s thesis. 2.1. Studies of vegetation-environment in B r i t i s h Columbia Studies of vegetation-environment relationships in B.C. include those carried out by Krajina and his students on Vancouver Island (Krajina 1969). Early studies were concentrated in the Douglas-fir region of the east coast. Krajina and Spilsbury (1953) developed the f i r s t c l a s s i f i c a t i o n of these forests, identifying seven associations for the Douglas-fir zone. Szczawinski (1953) studied the lichen and bryophyte plant communities in the same region. Later, McMinn (1957, 1960) investigated the water relationships of six plant associations occurring in the Douglas-fir forests of the Nanaimo River v a l l e y . He concluded that s o i l moisture regime i s a very important factor contributing to s i t e v a r i a t i o n . In the same vall e y , Mueller-Dombois (1959, 1965) studied secondary succession in recently-logged Douglas-fir forests and compared vegetation structure and composition with that of old-growth and older cut-over s i t e s . A complete description of the vegetation, s o i l s and successional stages of six associations was presented along with a discussion of the r e l a t i v e variation in vegetation c h a r a c t e r i s t i c s with regard to time since disturbance and i t s implication for s i t e i d e n t i f i c a t i o n after logging. 18 More recently, Kojima (1971) described the phytogeocoenoces of the Coastal Western Hemlock biogeoclimatic zone in Strathcona Pr o v i n c i a l Park, located in central Vancouver Island. Eight associations were recognized and analyzed s t a t i s t i c a l l y for correlations with environmental a t t r i b u t e s . It was concluded that moisture regime i s the most i n f l u e n t i a l factor c o n t r o l l i n g vegetation, given the same macroclimate and parent material, and that moisture regime is highly correlated with nutrient regime because of seepage water. Other studies of plant associations on Vancouver Island that are less related to the present study include Kuramoto (1965), Wade (1965), Cordes (1972) and Roehmer (1972). Several studies on the south coastal mainland of B.C. also pertain to t h i s thesis. Orloci (1961) c l a s s i f i e d forest types of the Coastal Western Hemlock biogeoclimatic zone between Howe Sound and P i t t Lake and investigated environmental relationships in these ecosystems (1964). Eis (1962) studied the influence of individual environmental factors as well as groups of factors on plant communities and tree productivity in t h i s same study area. Many factors were measured quanti t a t i v e l y and analyzed s t a t i s t i c a l l y . Eis concluded that: 1) topography was the primary factor influencing s o i l and water conditions within a macroclimatic region, 2) environmental features were more clo s e l y correlated with plant communities than s i t e index, and 3) most of the v a r i a b i l i t y between the communities studied was accounted for by s o i l and moisture regime, seepage water and s o i l permeability being the most important factors. Other pertinent studies in coastal B r i t i s h Columbia include 19 G r i f f i t h (1960), Lesko (1961), Klinka (1976) and Packee (1976). Krajina (1969) has compiled a l i s t of biogeocoenoses (plant associations and their corresponding s o i l s ) that occur in B r i t i s h Columbia from f i e l d studies and the work of his students. The d i s t r i b u t i o n of these types and the associated major tree species are related to moisture and nutrient regimes in two-dimensional grids for biogeoclimatic zones in B.C. 2.2. Studies of vegetation-environment in the P a c i f i c Northwest United States Results of c l a s s i f i c a t i o n s of forest communites in Oregon and Washington have been compiled by Franklin and Dyrness (1973). Descriptions of forest associations in the coast range have been made by Barrett (1962), C o r l i s s and Dyrness (1965), Bailey (1966), Waring (1969), Bailey and Hines (1971), Bailey and Poulton (1968), Meurisse and Youngberg (1971) and Hines (1971). Forest communities similar to those of the coast range have been c l a s s i f i e d in the western Cascades (Franklin 1966, Fonda and B l i s s 1969, Dyrness et a l . 1974, M i t c h e l l and Moir 1976). A l l of the studies followed the general approach of the habitat type c l a s s i f i c a t i o n scheme (Daubenmire 1952). P f i s t e r (1977) presents a review of habitat type c l a s s i f i c a t i o n s for the western United States. A c l a s s i f i c a t i o n has recently been completed for Mt. Rainier National Park (Franklin et a l . 1979), and one i s in progress for Olympic National Park. Recent studies have related habitat types to environmental gradients in southwest coastal Oregon by measuring several plant responses ( i . e . water stress, stomatal behavior, f o l i a r 20 nu t r i t i o n and phenology) and r e l a t i n g these to environmental variables to obtain plant response indices for s o i l moisture, temperature, transpiration and s o i l f e r t i l i t y . A temperature-growth index (Waring and Cleary 1967) and plant moisture stress during the growing-season measured by the Scholander pressure chamber (Scholander e_t a_l. 1965) were used to define the two axes of an environmental g r i d , on which the d i s t r i b u t i o n s of species and communities were plotted (Waring et a l . 1972). Certain indicator species (a t o t a l of 47) were then correlated with plant response indices to predict environment for s i t e s not d i r e c t l y measured. When tested on s i t e s where temperature and moisture had been measured, the indicator species estimates successfully predicted plant moisture stress and temperature growth index. This same approach was used by Zobel et a_l. (1976) in the western Cascades of Oregon. In addition, a Master Environmental Index (MEI) and S o i l P r o f i l e Index (SPI) were developed to show the r e l a t i o n s h i p of communities to complex gradients. The SPI estimated moisture status by assigning weights to each plot for s o i l c h a r a c t e r i s t i c s such as texture, stoniness and rooting depth. The MEI added a factor for topography to SPI. These indices were plotted against elevation to obtain two-dimensional charts of the d i s t r i b u t i o n of reference stands. Results showed measurement of gradients to be superior, though the indices did s a t i s f a c t o r i l y show moisture relationships between stands. Ordination was used by Dyrness et a l . (1974) to help c l a s s i f y forests in the aforementioned study area, and to show the r e l a t i o n s h i p between climax associations and environmental 21 features. The technique that they used (SIMORD) i s e s s e n t i a l l y a polar ordination (Dick-Peddie and Moir 1970). The d i s t r i b u t i o n of habitat types was described in terms of complex moisture and temperature gradients. Polar ordination was also used in the c l a s s i f i c a t i o n of habitat types in Montana ( P f i s t e r and Arno 1980) . Other work in this region comparing measured environmental data or environmental indices to vegetation composition and d i s t r i b u t i o n includes Waring and Major (1964), G r i f f i n (1967), Whittaker (1960) and numerous unpublished M.S. and Ph.D. theses (Franklin and Dyrness 1973). The autecological c h a r a c t e r i s t i c s of individual tree species in the P a c i f i c Northwest are also pertinent to this thesis and have been the subject of numerous studies; a comprehensive l i t e r a t u r e review summarizing t h i s information has been prepared recently by Minore (1979). 22 CHAPTER 3 STUDY AREA 3.1. Location and physiography The study area is located along the east coast of Vancouver Island between Campbell River (50°01'N, 125°18'W) and Ladysmith (49°00'N, 123°50'W), a distance of 150 km (Figure 1). Specific study s i t e s were located at elevations between sea le v e l and 1100 m, ranging from the coast up to 19 km inland. The study area l i e s within two physiographic subdivisions: the Nanaimo Lowland and the Vancouver Island Ranges (Holland 1964). The former i s a narrow s t r i p of coastal p l a i n below 610 metres in ele v a t i o n 6 which ri s e s gradually to the slopes of the Vancouver Island Ranges. The mountains p a r a l l e l the coast from southeast to northwest, and are dissected by several broad U-shaped v a l l e y s . The highest peak within the study area is Mt. Arrowsmith (1817 m). There are eight p r i n c i p a l r i v e r drainages in the study area (north to south): Oyster, Tsolum, Tsable, Qualicum, Cameron, Englishman, China and Nanaimo. A l l flow into the S t r a i t of Georgia except China Creek, which enters the Alberni Inlet. Three large lakes, Comox, Cameron and Home, also l i e within the study area. Cowichan Lake and the Chemainus River form the southern boundary of the study area. 'Fyles (1963a) records t h i s boundary as 250 metres. 23 F i g u r e 1. Study area and l o c a t i o n of c l i m a t o l o g i c a l s t a t i o n s . • A E S -30 year ...sciaC-- I cWfc ta i 1 24 3.2. Bedrock geology Most of the Nanaimo Lowland i s underlain by sedimentary rocks of the Nanaimo group. These are of Upper Cretaceous o r i g i n and consist of sandstone, shale, carbonaceous shale, conglomerate, greywacke and coal (Northcote 1973). The r o l l i n g topography i s comprised of low ridges underlain by sandstone and conglomerate, with narrow valleys formed by the erosion of less resistant, shales (Holland 1964). The Vancouver Island mountains are composed' of pre-Cretaceous faulted and folded sedimentary rocks and volcanic rocks, with frequent g r a n i t i c batholiths. The sedimentary rocks consist of limestone and more resistant chert, a r g i l l i t e , tuff and greywacke (Day et a_l. 1959; Muller and Carson 1969). The present geology of the island i s largely the result of three cycles of volcanism as described by Northcote (1973). Detailed descriptions of the geologic events (accounting for the heterogeneity of the present formations) and l o c a l maps are given by Muller (1965, 1971), Muller and Carlson (1969), Muller and Jeletsky (1970), Northcote and Muller (1972), Fyles (1955), Mathews, (1947), and Stevenson (1945) and are summarized by Packee (1976). 3.3. S u r f i c i a l geology The s u r f i c i a l geology of eastern Vancouver Island is predominantly the result of the most recent (Fraser) g l a c i a t i o n , which ended ten to twelve thousand years ago, and p o s t - g l a c i a l events. The predominant materials are g l a c i a l t i l l s , g lacio-f l u v i a l deposits and marine sediments. The sequence of events 25 res u l t i n g in the present mosaic of deposits may be summarized as follows, after Day et al..(1959): 1. Prior to the Fraser g l a c i a t i o n , the eastern lowlands were buried by hundreds of feet of r i v e r deposits, predominantly gravels and sands. 2. With the advance of the Fraser g l a c i a t i o n , e x i s t i n g s u r f i c i a l deposits were eroded and g l a c i a l t i l l of varying thickness (1 to 31 metres) was deposited in the coastal lowland, valley f l o o r s and lower mountain slopes. 3. At the height of g l a c i a t i o n , the Cordilleran ice sheet moved across the island in a southerly d i r e c t i o n , pushing westward through lower valleys such as the Cameron and Nanaimo. The ice sheet modified a l l topography below approximately 1220 m (Holland 1964), and depressed the land surface s i g n i f i c a n t l y . 4. Upon retreat of the g l a c i e r , the melting ice deposited g l a c i o f l u v i a l materials. The sea then re-entered the Georgia S t r a i t , eroded lowlands and l e f t marine deposits, and the land surface subsequently rebounded leaving marine deposits at various elevations above sea l e v e l , dependent upon the thickness of the former i c e . 7 5. Since g l a c i a t i o n , f l u v i a l , c o l l u v i a l and organic deposits have formed. 'Marine influence is found at 91 m in the V i c t o r i a and Alberni areas, 122 m at Nanaimo, 152 m at Qualicum and 183 m at Campbell River (Day et a_l. 1959). Other estimates d i f f e r (Heusser 1960; Holland 19647. 26 The g l a c i a l t i l l or morainal deposits vary considerably in thickness and texture. The deepest deposits are on the coastal lowland, with thinner materials in mountainous areas. Texture ranges from very gravelly sandy ablation t i l l to compact boulder clay basal t i l l . Surface t i l l s of the Nanaimo Lowland are predominantly gravelly, s i l t y sands. Fyles (1963a) c a l l s these materials "Vashon and sub-Vashon t i l l " , r e f e r r i n g to their apparent o r i g i n in the Vashon stade of the Fraser g l a c i a t i o n . In the glacially-scoured divide between the Alberni basin and Cameron val l e y , much exposed bedrock i s present with overlying t i l l less than 10 cm thick (Packee 1976). G l a c i o f l u v i a l deposits of primarily sands and gravels occur as fans and terraces in the knob and kettle topography of the higher uplands of the Nanaimo Lowland, the most extensive of which are found just above the highest marine deposits where valleys led to former sea levels (Day et_ a_l. 1959). Marine and glaciomarine deposits confined to the coastal lowland vary from 0.3-1.5 m in thickness on slopes to 1.5-9.0 m in f l a t s and depressions, found mostly over compact t i l l or rock (Day et a l . 1959). Texture of these deposits varies from massive stony clays to s i l t s and clays, depending upon the degree of wave action and mixture with g l a c i a l material during their format ion. The remainder of the s u r f i c i a l deposits on the east coast of Vancouver Island, including a l l u v i a l , c o l l u v i a l , lacustrine and organic materials, occupy a smaller proportion of the landscape.- Recent a l l u v i a l deposits in rive r v alleys consist of sands and gravels, often with a thin overlying veneer of f i n e r -27 textured mineral or organic deposits. C o l l u v i a l debris i s found on steep valley slopes, and i s composed of either bedrock fragments (talus) or slumping morainal material. Some buried lacustrine s i l t s and beach sediments have been found upstream from Cameron Lake beneath recent f l u v i a l deposits (Packee 1976). Organic accumulations occur sporadically in swampy areas throughout the study area (Maas 1972). A chronology and description of the s u r f i c i a l geology in the south coast region of B r i t i s h Columbia i s given by Armstrong et a l . (1965) and Packee (1976). Maps and descriptions for eastern Vancouver Island are given in several publications by Fyles (1959, 1960, 1963a', 1963b). 3.4. S o i l s Several broad s o i l landscapes have been described for the province of B r i t i s h Columbia. S o i l landscapes are defined at the s o i l great group l e v e l in the Canadian System of S o i l C l a s s i f i c a t i o n as "the t o t a l ecosystem with which a s o i l i s associated" (Valentine et a_l. 1978). The study area l i e s within two of these units: the Humo-Ferric Podzol and Dystric Brunisol landscapes. The l a t t e r occurs only to a limited extent at low elevations in the extreme southeastern part of the study area. The podzols of Vancouver Island are derived predominantly from morainal materials. Lewis (1976) studied these s o i l s and found that the Vancouver Island "model" of the podzol was somewhat d i f f e r e n t from the c l a s s i c model. The c l a s s i c model has an LFH surface organic layer; a l i g h t gray, s i l i c a - r i c h Ae horizon formed by eluviati o n (removal) of bases, organic matter, 28 iron and aluminum; and a reddish brown B horizon enriched with organic matter, iron and aluminum, changing gradually to a more yellowish C horizon (Valentine and Lavkulich 1978). In contrast, podzols on Vancouver Island generally lack a well-developed Ae horizon. The addition of organic matter and weathering of iron and aluminum in upper horizons result in no net depletion of materials despite heavy leaching. An Ae i s also absent from s o i l s derived from ba s a l t i c and andesitic parent materials which have no s i l i c a to leave behind after weathering. Vancouver Island podzols t y p i c a l l y have large accumulations of sesquioxides (Fe and Al) throughout a thick B horizon. Organic matter accumulation in this horizon i s also very s i g n i f i c a n t . Horizons are commonly patchy due to frequent churning of s o i l from root action, windthrow, and downslope movement in the mountainous forest environment (Lewis 1976). Podzols at elevations greater than 900 m within the study area generally receive enough p r e c i p i t a t i o n input to develop into Ferro-Humic podzols (Valentine et a l . 1978), which predominate on the north and west coasts of Vancouver Island. In addition to the podzolic s o i l s , several other s o i l orders are represented. Eluviated and Humic Gleysols occupy water-saturated areas such as swamps, floodplains, and l o c a l depressions, and are usually associated with marine clays, lacustrine s i l t s and clays, and fine-textured t i l l s . Regosols are found on younger parent materials, mainly c o l l u v i a l and a l l u v i a l deposits. Luvisols are most commonly found on marine and finer-textured f l u v i a l landforms. Some Dystric Brunisols occur in the coastal plain south of Nanaimo. Chernozems are 29 rare, occupying only 0.1%.of the Nanaimo Lowland and Alberni Basin (Day et a l . 1959), and are mostly in a g r i c u l t u r a l production or housing development. Occasional organic s o i l s may be found in low-lying swamps and bogs. A compendium of the s o i l s of Vancouver Island compiled by the B.C. Ministry of Forests from previous studies (Keser and St. Pierre 1973) gives descriptions of s o i l s found within the study area. Day e_t a_l. (1959) described s o i l s suited to agriculture on eastern Vancouver Island. Several of the vegetation studies discussed in the previous chapter described s o i l s for parts of central and east coastal Vancouver Island. For a review of s o i l s in the area as related to the USDA s o i l c l a s s i f i c a t i o n system, the reader is referred to Packee (1976). 3.5. Climate The climate of the study area can be described according to Koppen's scheme under three types: Cfb, Csb and Dfc. The Cfb Marine West Coast type (Trewartha 1968) is t y p i c a l of middle elevations (300-900 m). It i s characterized by mild temperatures and abundant r a i n f a l l , with no d i s t i n c t dry season. The west coast of Vancouver Island i s also a Cfb climate, yet the east coast Cfb has greater temperature extremes, more sunshine hours, and less p r e c i p i t a t i o n due to the rain shadow eff e c t of the Vancouver Island Ranges. This effect i s most pronounced in the Oyster River valley south of Campbell River, being d i r e c t l y east of the highest peaks on the island in Strathcona Provincial Park. Table II gives Atmospheric Environment Service (AES) 30-year Normal climatic data for several stations within the study Table I I . Selected 30 year normal (1941-1970) climatic data for the study area (Atmospheric Environment Service 1973). Station ELEV. MEAN DAILY TEMP. (°C) MEAN DAILY MAX. . TEMP (°C) MEAN DAILY MIN . TEMP (m) JAN. JULY ANNUAL JAN. JULY ANNUAL JAN. JULY ANNUAL ALBERNI LUPSI CUPSI . 9 1.2 17.8 9.4 3.0 24.6 14.2 -1.5 11.0 4.7 CAMPBELL RIVER 30 1.3 17.4 8.9 3.9 23.5 13.2 -1.3 11.3 4.7 COMOX AIRPORT 24 2.1 17.3 9.3 4.9 22.8 13.4 -0.8 11.9 5.2 COWICHAN LAKE FOR. 177 1.0 17.4 9.0 3.6 24.3 • 13.7 -1.5 10.5 4.3 NANAIMO 8 2.6 17.7 9.9 5.5 23.4 14.3 -0.3 12.2 5.4 MEAN RAINFALL (cm) MEAN SNOWFALL (cm) MEAN TOTAL PRECIPITATION JAN JULY ANNUAL JAN. APRIL ANNUAL JAN. JULY ANNUAL ALBERNI LUPSI CUPSI 9 25.3 2.6 174.8 40.6 Tr 94.7 29.4 2.6 184.2 CAMPBELL RIVER APT. 30 18.3 3.9 143.6 43.7 0.5 104.1 22.7 3.9 153.8 COMOX AIRPORT 24 15.3 2.a 110.1 43.2 1.3 106.2 19.6 2.8 120.7 COWICHAN LAXE FOR. 177 26.7 3.2 194.2 74.4 0.5 179.8 34.1 3.2 212.1 NANAIMO 8 13.5 2.5 101.1 28.4 0.3 69.9 16.7 2.5 108.5 31 area. Data for Campbell River t y p i f i e s the Cfb cl i m a t i c type in the study area. The Csb Mediterranean Subhumid type is characterized by mild temperatures and less abundant r a i n f a l l than the Cfb, with a summer drought period. It is ty p i c a l of lower elevations (below 300 m) on the central and southeastern coast of Vancouver Island. Here the rainshadow effect of the Vancouver Island Ranges and Olympic Mountains is considerable. Climatic data for Nanaimo is c h a r a c t e r i s t i c of thi s type. Higher elevations (above 900 m) are c l a s s i f i e d as the Dfc Humid Continental type. Cool short summers, abundant p r e c i p i t a t i o n and heavy winter snowfall are t y p i c a l . Some short-term Resource Analysis Branch climatic stations occur within t h i s type, but no AES 30-year records exist for thi s type on Vancouver Island. The closest long-term station at high elevation i s on Grouse Mountain near Vancouver. The island is influenced by the Aleutian low-pressure system in winter and the Hawaiian high-pressure system in summer, creating prevailing winds from the southeast and northwest, respectively (Chapman 1952). The Vancouver Island mountains modify the eastward flow of moist P a c i f i c a i r . The rainshadow effect of this range varies, depending upon the height of the mountains and the orientation of v a l l y systems through which moist a i r may flow. For example, the Alberni i n l e t extends the wet, west coast climate further inland by channeling prevailing winds. Topography and prevailing winds can, therefore, create standing cloud patterns that may s i g n i f i c a n t l y a f f e c t the long-term climate in some areas. E f f e c t s of cloud 32 patterns have been correlated with insect outbreaks • on southeastern Vancouver Island (Wellington 1965). Krajina (1969) has correlated the climatic types of Koppen with biogeoclimatic zones. Packee (1976) takes exception to t h i s because of the broad geographical range and d i v e r s i t y of vegetation with which the types are associated on a worldwide scale. Nevertheless, biogeoclimatic zones and subzones can be correlated with certain climatic variables, i f not to the broadly-defined types of Koppen. 3.6. Vegetation The study area includes portions of the Coastal Douglas-f i r , Coastal Western Hemlock, and Mountain Hemlock biogeoclimatic zones, which have been mapped for a l l of Vancouver Island (MacMillan Bloedel Limited 1974). Several subzones and variants have been i d e n t i f i e d on the most recent biogeoclimatic map of Vancouver Island for the Nootka-Nanaimo sheet at a scale of 1:500,000 (Klinka et a l . 1979), which gives climatic and edaphic information as well as c h a r a c t e r i s t i c combinations of species for zones, subzones and variants. According to the habitat type system, the following six vegetation zones are represented in the study area: Douglas-fir (PSME), Douglas-fir-Western Hemlock (PSME-TSHE), Western Hemlock (TSHE), Amabilis Fir-Western Hemlock (ABAM-TSHE) Amabilis F i r -Mountain Hemlock (ABAM-TSME) and Mountain hemlock-Alpine F i r (TSME-ABLA). Nineteen vegetation series have been i d e n t i f i e d for the area (Appendix I ) . Habitat type descriptions have yet to be published for Vancouver Island. 33 CHAPTER 4 METHODS OF STUDY 4.1. F i e l d studies 4.1.1. Study strategy and reconnaissance F i e l d operations were based in Nanaimo and extended over a period of four months beginning in May, 1979. Three weeks were spent in reconnaissance of the study area to locate suitable areas for sampling. A e r i a l photographs and the exis t i n g forest cover maps of MacMillan Bloedel Limited were also used. A systematic approach to sampling was chosen in order to reduce subjective bias in locating samples. Elevational transects were established on portions of mountain .slopes supporting "old-growth" stands free of major disturbance. The objective of sampling was to c o l l e c t data from which the potential climax vegetation for these s i t e s could be inferred. In a few cases, second-growth stands of natural ( f i r e ) o r i g i n were sampled when suitable old-growth forests were unavailable. These stands were generally over 120 years old. Leave s t r i p s in the Cameron and South Nanaimo River valleys provided rare opportunities to sample old-growth forests from valley floor (400 m) to ridgetop (1100 m) on northeast and southwest aspects (Figures 2 and 3). Other transects of more limited elevational range were established in the China Creek and Oyster River valleys. Transects were sampled at regular intervals of approximately 80 m. Reconnaissance of these areas suggested that t h i s interval would be adequate to characterize F i g u r e 3 . Cameron River v a l l e y l o o k i n g southeast. 35 changes in vegetation with elevation while conforming to the time l i m i t a t i o n s and objectives of the study. The Englishman River, Tsable River, and Cameron Lake areas were also sampled at regular elevational i n t e r v a l s ; however, due to the discontinuous occurrence of old-growth stands in these vall e y s , transects had to be "constructed" from available stands, making elevation gaps in these areas inevitable. The remainder of the t o t a l of 96 plots were located in small remnant stands along the coast. 4.1.2. Plot selection and sampling Ci r c u l a r 500 m2 plots were used to sample a l l vascular vegetation strata within representative portions of stands at pre-determined elevations along each transect (Franklin e_t a l . 1979). In a few cases, larger plots (1000 m2) were used where necessary to characterize v a r i a t i o n ; smaller plots (250 m2) were used in a few extremely homogeneous stands. Choice of plot centre was made subjectively in an attempt to avoid areas of windthrow, rock outcroppings, streambanks, logging and road disturbance and other microsites. Boundaries of the plot were measured and marked with flagging to f a c i l i t a t e location of plots on return v i s i t s . Trees were t a l l i e d by species and canopy layer in 10 cm DBH (Diameter Breast Height) classes. Canopy layers follow those of Smith (1962), (Appendix II, Table A l ) . Trees less than 1.5 m t a l l were recorded as seedlings. Where seedling cover was sparse, a l l seedlings on the plot were t a l l i e d by species. Where common to abundant, four subplots of 1.5-m radius were established 6 m from plot centre in the four cardinal directions 36 (Franklin e_t a_l. 1979). Young trees less than 20 cm t a l l were not counted as seedlings because germinants on nurse logs frequently numbered in the hundreds per subplot. Estimates of canopy cover for the overstory and seedling layers were also recorded. A complete species l i s t of understory vascular plants was made after a thorough examination of the p l o t . Nomenclature follows Hitchcock and Cronquist (1973) (Appendix I I I ) . Many of the common bryophytes were also noted, following nomenclature of Lawton (1971) and Schofield (1979). For each species in each layer the percentage cover for the plot surface was estimated. Understory layers follow those of Brooke et al. 1970 (Appendix II, Table A2). Vigor, s o c i a b i l i t y and abundance were also estimated for a l l vascular plants following the scales of Peterson (1964), Shimwell (1971) and Tansley and Chipp (1926), respectively (Appendix II, Table A3). Bryophytes and vascular plants not i d e n t i f i e d to genus and species in the f i e l d were co l l e c t e d . In some cases, incomplete flowering prevented posit i v e i d e n t i f i c a t i o n to the species l e v e l , p a r t i c u l a r l y in the case of grasses and sedges under dense canopies. Representative specimens of most of the vascular plants found on plots and adjacent areas were deposited in the MacMillan Bloedel Limited and UBC Forest Ecology herbariums. Some common lichens were noted, but none were sampled. The average height of the tree strata was measured using a Suunto clinometer and tape. Breast height age of at least three dominant trees was determined from increment cores. Estimates were made where trees were too large to determine age from 37 increment cores. Where possible, ring counts from stumps in adjacent clearcuts were obtained. An estimate of tree canopy density was made using a mirror densiometer. Five readings from d i f f e r e n t locations on the plot were averaged. Aspect, elevation (from an altimetre and topographic maps), percent slope, slope position and configuration, and location (latitude and longitude) were also recorded. Plot location was marked on topographic and forest cover maps and a description of the topography and nearby landmarks was made. Observations were made on insect and disease occurrence, f i r e history, and w i l d l i f e use. Each plot was i d e n t i f i e d to tentative vegetation zone and series using a f i e l d key developed by Packee (1979). This dichotomous key assists i d e n t i f i c a t i o n of climax vegetation units at the zone and series l e v e l by assessing the reproductive success of tree species (Appendix I ) . In addition, a biogeoclimatic zone, subzone, and "edatopic g r i d " position were also estimated using the B.C. Ministry of Forests "Guide to species selection and prescribed burning in the Vancouver Forest D i s t r i c t " (Klinka 1977) for comparative purposes. S o i l was described and sampled on each plot from a p i t approximately one metre square and at least one metre deep, unless a r e l a t i v e l y impermeable layer of otherwise unconsolidated material or bedrock was reached. Unconsolidated material included g l a c i a l t i l l , marine or c o l l u v i a l deposits. F i e l d descriptions of a l l forest floor and mineral horizons included depth, thickness, boundary distinctness and form, structure, consistence (dry, moist, wet), coarse fragment 38 content (size, shape, and volume), root abundance and size , and s o i l texture. V e r t i c a l and horizontal drainage were estimated. Samples of each horizon were c o l l e c t e d for analysis. Samples were a i r - d r i e d after each f i e l d day and were examined for moist and dry color using a Munsell color chart. Some physical s o i l analyses were done for t h i s thesis, but no chemical analyses. These w i l l be completed in future investigations. 4.2. Data analysis 4.2.1. Ordination techniques 4.2.1.1. Choice of methods The most complete and well-documented package of computer programs for ordination that i s currently available is produced by the Ecology and Systematics group at Cornell University, Ithaca, New York. Two of the programs available from the Cornell Ecology Program series are ORDIFLEX (Gauch 1977) and DECORANA ( H i l l 1979). ORDIFLEX i s a FORTRAN program that performs four ordination techniques: weighted averaging, polar ordination (PO), p r i n c i p a l component analysis (PCA), and reciprocal averaging (RA). The program provides f l e x i b l e data input and output options, allows several ordinations with a single run, and provides for simple editing and transformation of data. Two ordination techniques were chosen for this study from ORDIFLEX: polar ordination and reciprocal averaging. Both are well-documented and tested, and have been found to produce results superior to other techniques 39 (Gauch et a l . 1977). DECORANA is a recent improvement upon reciprocal averaging that eliminates certain problems with e a r l i e r versions. It is written in FORTRAN and may be run i n t e r a c t i v e l y from a computer terminal. Tests at Cornell indicate that i t i s "the best general purpose ordination" (Gauch, in preface to H i l l 1979). The following sections describe the theory and general mathematical basis for these . methods. For a complete description of the algorithms and computer programs, the reader i s referred to the l i t e r a t u r e c i t e d . 4.2.1.2. Polar ordination Polar ordination (PO), also known as Bray-Curtis and Wisconsin PO, i s mathematically and conceptually simple. The user chooses two samples from a data set that are d i s s i m i l a r in terms of species composition and/or environmental features to serve as opposite poles or endpoints with which a l l other samples are to be compared. The choice of endpoints depends upon the variation the user wishes to analyze ( i . e . nutrient status, moisture, elevation, e t c . ) . Samples are mathematically compared with endpoints using one of several s i m i l a r i t y measures. Three commonly used measures are: percentage s i m i l a r i t y (Odum 1950), c o e f f i c i e n t of community (Sorensen 1948) and Euclidean distance. There are several modifications of each (Goodall 1973). The measure chosen for th i s study i s percentage distance (PD), the complement of percentage s i m i l a r i t y . PD i s defined as: 40 1 PDjk = 100 - 200 x f=j min ( S i j , Sik) i i < s i3 + s i R ) where: PDjk = percentage d i f f e r e n c e between samples j and k. S i j , Sik = percent cover of s p e c i e s i i n samples j and k. I = t o t a l number of s p e c i e s i n samples j and k. PD may be c a l c u l a t e d f o r a l l p o s s i b l e p a i r s of samples to choose endpoints with the g r e a t e s t d i s s i m i l a r i t y . The next step i s the computation of o r d i n a t i o n v a l u e s . The procedure may be i l l u s t r a t e d with a g r a p h i c a l r e p r e s e n t a t i o n (Gauch 1977): sample j endpoint 1 L endpoint 2 where: X o r d i n a t i o n d i s t a n c e L b a s e l i n e d i s t a n c e between endpoints Dl d i s t a n c e between j and endpoint 1 D2 d i s t a n c e between j and endpoint 2 E d i s t a n c e o f f the a x i s the computed PD values f o r sample " j " are Dl " and D2. The 41 c a l c u l a t i o n s can be written a l g e b r a i c a l l y as: X = L 2 + D l 2 - D22 / 2L and E = V D l 2 - X 2 Samples equally d i s s i m i l a r to both endpoints w i l l ordinate in the centre of the axis. The distance of a sample from the axis (E) i s inversely related to the va r i a t i o n accounted for by t h i s axis. Second and subsequent axes can be defined by choosing additional endpoint p a i r s . A common approach i s to choose samples from the middle of the f i r s t axis that are most d i s s i m i l a r . "E" values can be useful for t h i s . 4.2.1.3. Reciprocal averaging Reciprocal averaging or "correspondence analysis" as described by Benzecri (1969) and Guihochet (1973) in France i s similar in nature to p r i n c i p a l component analysis. A complete description of the algorithms involved i s quite lengthy ( H i l l 1973), but the conceptual basis i s simple. The procedure begins by assigning a r b i t r a r y species scores that are used to obtain sample scores. Each sample score i s then calculated as the mean value of the scores for the species that occur • in i t . New species scores are then computed by averaging the scores for the samples in which that species occurs. This procedure i s repeated through successive i t e r a t i o n s u n t i l both species and sample scores converge to a unique solution. The solution i s not affected by the choice of i n i t i a l species scores, except in the number of it e r a t i o n s required to arr i v e at stable values. E s s e n t i a l l y , this i s a two-way weighted averaging computation. 42 Additional axes are derived using algorithms described by H i l l (1973). Reciprocal averaging produces both a species and a sample ordination. The method is extended to u t i l i z e quantitative data by weighting the sample scores proportional to the abundance of each species. In practice, RA axes scores are calculated from a species cross products matrix using the mathematical techniques of eigenanalysis; therefore, RA is computationally similar to PCA, though because of a di f f e r e n t data standardization i t produces more desirable results for vegetation analysis (Gauch e_t a l . 1977). For a complete treatment of the matrix algebra involved, the reader i s referred to Orloci (1978). 4.2.1.4. DECORANA Reciprocal averaging has two problems that detract from the interpretation of re s u l t s . The f i r s t stems from the method of cal c u l a t i o n of second and subsequent axes. Calculations ensure that higher axes are uncorrelated with the f i r s t , but not necessarily independent. This causes a frequent arching or "horseshoe" effect in graphs of the f i r s t two ordination axes resulting from a quadratic dependency of the second axis on the f i r s t axis. A second problem is the compression of scale at the axis ends. In other words, two samples with the same d i s s i m i l a r i t y w i l l ordinate closer together i f at the end of the axis than i f at the middle. To correct for these two problems, a program named DECORANA (DEtrended CORrespondence ANAlysis) has been written by H i l l (1979). The algorithms for deriving higher axes demand that they 43 not only are uncorrelated but that no systematic relationship of any kind exists with the f i r s t axis. This i s accomplished in a "detrending" procedure. The f i r s t DECORANA axis is i d e n t i c a l with RA. The second axis is detrended by dividing the f i r s t axis into segments and readjusting the sample scores in each segment to have zero mean. Detrended sample scores are used to calculate new species scores as in RA. The usual RA i t e r a t i o n procedure is followed u n t i l scores s t a b i l i z e . A similar detrending procedure is applied to each additional axis with respect to the preceeding one. The systematic arch effect i s thereby avoided, although a less s i g n i f i c a n t analogous effect does occur with the t h i r d and higher axes ( H i l l 1979). The second problem is minimized by rescaling the axes. This is accomplished by c a l c u l a t i n g a l o c a l mean standard deviation for species scores at intervals along the axis and using t h i s value to rescale the species so that within-sample standard deviation along the axis i s approximately unity. This assumes a f a i r l y even turnover of species along the gradient. The procedure was tested with simulated data with a regular structure and was found to preserve the known pattern. A recent a r t i c l e by H i l l and Gauch (1980) presents a more detailed description of the technique along with results of tests using both simulated and f i e l d data. 4.2.1.5. Application of ordination methods Ordinations were f i r s t performed on a subset of the t o t a l data set consisting of 31 plots from transects in the Cameron and South Nanaimo River va l l e y s . Reciprocal averaging and 4 4 DECORANA were used for a f i r s t ordination of plots and species. Several data transformations and species editings were performed to test their e f f e c t s , and to gain f a m i l i a r i t y with the properties of the data. Separate ordinations for understory and overstory species were performed, as well as for a l l species combined. Understory ordinations included only vascular plants because of incomplete c o l l e c t i o n of bryophyte data. In addition, because many mosses and liverworts are d i s t r i b u t e d in response to microsite conditions (e.g. base of trees, rocky seepage slopes) and few respond to broader s i t e features, i t was f e l t that their inclusion in the analysis might obscure the broader scale patterns of interest in thi s study. Ordinations of tree species using both calculated basal area values and percent cover estimates were compared in order to determine the value of c o l l e c t i n g mensurational data for thi s purpose. Basal area tree data was combined with percent cover estimates of understory species for complete species ordinations by converting both to a 10 point scale. This placed a l l data within the same range of values. Once RA and DECORANA ordinations were completed, suitable endpoints were chosen for polar ordinations. Several combinations of species and sample endpoints were tested to investigate d i f f e r e n t environmental gradients. Following each ordination, environmental data ( i . e . WSI, AWSC, potential solar radiation, temperature, elevation) were plotted on the ordination graphs in an attempt to reveal patterns and to correlate axes with environmental gradients. Exceptions to general trends were investigated for causal factors. Vegetation zone, vegetation series and habitat type 45 d e s i g n a t i o n s were a l s o p l o t t e d on o r d i n a t i o n diagrams to analyze the r e l a t i o n s h i p between these c l a s s i f i c a t i o n u n i t s . Assessment of v e g e t a t i o n and environmental r e l a t i o n s h i p s f o l l o w i n g each o r d i n a t i o n suggested p o s s i b l e f u r t h e r o r d i n a t i o n s . A s e r i e s of o r d i n a t i o n s was performed f o r both the e n t i r e data set and f o r data for each i n d i v i d u a l zone f o l l o w i n g t h i s same procedure. 4.2.2. Tabular a n a l y s i s technique The B.C. M i n i s t r y of F o r e s t s uses the t r a d i t i o n a l Braun-Blanquet (1932) t a b u l a r comparison technique to i d e n t i f y v e g e t a t i o n u n i t s . T h i s approach i n v o l v e s the p r e p a r a t i o n of s u c c e s s i v e l y r e f i n e d a s s o c i a t i o n t a b l e s to group samples ( r e l e v e s ) on the ba s i s of s i m i l a r combinations of s p e c i e s . M u t u a l l y - e x c l u s i v e groups of s p e c i e s are sought from a rearrangement of samples and sp e c i e s ( i . e . columns and rows i n the t a b l e ) and are used to i d e n t i f y s i m i l a r a s s o c i a t i o n s . The method i s d e s c r i b e d in d e t a i l by Shimwell (1971). Because t h i s i s a time-consuming procedure when done manually, a computer program has been developed to a s s i s t s o r t i n g and p r e p a r a t i o n of f i n a l t a b l e s ( K l i n k a and Phelps 1979). The program does not a s s i s t i n i d e n t i f y i n g groupings i n any a n a l y t i c a l way, i t simply speeds up the manual s o r t i n g process and e l i m i n a t e s t r a n s c r i p t i o n e r r o r s r e s u l t i n g from hand-copying of t a b l e s . The program does, however, c a l c u l a t e a mean and range of species s i g n i f i c a n c e f o r a l l s p e c i e s and samples, computes s p e c i e s presence values (percent occurrence i n the 46 u n i t ) , and assigns species to constancy classes. The program allows user sorting of samples, but not of species. Species are arranged in decreasing order of presence and mean species sig n i f i c a n c e by strata. This feature is advantageous for characterizing units, but does not meet the objective of forming species groupings. The tables in Appendix VI were prepared using t h i s program. A second program, which accompanies the vegetation table program, produces environmental tables that summarize environmental data for each group of samples. Comparison of s i t e data with vegetation groupings helps in defining biogeocoenotic uni ts . The tabular comparison method was applied to the sample data following ordinations. The f i r s t axis ordination sample scores were used for the f i r s t arrangement of association tables. Tables were used for comparison of plot groupings i d e n t i f i e d from the ordinations, and for determination of c h a r a c t e r i s t i c s of vegetation associated with segments of the environmental gradients. 4.2.3. Correlation with environmental data 4.2.3.1. Climatic data Climate for each of the 96 plots was predicted from 30-year average (1941-1970) Atmospheric Environment Service (AES) c l i m a t i c stations, and from short-term (3-5 years) Resource Analysis Branch (RAB) stations for the f i v e growing-season months; (May through September) (Appendix IV, Table A4). Because 47 of differences in elevation and aspect between stations and plots, values of temperature, p r e c i p i t a t i o n and solar radiation had to be adjusted. Mean monthly temperature was adjusted using a May through September lapse rate of 1.66°C/304.8 m (2.98°F/1000 f t . ) determined from RAB transects on Mt. Cain, Mt. Puzzle and Mt. Crest on Vancouver Island (M. C. Coligado, pers. comm.). Plot temperatures were derived with the following formula: Tp = Ts + (Es - Ep) L where: Tp = plot adjusted temperature Ts = station temperature Ep = plot elevation Es = station elevation L = lapse rate Lapse rates d i f f e r for other months of the year and for minimum and maximum temperature determinations. The standard free surface lapse rate of 1.67°C/304.8 m (3°F/1000 f t . ) is confounded in mountainous t e r r a i n because of turbulence, u p l i f t , cold a i r drainage and other weather-topography interactions. This should be kept in mind in interpreting any estimates. Observations on Mt. Cain also showed a difference between mean monthly temperature values for north and south aspects (M. C. Coligado, pers. comm.). This difference was not used to adjust plot values since confounding factors made such an adjustment beyond the degree of accuracy possible for these estimates. Average monthly p r e c i p i t a t i o n for May through September was 48 predicted from a provisional formula derived from RAB transects (M. C. Coligado, pers. comm.): Pe = Psl + [ 11502.,7(Psl)" 1 • 2 4 ' ] [E] where: Pe = May-September p r e c i p i t a t i o n in mm for a given elevation Psl = May-September p r e c i p i t a t i o n in mm at sea l e v e l E = Elevation in hundreds of metres This equation describes an inversely proportional r e l a t i o n s h i p between p r e c i p i t a t i o n at sea le v e l and increase with elevation. In other words, in d r i e r areas the orographic e f f e c t of mountains i s greater (greater increase in p r e c i p i t a t i o n with elevation) than for areas of higher p r e c i p i t a t i o n at sea l e v e l . Solar radiation data are rare for the study area. The only station recording actual net dai l y short wave radiation i s Nanaimo Departure Bay (AES). Hours of bright sunshine are recorded at Alberni Lupsi Cupsi, Nanaimo Airport, Campbell River and Cowichan Lake Forestry. These data were converted to net solar radiation (cal/cm 2) using one of several formulae derived for d i f f e r e n t data inputs from a Douglas-fir forest at the UBC Research Forest, Haney B.C. (Black 1973). The equation applicable for the data available i s : Rn4=0.0338Ro(n/N)2+0.4895RO(n/N)-184(n/N)+0.206RO-46 where: Rn4 = net solar radiation (cal/cm 2/day) Ro = Radiation at the top of the atmosphere (available 49 from Smithsonian Meteorological Tables ( L i s t 1949) N = Daylength (from tables of Black (1979) calculated from Furnival et a_l. (1969) ) n = number of sunshine hours (from AES stations) An average albedo (degree of surface reflectance) of 0.12, calculated from a Douglas-fir stand, was used in the derivation of this formula. This was considered adequate for the qu a l i t a t i v e comparisons of the present study (T. A. Black, pers. comm.), though i t should be noted that lower actual albedo for species with darker foliage, such as Abies amabilis, causes this formula to underestimate net radiation for some stands. While the l o c a l sunshine data account for cloudiness, there i s no way to adjust t h i s value for individual plots where cloud and fog patterns may be e n t i r e l y d i f f e r e n t from the climatic stations. There is also some reason to doubt the v a l i d i t y of the data from Nanaimo Departure Bay, due to the effe c t of the nearby Harmac pulp m i l l (E. C. Packee, pers. comm.). This effect is probably not as s i g n i f i c a n t for the five summer months with pre v a i l i n g northwesterly winds as i t i s in winter when southeast winds p r e v a i l . The same effect (a nearby pulp m i l l ) may d i s t o r t sunshine data from Alberni Lupsi Cupsi. Potential annual d i r e c t solar radiation (cal/cm 2/yr) for each plot was estimated from tables by Buffo et a l . (1972) for various slopes and aspects from 0° to 60° N l a t i t u d e . These data allow a q u a l i t a t i v e comparison of solar radiation input for evaluation of aspect, slope and latitude differences for individual plots, since measured AES data could be used only to 50 characterize different sampling areas, not individual p l o t s . 4.2.3.2. Water Stress Index Moisture a v a i l a b i l i t y for plant growth on a s i t e - s p e c i f i c basis cannot be evaluated simply by comparing p r e c i p i t a t i o n input. Complex interactions of many topographic, edaphic, geologic and b i o t i c factors influence water relat i o n s h i p s . For this reason, some method of combining such influences into a single value for comparison of samples can be a valuable t o o l . The hygrotopes of Krajina (1969) are a q u a l i t a t i v e attempt at locating regions on a moisture gradient, but are inadequate for any comparative attempts that cross zone boundaries, because the absolute moisture status associated with each hygrotope d i f f e r s from zone to zone. Also, s i t e water regimes cannot necessarily be characterized adequately by a one-dimensional system (from dry to wet), because of differences in evapotranspiration, seasonal v a r i a t i o n , etc. The USDA (So i l Survey Staff 1975) and CSSC (Canada S o i l Survey Committee 1978) s o i l moisture regimes refer to a v a i l a b i l i t y of water during specified periods of the year, but the s o i l moisture subclasses (CSSC) and s o i l moisture regimes (USDA) are often too broad for a comparison of s i t e -s p e c i f i c moisture. Consequently, some other means of q u a l i t a t i v e comparison of s i t e moisture was sought which i s based upon quantitative data available for the s i t e . A method that attempts to evaluate s i t e moisture a v a i l a b i l i t y in this way has been developed by Ballard (1974) for forests in south coastal B.C. It is c a l l e d a Water Stress Index (WSI). The method uses a model that simulates water 51 balance by u t i l i z i n g l o c a l c l i m a t i c data and information that can be gained from a single v i s i t to the s i t e to predict potential water deficiency during the growing-season (May-September). The purpose of developing such an index was "to enable foresters to predict the r e l a t i v e severity of s i t e water d e f i c i e n c i e s and to compare d i f f e r e n t s i t e s in terms of potential drought problems" (Ballard 1974). The WSI index value is given in terms of the number of months, to the nearest half month, during which severe and moderate water deficiency may occasionally occur. For example, a WSI value of 2.0 + 1.5 would be interpreted to mean that severe water stress may occur at times during 2 months, with moderate stress during an additional 1.5 months for the 5 month period, May through September. Severe stress i s t a l l i e d whenever the model calculates s o i l water content at the end of a half-monthly period to be at the Permananent Wilting Point (PWP)8, which i s assumed to be near the l i m i t of available s o i l water content for most plants. Moderate stress is t a l l i e d for a half-month period i f the model calculates s o i l water content not reaching PWP, but f a l l i n g below half of the available water storage capacity, which i s defined as water held between f i e l d capacity (FC)' and PWP. The model predicts water depletion in the s o i l by simulating water balance from p r e c i p i t a t i o n , evapotranspiration, and s o i l water storage capacity at bimonthly i n t e r v a l s . Monthly r a i n f a l l i s assumed to f a l l in two equal amounts, at the 8Water held at a matric potential of -15 bars (-1.5 MPa). 'Water held at a matric potential of -0.1 bar (-10 kPa). 52 beginning and middle of the month. This was thought to be v a l i d for modeling purposes since two week summer drought periods are not uncommon for the south coastal area. A number of other modeling assumptions are made: 1. On May 1st, the s o i l i s charged to f i e l d capacity from spring p r e c i p i t a t i o n and/or snowmelt. 2. Water drains to f i e l d capacity immediately following p r e c i p i t a t i o n . 3. Water loss at s o i l water contents between FC and PWP occurs only by evapotranspiration. 4. Loss of water by evapotranspiration ceases when calculated water storage reaches PWP. 5. Only water storage in the rooting zone i s si g n i f i c a n t for stress c a l c u l a t i o n s . Evapotranspiration rate for each month i s calculated as (McNaughton 1974) : E = S/(S+Y) (Rn/L) cm/day where: S = the slope of the saturation vapor pressure curve (mbars/°C) Y = the psychrometric constant (mbars/°C) Rn = average d a i l y net radiation L = 590 cal/cm 3 (based on a 15°C approximate regional a i r temperature and inferred s o i l temperature, and l i q u i d water density of 1.0 g/cm3) The term "S/(S+Y)" varies with mean monthly a i r 53 temperature, and was determined from available tables (Campbell 1977) using derived temperature values for each p l o t . Net radiation was estimated as: Rn = 10.4 + 0.59 Rs cal/cm 2/day where Rs i s solar radiation (direct plus d i f f u s e ) 1 0 (Black, McNaughton and Tang 1973). Diffuse radiation i s assumed to be 0.15 Rs on l e v e l t e r r a i n . Direct radiation i s affected by slope and aspect; consequently, t h i s value i s adjusted for each s i t e using the r a t i o : Xa/Xo where: Xa = di r e c t radiation on the slope Xo = di r e c t radiation on l e v e l t e r r a i n The mean values of ranges in th i s r a t i o , calculated from tables by Buffo, Fritschen and Murphy (1972), were used to establish fiv e radiation factors, each assigned a Radiation Index l e t t e r (A through E) for easy determination of radiation adjustments for slope and aspect (Appendix IV, Table A5). Available water storage capacity (AWSC) for the rooting zone i s determined from: 1. Forest floor thickness 2. Rooting depth 3. Thickness of each mineral layer with a d i s t i n c t 1 "Where Rn was determined from sunshine hours as described in the previous section, Rs was calculated using t h i s relationship. Where Rs was available from measurements, Rn was calculated. 54 texture and/or coarse fragment content The e f f e c t i v e thickness of each layer i s calculated by adjusting for coarse fragments (> 2 mm). Texture and e f f e c t i v e thickness are used to obtain AWSC for each mineral layer. Storage capacity for organic layers i s also estimated from a table (Appendix IV, Table A6). Computer programs are available (de la Fuente 1977) that calculate WSI values from the necessary data input. Values may be calculated i n d i v i d u a l l y for each s i t e , or tables may be generated for a given area that give WSI values arranged according to t o t a l p r e c i p i t a t i o n , AWSC and Radiation Index. A set of such tables for south coastal B.C. based upon regional average data used for developing the WSI model accompanies the programs. In c a l c u l a t i n g WSI values for the samples, several adaptations of f i e l d data were made. Coarse fragment content was estimated in the f i e l d using four general classes: 1 = <20%, 2 = 20-50%, 3= 50-90%, 4 = >90% (Resource Analysis Branch 1976). Available Water Storage Capacity i s highly dependent upon coarse fragment content. Because of the wide range of these categories, i t i s possible to predict a wide range of AWSC from the f i e l d estimates. For instance, for a s i l t loam horizon 28 cm thick, e f f e c t i v e thickness using both extremes of category "3" would be either 14 cm or 2.8 cm. The corresponding AWSC would be 5 cm and 1 cm, respectively. The effect i s exaggerated with greater horizon thickness and finer textures. To account for t h i s problem in a consistent manner, i t was decided to use the lower 55 l i m i t of each c l a s s . In thi s way, any error in WSI calculations would be on the conservative s i d e — p r e d i c t i n g stress only in cases of greatest p r o b a b i l i t y . Another adjustment had to be made in the case of rooting depth. Pit depth varied around the one metre average. In many cases, roots were observed down to the bottom of the p i t . Where th i s occurred, a one metre rooting depth was assumed so that estimates of AWSC would not vary as a function of p i t depth. This would also y i e l d stress estimates on the conservative side. The WSI model assumes good drainage; consequently, i t cannot be applied to plots with water tables within the rooting zone during the growing-season. It also cannot account for seepage, which is an unavoidable problem with any moisture regime index. WSI values computed for plots with abundant seepage or evidence of high water tables were not considered v a l i d for comparison with other p l o t s . Other l i m i t a t i o n s of the index w i l l be discussed in subsequent chapters. 56 CHAPTER 5 RESULTS AND INTERPRETATION 5.1. Environmental calculations Temperature, p r e c i p i t a t i o n , potential solar radiation, s o i l available water storage capacity (AWSC) and Water Stress Index (WSI) values were determined for each sample (Appendix IV, Table A7). Average growing-season temperature ranged from 15.9° C at 90 m near Cowichan Lake to 7.8° C at 1350 m near China Creek. Average p r e c i p i t a t i o n for the growing-season ranged from 14.2 cm to 41.1 cm. Potential solar radiation varied from 88,000 cal/cm 2/yr on steep north-facing slopes to 225,000 cal/cm 2/yr on steep south-facing slopes. AWSC was also quite variable in a l l areas sampled. WSI values were highest and the frequency of moisture stress greatest for the PSME zone where as much as 5 months of severe stress was predicted. Stress was rarely predicted for plots higher than 900 m because of high p r e c i p i t a t i o n input. A problem arises in applying the Resource Analysis Branch p r e c i p i t a t i o n regression to the Oyster River area, where the closest p r e c i p i t a t i o n records near sea le v e l are for Campbell River (69 m). The Oyster River i s in the pronounced rainshadow of the Strathcona peaks, while Campbell River i s just outside of this influence. As a consequence, use of Campbell River data overestimates p r e c i p i t a t i o n for th i s area, p a r t i c u l a r l y at lower elevations. To make estimates more r e a l i s t i c , p r e c i p i t a t i o n for the Campbell River Airport (109 m) was used, even though higher in elevation, because i t is nearer the study plots and f a l l s 57 within the rainshadow area. It has 11.6 cm less May-September pr e c i p i t a t i o n than Campbell River, just 10 km north and 36 m lower in elevation. This situation i l l u s t r a t e s the caution with which any generalized derived climatic data should be used where complex topographic-moisture relationships e x i s t . Nevertheless, this approach has been used to derive c l i m a t i c data for characterization of upper elevation biogeoclimatic zones, since l i t t l e actual data are available (Klinka e_t a_l. 1979). To test the r e l i a b i l i t y of the p r e c i p i t a t i o n regression, p r e c i p i t a t i o n from the nearest sea l e v e l c l i m a t i c station was used to predict mean monthly values (May - September) for higher elevation RAB stations where measured values were known. The measured and predicted values are given in Table I I I . Results show that predictions had as much as a 25% error, but were usually within 6% of the measured values. Figure 4 shows the d i s t r i b u t i o n of samples in r e l a t i o n to p r e c i p i t a t i o n and temperature with sample location plotted on the diagram. The Oyster River samples are separated from the others because the rainshadow influence, combined with lower temperatures, makes them quite d i f f e r e n t from plots at similar elevations further south. Tsable River samples also appear separated because most are at low elevation and have higher p r e c i p i t a t i o n values than low elevation plots at more southerly l a t i t u d e s . Figure 5 plots solar radiation and temperature with vegetation zones i d e n t i f i e d . Samples in the upper right corner of the figure in the PSME and PSME-TSHE zones have the highest heat loadings during the summer months. A high proportion of 58 Table I I I . P r e c i p i t a t i o n estimates compared to measured p r e c i p i t a t i o n for selected RAB climatological stations. PRECIPITATION (cm) MAP CODE STATION MEASURED ESTIMATE % DIFFERENCE 5 Tsable River FP (235m) 21.0 26.4 25.7 6 2E2500 (762m) 34.9 33.1 5.4 7 2E1400 (427m) 27.9 28.9 3.6 8 Cathedral (248m) 21.5 23.9 11.2 8 + 10 Cathedral (248m) 0 > 4 Franklin (177m) 16 McKay (290m) 23.5 22.4 4.7 18 Lookout High (1113m) 37.2 38.0 2.2 average % difference 7.0% From RAB provisional formula (Coligado 1980) based on V.I. climate transects. 59 F i g u r e 4. P r e c i p i t a t i o n verses temperature f o r sample p l o t s . 15.of 14.0 13.Of 12.01-"HOT, DRY' s 10.0 9.0 8.0 W7 7 W76 E21 L75 L12 L0 9 E2S E l l T8 9 T88 X8 7 T90 T92 T91 T9 3 SAMPLE LOCATION T86 E15 T96 L78 G9 5 L10 E24 G28 G26 G i l T98 T97 T99 T18 Y02 Y01 Y00 Y80 Y79 Y04 Y84 Y85 Y03 Y83 E16 G27 E14 G20 G29 E13 N60 N61 G22 -„ N43 N 4 8 Q 1 5 N47 G34 China Creek = C a t h e d r a l Grove = Cameron V a l l e y = Cowichan Lake = Englishman R i v e r = L o c a l Nanaimo = Nanaimo V a l l e y = T s a b l e R i v e r = Oyster R i v e r ^ P r e c i p i t a t i o n and temperature are average growing season (May -September). N56 N51 Y81 V37 N55 V31 C72 G30 N59 N57 N52 V40 C71 V46 N58 C73 V33 E62 N53 Y82 V41 C64 C69 C70 V32 C68 7-?2.0 16.0 20.0 24.0 28.0 32.0 PRECIPITATION (cm) 36.0 C67 C44 C66 C65 "WET, COLD" 40.0 POTENTIAL ANNUAL SOLAR RADIATION (M c a l / c m 2 / y r ) o o < • • > o o o -4— o o o H — <3 • • • IP Or • • < • o • • • n - P • •>> t> O • • > > > o Q O < • > O D *• *• -H u ro ro oo oo *» *» 31 % S S S GO OO CO o e H fD t-3 CD 3 TJ fD i-( SD r+ C fD < fD H V) fD cn 'Tj O rt fD D rf H-&> cn O &) n t-i PJ a H-OJ rt H-O Ml O M cn 3 t) M fD TJ M O r+ cn 09 61 samples in these two zones were on f l a t or gentle slopes with average radiation input. A better d i s t r i b u t i o n of radiation environments were sampled in the other vegetation zones. Some of the ABAM-TSHE zone samples are mixed within the warmer portion of the temperature axis because the extrapolated temperature values for these plots (based on an elevation regression) was unable to account for cold a i r drainage in the valley bottom si t e s these samples represent. The predicted temperature values for these s i t e s is not a true r e f l e c t i o n of the temperature environment influencing the vegetation. In Figure 6, p r e c i p i t a t i o n and AWSC are the two axes with vegetation series plotted. From t h i s diagram, one can ide n t i f y the d r i e s t and wettest samples in the study area, neglecting seepage water and evapotranspiration influences. There is considerable v a r i a b i l i t y in these two properties within most series, p a r t i c u l a r l y in the TSHE serie s . With seepage not accounted for, i t i s impossible to assess moisture relationships between series on the basis of the AWSC axis. There is a general macroclimatic trend associated with the p r e c i p i t a t i o n axis that largely r e f l e c t s elevational differences, with some effect from latitude and rainshadow. 5.2. Ordination of the entire data set An i n i t i a l series of ordinations was performed with the objective of analyzing relationships between zones. Results of dif f e r e n t methods and data transformations were also compared. Figure 7 shows an ordination of a l l samples and species using DECORANA. The lengths of both axes are given to show the F i g u r e 6 . P r e c i p i t a t i o n v e r s e s A v a i l a b l e Water S t o r a g e C a p a c i t y f o r sample p l o t s . • VEGETATION SERIES • • • • • o o I A • • m • • Q A A 1 A* " m A • • " • D A • 0 A A A D ° v O • v O A Q ° Q v Q Q • Q Q o Q O PSME • ABGR A THPL A THPL-TSHE(PSME) • TSHE • TSHE-ABAM Q ABAM A ABAM-TSHE(THPL) V ABAM-TSHE(CHNO) T CHNO <0> ABAM-TSME o 1 2 16 20 24 28 32 36 40 44 AVERAGE GROWING SEASON PRECIPITATION (cm) Figure 7. DECORANA o r d i n a t i o n of a l l samples showing the d i s t r i b u t i o n of vegetation zones. 320 CO -H X (0 O O oo° o o o • A A 0 ° . A * o o r A A A o o o O A a O O H A > © O H A O o 0 < > " A A O ° O A B A A . B O A A ^ A B Q A O A A gg B ^ a • A O B A I B a axis 1 © PSME O PSME-TSHE B TSHE O A ABAM-TSHE Or O ABAM-TSME Co 64 r e l a t i v e v a r i a t i o n that they represent. Vegetation zones were separated along the f i r s t axis of variat i o n , with considerable overlap in the positioning of samples from lower zones. This f i r s t axis has been interpreted as an elevation gradient, which r e f l e c t s macroc1imatic temperature and p r e c i p i t a t i o n di f ferences. The sharpest separation occurs between the ABAM-TSHE zone and a group of samples consisting of the highest elevation ABAM-TSHE samples and those from the ABAM-TSME zone. This separation r e f l e c t s the introduction of several species at approximately 1000 m, above which snow p e r s i s t s u n t i l late May. Comparison of samples on opposite ends of the second axis reveals differences in topo-edaphic moisture. The greatest variation occurs within the PSME and PSME-TSHE zones. Sites on r i v e r terraces, poorly-drained depressions and lower slopes were separated from dry upper slopes and rocky outcrops. The spread of plots along the second axis is reduced with increasing elevation, suggesting that species variation within the ABAM-TSME zone i s only about one-third that of the PSME or PSME-TSHE zone. A DECORANA ordination was performed using only understory species. Results (not shown) were very similar to the f u l l species ordination. There was s l i g h t l y more clumping of plots in the PSME-TSHE and ABAM-TSME zones, resulting from the removal of tree species from the comparison. Tree d i s t r i b u t i o n tends to overlap d i f f e r e n t understory communities and diminish the o v e r a l l species differences between plots. The s i m i l a r i t y of these two ordinations, however, implies that zones may be 65 distinguished on the basis of c h a r a c t e r i s t i c combinations of understory species alone. A t h i r d sample ordination using tree species alone produced generally similar r e s u l t s , but with a higher degree of overlap between vegetation zones. The i n i t i a l scattergram i d e n t i f i e d ' o u t l i e r samples as f i r s t axis endpoints, which caused clumping of other samples in the centre of the axis. DECORANA, as with normal reciprocal averaging, i s sensitive to outliers—samples very unlike a l l others. In thi s case, one plot having only Pseudotsuga menziesii in the overstory and two plots having primarily Abies lasiocarpa overstories were separated as axis endpoints. Deletion of these plots produced an improved ordination, though less s a t i s f a c t o r y than either the f u l l species or understory ordinations. Polar ordination produced poor results for the complete data set because there i s such high "beta" (between sample) d i v e r s i t y . Because chosen endpoints had no species in common, and intermediate plots had few species in common with either endpoint, plots were clustered in the centre of the f i r s t axis. This e f f e c t was exaggerated in ordinations of tree species alone. 6 6 5.3. Description and analysis of the Pseudotsuga menziesii (PSME) and Pseudotsuga menziesi i - Tsuga heterophylla (PSME-TSHE) vegetation zones In order that the reader may more f u l l y interpret the presentation of the ordination results, the description of vegetation series and habitat types i s presented f i r s t for each zone, even though these c h a r a c t e r i s t i c s were compiled after the analysis. These two zones were combined in the analysis because of their similar plant communities and because of limited sampling in each zone. In the biogeoclimatic c l a s s i f i c a t i o n system, these two units are considered dry and wet subzones of the Coastal Douglas-fir biogeoclimatic zone. 5.3.1. Description of the vegetation series Four vegetation series were i d e n t i f i e d in the PSME and PSME-TSHE zones. Several others are given by Packee (Appendix I ) . The PSME vegetation series designates an area where Douglas-fir is the potential climax tree species ( i . e . i t is able to reproduce successfully under i t s own canopy and is more abundant than a l l other tree species). The PSME series i s the zonal series for the PSME zone. In the PSME-TSHE zone the PSME series occurs commonly on south aspects, droughty s o i l s or rock outcrops. This series represents the driest conditions for tree growth sampled, though d r i e r series exist in these two zones. 67 The ABGR series occurs where grand f i r i s the most successfully reproducing conifer. Red cedar may also be present as a co-climax species. This series i s t y p i c a l l y found on river terraces, in sandy or gravelly alluvium and along the coastal plai n in coarse-textured t i l l . It i s common in both the PSME and PSME-TSHE zone. The THPL-TSHE(PSME) series i s defined as an area where red cedar and western hemlock are reproducing successfully. Douglas-f i r reproduction i s only marginally successful. This series represents the zonal overstory association of the PSME-TSHE zone. In the PSME zone, th i s series i s usually found on northerly aspects, river f l a t s , or lower slope positions where there i s groundwater influence but no imperfect drainage. The THPL series i d e n t i f i e s areas where red cedar i s the potential climax tree species. In these areas, red cedar i s usually the sole regenerating species with occasional grand f i r or western hemlock but rarely Douglas-fir. Red alder i s usually common. Areas in the PSME and PSME-TSHE zones with a high water table and/or poorly-drained s o i l support this edaphic climax. 5.3.2. Description of habitat types Site c h a r a c t e r i s t i c s for samples in each habitat type of the PSME and PSME-TSHE zones are presented in Appendix V, Tables A9 - A13. Association tables for each type are given in Appendix VI, Tables A24 - A28. Table IV summarizes the vegetation c h a r a c t e r i s t i c s for each habitat type in these two zones. To simplify the naming of the PSME/GASH-BENE and PSME/HODI/POMU habitat types, Douglas-fir was the only tree species used, even 68 Table iv. Summary of major species for habitat types i n the PSME and PSME-TSHE veaetation zones PSME/ ARME/ GASH 1 PSME/ PSME/ ABGR/ TREE LAYER Abizi giancLU Ace* macAophylZum AlnuA faibnja Atbutui mznzizAii P-ccea AirfchzniiA PinuA coyitonXa PoputuM tni.chocan.pa P&zudot&uga mznzizi>Li Thuja ptlcata Tiuga hzt.zn.ophylla SHRUB LAYER EzAbzitt, nzAvoia Gautthznla. iliatton HolodZicui diicolon. P.06a gymnocaipa Rabm, ipzctab-itU Pubui uKiinui SymphoKi.can.poA atbui, Vacciyrfjum paAv-L^otium HERB LAYER Ackly-i ZnJLpkyXJLa AdznocauZon bicolon, Adiavrfwn pe.dcrf.um AAZYIOJUO. macnophyUUx AthynMam fattix.-fazmina Blzchnum ipicant EKomui vuZgaAti Campanula icouteA-i Cklmapkita umbztlata Vn.yoptz>vL$ auifiiaca Vzitaca occA-dzntaJUA GaZium t/u.{,lon.um Lactuca muKaSLLi, Linnaza bon.zatu> Ly&ickitum amzfLlcanum Polystichum munitum PtzAldtuni aquitinum StAzptopui, amplzXslfaoLuiA Tiaxzlla ZacinLata Tianztla. fu^otlata TnlzntaCU, lati^otia TfUZZiwn ovatum VznjxtAum vi/Udz PC MC IV 7.0 I 2.4 V 6.9 V V IV IV IV III 3.1 7.1 2.6 4.1 2.0 1.9 II 1.6 III 2.7 IV 5.6. I l l 1.0 II 2.6 3.3 + .2 II +.0 I +.2 III 1.0 IV 2.0 THPL/ Presence Class: 2 Mean Coverage: I = 1-20%; II = 21-40%; trace; 1 = 0.1-1.0%; 2 GASH/ HODI/ POMU LYAM , BENE POMU PC MC PC MC PC MC PC MC III 4.5 I 1.5 IV 3.2 II 1.7 III 3.0 III 4.4 II 2.1 I 1.4 I 1.5 V 4.9 IV 4.0 I + .7 I 1.0 IV 5.1 V 6.3 V 8.7 IV 5.9 IV 5.1 IV 5.2 III 1.8 III 2.4 IV 5.3 V 5.0 III 4.3 V 5.6 V 5.2 V 5.3 IV 3.3 V • 5.0 V 7.4 II 5.5 II 1.3 II 4.1 II 1.0 V 4.0 I + .0 II 3.2 IV 4.2 I +'.1 II + .8 III + .5 IV 3.5 III 1.1 I •1.5 III 1.4 II + .6 I + .0 I 1.5 I + .2 V 3.3 IV 3.8 V 3.0 V 3.0 IV 4.7 V 3.7 V 3.0 II + .6 II 1.7 III 1.2 I + .0 I + .2 IV 1.1 III 1.6 I 1.0 III + .9 V 4.3 I + .0 IV 3.1 I + .1 II + .8 III 1.0 V 1.7 II + .6 I + .3 IV 2.3 IV 2.1 II + .6 I + .0 III 3.1 III 1.2 IV 1.0 III 1.9 III 2.5 I + .0 I + .0 V 3.3 V 3.5 V 8.2 V 8.2 V 4.5 V 3.1 I 1.5 II + .4 II 2.1 II + .8 IV 1.0 III 2.1 IV 1.3 V 1.0 I + .0 III 2.1 V 3.7 IV 1.0 V 4.0 IV 3.3 IV 2.1 I + .0 I + .2 IV 2.1 II + .0 II + .0 IV 1.2 = 4 1 " 60%; IV = 61-80%; V = 81-100% : the following classes • 1.1-2.2%; 3 = 2 .3-5.0%; 4 = 5. 1-10.0%; 0%; 7 = 33 .1-50. 0%; 8 = 50. 1-75 .0%; over 75.0% 69 though western hemlock should be included for stands in the PSME-TSHE zone. 5.3.2.1. Pseudotsuga menziesi i - Arbutus menziesi i / Gaultheria  shallon (PSME-ARME/GASH) habitat type This habitat type occurs exclusively within the PSME vegetation ser i e s . Stands belonging to thi s type were found on rocky outcrops and ridgetops near the coast in the Nanaimo and Englishman River areas. Sampled stands range in elevation from 120 to 540 m on a variety of slopes and aspects. Parent material is of c o l l u v i a l and morainal o r i g i n occurring as a thin veneer over bedrock, commonly sandstone. With the exception of one s i t e , s o i l rooting depth is less than 50 cm. S o i l textures range from loams to loamy sands with moderate to high coarse fragment content. The tree canopy is t y p i c a l l y sparse, having an average canopy density of 82% and the lowest average basal area (70 m2/ha) for the zone. The average height of the dominant trees i s 26 m for stands ranging in age from 185 to 275 years. The tree layer is dominated by Pseudotsuga menziesii with a s i g n i f i c a n t amount of Pinus contorta on some s i t e s . The presence of Arbutus  menziesii in the lower canopy layers distinguishes t h i s type (Figure 8). Characteristic shrubs include Lonicera c i l i o s a , Lonicera  hispidula, Symphoricarpos albus and Symphoricarpos mollis, though their dominance and constancy are not high. Gaultheria  shallon and Berberis nervosa are constant dominants. Other frequently occurring shrubs are Holodiscus di scolor, Rosa 70 F i g u r e 8. PAzudotAuga mz.nzi.zMix. --JAbuttU mznziz&iJ. / GaixLthztujOL khoJULon (PSME-ARME/GASH) h a b i t a t type. F i g u r e 9. ?&zudo£6uga mznziz&il I GauZtkeAia. shallon - BzAbzAAj, nzAvoia (PSME/GASH-BENE) h a b i t a t type. Range p o l e i n photos i s 1.5m t a l l with 1 dm graduations. 71 gymnocarpa and Rubus ursinus. The herbaceous layer i s dominated by grasses, the most abundant of which are Festuca occidentalis and Bromus vulgaris. Other common to frequently occurring herbs are Goodyera oblongi f o l i a , Tr i e n t a l i s l a t i f o l i a , Hierac ium  albiflorum, Campanula s c o u l e r i , Collomia heterophylla, Madia  sativa and Arenaria macrophylla. Mosses that occur frequently include Rhacomitr ium canescens, Pl'eurozium schreber i , and Rhytidiadelphus triquetrus. Understory species richness i s high r e l a t i v e to other types in the zone. The PSME-ARME/GASH habitat type appears equivalent to the Cladonia - Pelt igera - Mahonia 1 1 [Berber i s] nervosa  Gaulther ia shallon - Arbutus menziesi i - Pinus contorta -Pseudotsuga menziesii phytocoenosis 1 2 of Krajina (1969). It i s also c l o s e l y related to the s a l a l - l i c h e n association of Mueller-Dombois (1959) and McMinn (1957), though these types include habitats further from the coast with no Arbutus. Similar Douglas-fir - Arbutus forest types with Gaulther i a - and Festuca-dominated understories have been described for the San Juan Islands (Franklin and Dyrness 1973). A l l authors describe this type for rock outcrops, exposed ridges or south-facing slopes for low elevation, dry coastal areas. 1 1 Berberis nervosa Pursh i s now c a l l e d Mahonia nervosa (Pursh) NuttT (Taylor and MacBryde 1977). Berberis was retained to remain consistent with Hitchcock and Cronquist (1973) because Mahonia has not yet gained widespread acceptance. 1 2 For the f u l l l a t i n i z e d biogeocoenosis (#19), the reader i s referred to Krajina (1969). Subsequent reference to Krajina's biogeocoenoses w i l l use s i m i l a r l y abbreviated names. Obsolete species names w i l l be given in brackets. Comparison with these types is d i f f i c u l t because no descriptions are provided. 72 5.3.2.2. Pseudotsuga menziesi i / Gaultheria shallon - Berberis  nervosa (PSME/GASH-BENE) habitat type This habitat type was found at lower elevations (60 - 185-m) within the Englishman, Nanaimo and Tsable River drainages on a variety of g l a c i a l and glacio-marine parent materials. S o i l s range from s i l t loams to loamy sands. Several plots show evidence of imperfect drainage. Samples occur in the PSME, ABGR and THPL-TSHE(PSME) vegetation ser i e s . A l l stands are on f l a t or northerly aspects with gentle slopes, though t h i s i s not a distinguishing c h a r a c t e r i s t i c for the type. Sampled stands range in age from 120 to 460 years with an average canopy density of 93%, excluding one stand composed of scattered veterans. Height of the dominant trees averages 33 m. Stand volume i s s i g n i f i c a n t l y greater than In the previous habitat type and intermediate for the zone. The tree canopy i s dominated by Pseudotsuga menziesii with some Thuja p i i c a t a and Tsuga heterophylla common in lower layers. Douglas-fir regeneration is common in these stands. The shrub layer i s dominated by high but variable cover of Gaultheria shallon and Berberis nervosa in a l l samples (Figure 9). Vacc inium parvifolium i s also a constant shrub, but has small cover. The only constant herb species i s T r i e n t a l i s  l a t i f o l i a . Polyst ichum munitum, Pteridium aquilinum and Achlys  tr i phylla occur in most stands but vary considerably in importance. Moss cover of Stokesiella oregana and/or Hylocomium  splendens i s often greater than 50%. Few old-growth stands of t h i s habitat type were found. The old-growth stands from which samples were taken are small 73 remnants with some disturbance; consequently, the samples are not ideal for a good characterization of the habitat type. The PSME/GASH-BENE habitat type appears to include the [Eurhynchium] St o k e s i e l l a oregana - Mahonia nervosa [Berberis] -Gaultheria shallon - Pseudotsuga menziesi i phytocoenosis (biogeocoenosis #5) of Krajina (1969). It i s equivalent to the mesic Gaulther ia shallon - Pseudotsuga menziesi i ecosystem for the CDF wet subzone (Klinka 1977), l a t e r described as the Oregon grape - s a l a l - Douglas-fir zonal biogeocoenotic type by Klinka et a l . (1979). A Dou g l a s - f i r - s a l a l association was described by Krajina and Spilsbury (1953), McMinn (1957) and Mueller-Dombois (1959) for eastern Vancouver Island. Franklin and Dyrness (1973) mention a PSME/GASH type for the San Juan Islands. Very similar PSME/GASH-BENE habitats occur in the Tsuga heterophylla zone as described by Orloci's (1961) Gaultheria- Mahonia forest type and similar types from studies in Oregon and Washington, as summarized by Franklin and Dyrness (1973). 5.3.2.3. Pseudotsuga menziesi i / Holodiscus discolor / Polystichum munitum (PSME/HODI/POMU) habitat type This habitat type was found on steep c o l l u v i a l slopes and rapidly-drained, gravelly a l l u v i a l terraces'in the Cameron and Nanaimo River valleys and near Cowichan Lake. Sites range in elevation from 95 to 440 m on various aspects. A l l are in the THPL-TSHE(PSME) vegetation series. Rooting depth i s generally 50 cm or le s s . A l l s i t e s have s o i l s with either sandy texture or high coarse fragment content. Canopy density i s greater for c o l l u v i a l slopes (average 74 93%) than for a l l u v i a l terraces (average 75%). Height of the dominant trees ranges from 44 to 60 metres. Basal area i s also quite variable (83 - 158 m 2/ha). A l l stands are between 275 and 360 years old. The dominant overstory species is Pseudotsuga menziesii with Thuja p i i c a t a , Tsuga heterophylla and Acer macrophyllum of varying amounts in the lower canopy layers. A few Douglas-fir seedlings are present on most plots, with abundant western hemlock seedlings in a few samples. The distinguishing c h a r a c t e r i s t i c of t h i s habitat type i s the constant presence of Holodiscus discolor in the t a l l shrub (BI) layer (Figure 10). With the exception of one sample, cover of t h i s species is 10% or greater. Rosa gymnocarpa, Berberis  nervosa and Vaccinium parvifolium are present in most samples. One sample has 90% cover of Gaulther ia shallon and appears to border the PSME/GASH-BENE habitat type except for i t s high cover of Holodiscus discolor (40%). Polystichum munitum dominates the herbaceous layer along with the constantly high cover of Achlys t r i p h y l l a . Several species that occur in most stands include Lactuca muralis, Linnaea borealis, T r i e n t a l i s l a t i f o l i a , Galium t r i florum, T i a r e l l a l a c i n i a t a , T i a r e l l a t r i f o l i a t a and Polypodium  glycyrrhiza in order of decreasing importance. Rhytidiadelphus  loreus and Rhytidiadelphus triquetrus are common mosses along with varying amounts of the ubiquitous S t o k e s i e l l a oregana and Hylocomium splendens. Plagiomnium insigne and Isothecium  stoloniferum also occur frequently. A type similar to the PSME/HODI/POMU habitat type has not 75 Figure 10. Psojxdo tsuga. mmziesLi / Holodli>cvu> discolor I Polystichum munUum (PSME/HODI/POMU) habitat type. F i g u r e 11. Abla gsiancLii / Polystichum munitum (ABGR/POMU) h a b i t a t t y p e . 76 been described in B. C. Apparently t h i s habitat has been included in the Polystichum-dominated biogeocoenoses of Krajina (1969) and the Polystichum munitum - Achlys t r i p h y l l a association of Klinka (1977). It i s similar to the PSME/HODI habitat'type described by Dyrness et a l . (1974) for the dri e s t habitats in the Tsuga heterophylla vegetation zone of the central western Cascades of Oregon where Douglas-fir i s the edaphic climax tree species. The cover of Polystichum is lesser and the frequency of Gaultheria i s greater in their type. 5.3.2.4. Abies grandis / Polystichum munitum (ABGR/POMU) habitat type The ABGR/POMU habitat type was the most extensively sampled type in the PSME-TSHE zone. Most sampling was done near the Oyster River where some of the few old-growth stands of thi s type remain. It occurs mostly in the ABGR vegetation series, but also in the THPL-TSHE(PSME) series. Samples are on middle to lower slopes (up to 75% slope) and a l l u v i a l terraces from 10 to 300 m in elevation. Most si t e s are on moderately well-drained to well-drained a l l u v i a l or g l a c i o - f l u v i a l parent materials with moderate coarse fragment content. S o i l texture ranges from clay loams to loamy sands. Many of the s o i l s have a well-developed Ah horizon. In stands ranging in age from 200 to 500 years, the average height of dominant trees is 50 m. Basal area on plots exceeds 200 m2/ha. A few 90- to 120-year-old stands were also sampled that are 35-40 m in height, with considerably smaller basal area. Canopy density for a l l samples i s high (average 93%). 77 The overstory i s dominated by Pseudotsuga menziesi i , but with s i g n i f i c a n t amounts of Thuja p i i c a t a and Abies grandis as codominants. Acer macrophyllum i s a common component of a l l canopy layers. Tsuga heterophylla is a constant component with high cover in the lowest canopy layers. Regeneration of a l l of these species except Douglas-fir is present in varying amounts. Vacc inium parvifolium, Rubus s p e c t a b i l i s and Berberi s  nervosa are nearly constant shrub species. Rubus ursinus i s frequently present. The herbaceous layer is almost completely dominated by Polystichum munitum on most s i t e s (Figure 11). Occurring constantly along with Polyst ichum munitum are T i a r e l l a  t r i f o l i a t a and Achlys t r i p h y l l a . In small patches where Polyst ichum mun i turn cover does not dominate, a r i c h herbaceous layer i s present. Species occurring in most samples include T r i e n t a l i s l a t i f o l i a , Galium t r i f l o r u m , T r i l l i u m ovaturn, T i a r e l l a lac in iata, Adiantum pedatum, Adenocaulon bicolor, Lactuca muralis, and Dryopteris austriaca. The most common grass species is Bromus p a c i f i c u s . Several other herbaceous species are found in scattered stands; none of these herbs have high constancy, but they are rarely found in other habitat types of this zone (Appendix VI, Table A27). A r i c h moss f l o r a i s found on humus, rock and trees, p a r t i c u l a r l y on hardwood species. The ABGR/POMU habitat type corresponds to three of Krajina's (1969) biogeocoenoces: 1, l a and 2a. Number l a has a [Mnium] Plagiomnium insigne - [Eurhynchium] • Stokesiella  stokesi i - Polystichum munitum - T i a r e l l a t r i f o l i a t a  Pseudotsuga menziesi i - Abies grandis - Thuja p l i c a t a phytocoenosis. Klinka (1977) recognizes a Polystichum - T i a r e l l a 78 association for the CDF wet subzone. The Douglas-fir - western red cedar - swordfern association of Krajina and Spilsbury (1953), McMinn (1957) and Mueller-Dombois (1959) are equivalent. The ABGR/POMU type is similar in understory composition to the TSHE/POMU habitat type of t h i s study that is also described for Oregon and Washington (Franklin and Dyrness 1973, Franklin et a l . 1979) and to the swordfern types of the CWH biogeoclimatic zone in B r i t i s h Columbia. 5.3.2.5. Thuja p i i c a t a / Lysichitum americanum (THPL/LYAM) habitat type The THPL/LYAM habitat type was found on imperfectly- to poorly-drained s i t e s at the base of slopes or in topographic depressions. A l l are in the THPL vegetation series from stands near the Tsable River and Haslam Creek. Clay loam s o i l s with few coarse fragments derived from glacio-marine and morainal deposits are t y p i c a l for t h i s type. Roots are concentrated in the surface organic layers. The three stands sampled were a l l under 200 years old with variable canopy cover. Dominant trees are less than 30 m t a l l . The upper canopy layers are dominated by Thuja p l i c a t a and Tsuga heterophylla. Alnus rubra is abundant in the intermediate canopy layer. Populus trichocarpa and Picea sitchensis are found in the Tsable River stands. Because of the small number of stands, i t i s d i f f i c u l t to give r e l i a b l e constancy values. The shrub layer commonly includes Vaccinium parvifolium, with Rubus s p e c t a b i l i s , Rhamnus  purshiana and Ribes bracteosum frequently associated. The herb 79 layer has several constant species: Lysichitum americanum, Athyrium f i1ix-femina, Polyst ichum munitum, Dryopteris  austriaca, and T i a r e l l a l a c i n i a t a (Figure 12). Other common herbs are Blechnum spicant, Equisetum spp., Veratrum v i r i d e , Streptopus amplexifolius, Lactuca muralis, Polystichum l o n c h i t i s and T i a r e l l a t r i f o l i a t a . Common mosses include Isothec ium  stoloniferum, Stokesiella oregana, Rhizomnium glabrescens, Plagiothecium undulatum and Plagiomnium insigne. Associations corresponding to the THPL/LYAM habitat type have been described by nearly a l l of the studies of coastal B. C. The phytocoenosis given by Krajina (1969) i s : Carex obnupta -Lysichitum amer icanum - Alnus rubra - Picea sitchensis - Thuja  p i i c a t a (biogeocoenosis #16). Krajina and Spilsbury (1953), McMinn (1957) and Mueller-Dombois (1959) describe similar associations for the Douglas-fir zone. Orloci (1961) and Kojima (1971) found similar types in the adjacent Coastal Western Hemlock biogeoclimatic zone. Franklin and Dyrness (1973) and Franklin et a l . (1979) describe t h i s habitat type as a forested swamp. The samples in the present study were of similar species composition to those of other authors except for the absence of Carex obnupta . 5.3.3. Ordination results 5.3.3.1. DECORANA The 29 samples within the PSME and PSME-TSHE vegetation zones were ordinated with DECORANA using a l l species, understory species only and tree species only. The most successful 81 ordinations, judged in purely subjective fashion, were obtained using understory data transformed to an octave 1 3 cover scale with some deletion or downweighting 1 4 of rare species. Results of. a DECORANA sample ordination of a l l species (with less than three occurrences deleted) are presented in Figures 13 and 14. Axis 1 and 2 represent the f i r s t two axes of var i a t i o n i d e n t i f i e d by DECORANA from a plot-by-species data matrix. The length of axis 2 is 64% of axis 1 and represents only 37% of the variation (indicated by i t s eigen value) of axis 1. A t h i r d axis (not plotted) represented only 12% of the var i a t i o n of axis 1. These figures suggest a strong major axis of var i a t i o n ( i . e . gradient) along which samples are di s t r i b u t e d . Vegetation series (Figure 13) were di s t r i b u t e d along the f i r s t ordination axis with samples from the PSME and THPL series clustered at the right and l e f t extremes of the diagram, respectively. The ABGR and THPL-TSHE(PSME) series occupy an intermediate position closer to the l e f t end of the diagram. These two series overlap considerably. The d i s t r i b u t i o n trend along the second axis was uninterpretable. The f i v e habitat types (Figure 14) showed a d e f i n i t e preference for certain portions of the ordination diagram. The sequence of types from l e f t to right along the f i r s t axis i s : THPL/LYAM, ABGR/POMU, PSME/GASH-BENE, PSME/HODI/POMU and PSME-1 3 Octave i s an 8-point scale based on logarithms (base 2) for input values from 0 to 100. 1 4 A procedure in which cover values of rare species are reduced by a factor proportional to the cover of the most abundant spec ies. 82 F i g u r e 13. DECORANA o r d i n a t i o n of samples i n the PSME and PSME-TSHE zones - ve g e t a t i o n s e r i e s . 200 • • • • • • • • ^ PSME ABGR DTHPL-TSHE(PSME! ^THPL • a x i s 1 320 F i g u r e 14. DECORANA o r d i n a t i o n of samples i n the PSME and PSME-TSHE zones - h a b i t a t types. 200 J *7 • • • A PSME/ARME/GASH • PSME/GASH/BENE • PSME/HODI/POMU ^ ABGR/POMU ^ THPL/LYAM • • a x i s 1 320 83 ARME/GASH. The types at the extremes of axis 1 formed the most d i s t i n c t c l u s t e r s . The ABGR/POMU type occupies a narrow band on axis 1 but i s spread out along axis 2. Part of the va r i a t i o n r e f l e c t e d in axis 2 may be the result of sample location.^ Several samples in the same area along the Tsable River are separated from the other samples. The PSME/GASH-BENE type overlaps the other two intermediate types along axis 1 but i s •separated f a i r l y well along axis 2. The PSME/HODI/POMU type forms an i n d i s t i n c t cluster adjacent to the ABGR/POMU type. One sample of t h i s type had a high percent cover of Berberis nervosa that caused i t to be grouped within the PSME/GASH-BENE c l u s t e r . Another sample had unusually high Gaultheria shallon cover; consequently, i t is grouped near the PSME-ARME/GASH type. Similar e f f e c t s causing the ordination of individual plots could be noted. An accompanying species ordination, generated along with the sample ordination, is shown in Figure 15. Only the more common species have been plotted. The reader may consult this ordination and the association tables (Tables A24 - A28) for further interpretation of the sample ordination. To i l l u s t r a t e the improvement that DECORANA makes over normal reciprocal averaging, an RA ordination with the dame data input as Figures 13 - 15 is presented in Figure 16. The "horseshoe e f f e c t " caused by the quadratic r e l a t i o n s h i p of the f i r s t and second axes i s obvious. This tends to obscure the fact that the ordering along the f i r s t axis is i d e n t i c a l to that of DECORANA. Figure 17 presents a DECORANA ordination of understory species alone. Habitat types form much more d i s t i n c t c l u s t e r s in Figure 15. DECORANA ordination of species in the PSME and PSME-TSHE vegetation zones.-'-•ACMA •ACMA2 •BRPA *CAREX *STAM •ATFI ABGR* •RUSP 'TILA *ADPE •TITR •GATR •LAMU •EQUIS *DRAU •LYAM •POTR •BLSP •THPL THPL2* * •POMU •ACMA3 •ABGR2 •VAPA •ABGR3 •POGL4 •TRLA2 •LIB02 •RUUR •PSME •BENE •PSME2 •ELGL •TBOC . *LANE "ARMA3 •CHUM SYAL* •HODI •ROGY •GOOB •PRVE •TSHE2 •TSHE •PTAQ ^Frora the sample ordination shown in Figures 13 and 14. axis 1 560 85 Figure 16. Reciprocal averaging ordination of samples i n the PSME and PSME-TSHE zones - habitat types. 72 CO •H X re! \7 X7 ^7 ^7 \7 • • • \7 A PSME/ARME/GASH • PSME/GASH/BENE • PSME/HODI/POMU ^7 ABGR/POMU • THPL/LYAM ^7 \7 A AA axis 1 100 Figure 17. DECORANA ordination of samples in the PSME and PSME-TSHE zones - habitat types (understory only). 240 CN •rH X \ 7 ^7 \7 • ^7 ^7 \7 A \7 • • • A • axis 1 360 86 th i s ordination than when trees species were included. The d i s t r i b u t i o n pattern of types is very similar; however, the PSME/GASH-BENE type i s now intermediate between the PSME-ARME/GASH and PSME/HODI/POMU types. By eliminating tree species, plots in the PSME/GASH-BENE type having some Alnus rubra, Acer  macrophyllum, Taxus brevi f o l i a and Cornus nuta l i i no longer show as great a s i m i l a r i t y to the ABGR/POMU type. The great d i v e r s i t y of tree species in the ABGR/POMU type contributed to a wide spread of plots along the second axis in the f u l l species ordination. When trees are eliminated, t h i s type forms a tight c l u s t e r , r e f l e c t i n g the high dominance of Polyst ichum munitum and the nearly constant presence of T i a r e l l a t r i f o l i a t a , T i a r e l l a l a c i n i a t a , Achlys t r i p h y l l a , Galium tri f l o r u m , T r i l l i u m  ovatum, Berberis nervosa and Vacc inium parvifolium. Conversely, the absence of trees in ordinating the PSME-ARME/GASH type caused a greater spread of samples because the two dominant overstory species, Arbutus menziesi i and Pseudotsuga menziesi i , tended to draw samples together with an otherwise diverse understory. In the understory ordination, two plots were segregated from the other PSME-ARME/GASH samples because of lower cover of Gaulther ia shallon and Festuca occ i d e n t a l i s , more Berberis nervosa and Bromus vulgaris and the addition of few minor species. These properties caused them to be grouped closer to the PSME/GASH-BENE type, while three plots with Holodiscus  discolor present ordinated toward the PSME/HODI/POMU type. Three samples in the PSME/GASH-BENE type are closest to the ABGR/POMU group due to the presence of Achlys t r i p h y l l a . Within the PSME/HODI/POMU type, two separate groupings were formed on 87 the basis of the amount of Polystichum munitum and T i a r e l l a  t r i f o l i a t a present, the plots having greater abundance of these species ordinating closer to the ABGR/POMU type. The THPL/LYAM type was not s i g n i f i c a n t l y affected by the elimination of tree species. Overall, the understory ordination produced the best segregation of types. Ordination of samples using overstory basal area data alone was least e f f e c t i v e for id e n t i f y i n g communities (Figure 18). Because of high frequency of common species (Douglas-fir, red cedar, western hemlock), rare or less common species had the most e f f e c t upon the ordination. There is a large cluster of samples, representing nearly a l l habitat types, with similar overstory composition. The THPL/LYAM habitat type appears most d i s t i n c t i v e in terms of tree species because of abundant Alnus  rubra and the presence of two species not found in other types: Pppulus trichocarpa and Picea s i t c h e n s i s . Two plots in the ABGR/POMU habitat type were separated from the others because of large Abies grandis dominants. Other plots in t h i s type were separated because of varying amounts of Alnus rubra, Abies  grandis, Acer macrophyllum, and Thuja p l i c a t a . There was much more tree species v a r i a b i l i t y in wet types than in moist or dry habitat types. 5.3.3.2. Polar ordination A polar ordination was performed where endpoints were chosen to represent the dri e s t and wettest conditions sampled (Figure 19). Plots from the PSME-ARME/GASH and ABGR/POMU habitat types represent the dry and wet extremes of the f i r s t ordination Figure 18. DECORANA ordination of samples in the PSME and PSME-TSHE zones (overstory only) using basal area. 300 + I N A P S M E / A R M E / G A S H • PSME/GASK/BENE • P S M E / H O D I / P O M U \7 A B G R / P O M U • T H P L / L Y A M A A D A ° * A A • • • ^7 • ^ ^ 7 \7 axis 1 320 Figure 19. Polar o r d i n a t i o n of samples i n the PSME and PSME-TSHE zones - h a b i t a t types (understory o n l y ) . 100 + ^ 4 PSME/ARME/GASH „ ^ , . ,,„ © PSME/GASH/BENE wet . p o o r l y - d r a i n e d • PSME/HODI/POMU _ \7 ABGR/POMU •V THPL/LYAM A A A A " d r y , w e 1 1 - d r a i n e d " \7 ^7 ^7 _ _ • • ^7 • • " w e t , w e l l - d r a i n e d " a x i s 1 100 89 axis, respectively. To separate samples along a second axis, a well-drained ABGR/POMU sample and a poorly-drained THPL/LYAM sample were chosen. The fiv e plots in the PSME-ARME/GASH type form a cluster at the 'dry' and of axis 1. At the other extreme, the ABGR/POMU type forms a well-spread grouping which is separated along the second axis from the THPL/LYAM type. The PSME/HODI/POMU and PSME/GASH-BENE types occupy an intermediate position along the f i r s t axis, with the former type closer to the Polystichum  munitum type. These two types do not form d i s t i n c t c l u s t e r s , though they are separated f a i r l y well from each other along the second axis. The resulting diagram i s a triangular shape, the points of which represent dry and well-drained, wet and well-drained, and wet and poorly-drained conditions. The differences in the spread of plots within types indicate much greater homogeneity within samples from the PSME-ARME/GASH type than any of the other types. This is p a r t i a l l y due to the high dominance of Douglas-fir throughout the type, which would have a large numerical weight in ordination c a l c u l a t i o n s . The two intermediate types are least d i s t i n c t ; however, the tighter clusters formed at the extremes of axis 1 are deceiving because there are no wetter or dr i e r plots beyond the axis ends that would tend to spread them out further. The lack of sharp breaks between types and the spread of plots within types suggests that species composition changes gradually between types along moisture and drainage gradients rather than abruptly. To test the effect of changing data input and choice of endpoints on the overall interpretation of polar ordination, a 90 number of data transformations and sample endpoints were chosen. Results showed a similar o v e r a l l relationship between habitat types when samples from the same types as in Figure 19 were used as endpoints, though the positioning of individual samples changed. A d e f i n i t e trend in species richness was i d e n t i f i e d for samples in the zone. The number of species (understory and overstory) was greatest at the dry and wet extremes of the gradient. Canopy cover was investigated as a possible factor a f f e c t i n g d i v e r s i t y . A simple li n e a r regression of percent canopy cover and t o t a l number of species for each sample showed no c o r r e l a t i o n whatsoever. Canopy cover data did, however, show that the smallest canopy cover is concentrated at the dry end of the moisture gradient. Canopy cover on wet habitats remained high due to the presence of hardwood species despite the few number of stems. 5.3.3.3. Relationship of ordination axes to environmental gradients Environmental variables were plotted on the results of the DECORANA ordination of understory species in an attempt to interpret the axes in r e l a t i o n to environmental features. Results show that axis 1 of Figures 13 and 14 corresponds to a "complex" moisture gradient influenced by slope p o s i t i o n , s o i l , drainage, and s o i l AWSC (texture, depth and coarse fragment content). The word complex is used to refer to a gradient that is the sum t o t a l of several factors, as opposed to a single factor (e.g. elevation). Plotting elevation, temperature, 91 p r e c i p i t a t i o n and solar radiation revealed no strong o v e r a l l patterns, though the positioning of some individual plots could be explained by one or more of these factors. No single gradient could be i d e n t i f i e d as corresponding to the second axis. S o i l texture becomes progressively finer from the sandy loams of the PSME-ARME/GASH type to the clay loams of the THPL/LYAM type (Tables A9 - A13). V e r t i c a l drainage r e f l e c t s these textural changes. The trend along axis 1, from l e f t to right, i s from rapid to imperfect drainage. Topographic position is also associated with these differences. The types at the l e f t occupy upper slope positions and the types on the right occupy river f l a t s and depressions. Figures •20a and 20b plot AWSC and WSI on the ordination diagram of Figure 14. Samples with less than 12 cm of AWSC are concentrated on the right half of axis 1. A l l of the PSME-ARME/GASH and PSME/GASH-BENE samples are included in thi s group along with three of the six PSME/HODI/POMU samples. Of the remaining three PSME/HODI/POMU samples, two have an AWSC of 12.5 cm. One ABGR/POMU sample also had low AWSC; however, i t i s located on an a l l u v i a l f l a t and would receive input of groundwater. Results of WSI calc u l a t i o n s show that potential water stress is commonly severe in the PSME-ARME/GASH and PSME/GASH-BENE types, variable in the PSME/HODI/POMU type and rare in the ABGR/POMU and THPL/LYAM types. This i s shown in a a general trend along the f i r s t axis. As Figure 20b suggests, there i s considerable v a r i a b i l i t y among WSI values in the central portion of the axis. Imperfectly-drained s i t e s have been i d e n t i f i e d on 92 Figure 2 0 a D E C O R A N A ordination of samples in the P S M E and P S M E - T S H E zones - vegetation s e r i e s , with W S I plotted. 200. CN LO •H X rti • 0+.5 0*2 • .5+2 • 0+5 • • * 0+1 • • .5+3.5 A P S M E • A B G R D T H P L - T S H E ( P S M E ) ^ T H P L 3+2 • 4+1 A axis 1 A U 4 A' 5+0 2.5+1.5 1+1.5 4*1 A 1+2.5 A W S I : months of stress severe + moderate *imperfect drainage 320 Figure 20b. D E C O R A N A ordination of samples in the P S M E and P S M E - T S H E zones, with A W S C plotted. 200 CN CO •H X rti • Q • Q A W S C Q 12cm A 12cm Q ° Q Q Q Q D Q axis I Q 3 2 0 93 the diagram because the WSI calculations assume drainage to f i e l d capacity following p r e c i p i t a t i o n input. WSI values for these s i t e s are questionable, yet even s i t e s which are imperfectly - drained part of the year can exhibit some stress during summer drought periods. 5.4. Description and analysis of the Tsuga heterophylla (TSHE) vegetation zone 5.4.1. Description of the vegetation series Only three vegetation series are represented in samples from the Tsuga heterophylla zone. Most samples are from the zonal TSHE serie s . This series is i d e n t i f i e d as any area where Tsuga heterophylla is reproducing successfully and reproduction of either Pseudotsuga menziesi i or Abies amabilis i s absent. Thuja p l i c a t a may be present as a co-climax species. This series occupies t y p i c a l , well-drained s i t e s within the zone. The THPL-TSHE(PSME) series is i d e n t i f i e d by the successful reproduction of Tsuga heterophylla and Thuja p i i c a t a with the addition of marginally successful reproduction of Pseudotsuga  menz i e s i i . Abies grandis is also frequently present. This series occupies somewhat warmer and drier sites than the TSHE serie s . The TSHE-ABAM series is defined as an area having successful reproduction of Abies amabilis, but Tsuga  heterophylla s t i l l dominates the understory reproduction. Thuja  p l i c a t a i s also present as a co-climax species. Areas supporting th i s series are cooler and wetter than average for the zone and are usually found as t r a n s i t i o n a l types bordering the ABAM-TSHE 94 zone, or in areas with frost pockets or cold a i r drainage. 5.4.2. Description of habitat types Site c h a r a c t e r i s t i c s for samples in each habitat type of the TSHE zone are presented in Appendix V, Tables A14 - A16. Association tables for each type are geven in Appendix VI, Tables A29 - A31. Un c l a s s i f i e d samples are l i s t e d in Table A32. Table V summarizes the vegetation c h a r a c t e r i s t i c s for habitat types in t h i s zone. 5.4.2.1. Tsuga heterophylla / Gaultheria shallon (TSHE/GASH) habitat type Stands belonging to this habitat type were found between 440 and 740 m in elevation on c o l l u v i a l and morainal parent materials. Slope and aspect are quite variable. S o i l texture is generally loam to loamy sand with moderate to high coarse fragment content. The majority of stands occupy well-drained middle slopes. Sample stands are from the Cameron, Nanaimo and Englishman River valleys, and are mostly in the TSHE vegetation seri e s . The majority of the stands are from 250 to 300 years old, and have f a i r l y open canopies, the average density is 85% with no stand exceeding 89%. Timber volume is low in t h i s habitat type as indicated by an average height of 31 m and an average basal area of 65 m2/ha, excluding one low elevation (440 m) stand in the valley floor with high volume. The overstory i s characterized by Pseudotsuga menziesii in the dominant and co-dominant canopy positions with a lower canopy of Tsuga heterophylla and Thuja p l i c a t a . Western hemlock Table V. Summary of major species f o r habitat types i n the TSHE vegetation zone. TSHE/ TSHE/ TSHE/ TYPE: G A S H GASH/BENE/ POMU ACTR TREE LAYER 1 PC MC2 PC MC PC MC Abtej> amxbAJLLt, i 1.7 I 1.4 I + .0 Ptnu6 morvtlcola i + .3 I 1.4 I + .0 P6eM.dotAu.ga me.nztzA-11 V 7.9 V 7.9 V 6.7 Thuja pLLcata IV 4.0 V 4.4 IV 2.2 Ttuga hoXzAophylta. IV 5.0 V 6.8 V 5.8 SHRUB LAYER AmttanckleA a l n l ^ o t l a I + .0 III + .4 V 4.7 III + .9 V 3.1 III 1.2 GadxJUJtiznJjx AhaLLon V 9.3 V 7.2 POACL gymnocaApa II + .0 V 4.6 II + .0 Rubai uAAtmu IV 1.7 SymphoAA.caApa6 albui III 1.9 Vaccixuixm alxu>ka.e.n6e. II 1.8 3.3 Vajzcjjvum ovaLi^oZtum III + .8 Vacctntum paA.vt^oZtum II 3.0 V 2.4 V 2.4 HERB LAYER kchtyi, tfUphyZIa I + .0 V 5.0 V 4.0 Adunocouuton btcoloh. V 1.6 I + .0 ChimaphUji me.nzt&A<U. III 1.4 IV + .1 III 1.1 Ckunaphtla. umboJUtxita. V 3.2 V 1.8 IV 1.8 COAJOUA canadzn&AJ, I + .3 V 3.2 I + .7 Futuca. occidtntatl6 II + .6 IV 1.7 GoodyeAa obtongt^otta IV + .5 V 1.2 IV + .5 Lactuca mufuxZJJ> II 1.3 III + .0 I I I 1.1 Lathytux& nevaden^ -cA V 4.1 Linnaoji boA.zatu> IV 4.9 V 2.7 I 1.4 LustoAa. cawvina. I + .0 I + .0 I + .7 LL&teAa coidata IV 2.0 II + .0 II + .0 QAmonklza. cht£e.n&-L& IV + .5 PzdtcuLa/uA fiacmo^a. V 3.0 Poty^tichum mujuXum II 1.1 III + .0 V 7.4 PtVujLhxm aquXJLinum IV 2.2 I + .0 TJjLVieXla ZacUruata I + .0 II + .0 IV 2.0 TiaKoJULa. tnJi^oUjxta. I + .0 II 3.2 V. 2.5 ThlcYVtaJLu> l£Lti{,oLia II 1.0 II 1.4 I l l 1.5 TAAAVtium ovatum II + .0 V + .4 Vtola ohJbtcutata. II + .0 V 3.7 IV 1.1 ^"Presence Class: I = 1-20%; II = 21-40%; III = 41-60%; IV = 61--80%; V = 81-100% 2 Mean Coverage: Values given are averages of the following c l a s s e s + = trace; 1 = 0.1-1.0%; 2 = 1.1-2.2%; 3 = 2.3-5.0%; 4 = 5.1-10.0%; 5 = 10.1-25.0%; 6 = 25.1-33.0%; 7 = 33.1-50.0%; 8 = 50.1-75.0%; 9 = over 75.0% 96 and red cedar regeneration predominates. The shrub layer i s completely dominated (80-90% cover) by Gaultheria shallon with occasional Vaccinium parvifolium and Berberis nervosa (Figure 21). Linnaea borealis and Chimaphila  umbellata are nearly constant in the herb layer. Cover of these two species is considerable in some samples. L i stera cordata, Chimaphila menziesi i , Goodyera oblongi f o l i a and T r i e n t a l i s  l a t i f o l i a are present in most samples. Most other herb species are infrequent because of the dominant s a l a l cover. Many other herb species occur infrequently and with low abundance. The moss layer i s characterized by high cover of Hylocomium splendens. Rhytidiadelphus loreus i s also abundant with frequent presence of Pleurozium shreber i , Rhyt i d i o p s i s robusta, Rhyt idiadelphus  triquetrus, and Brachythecium asperimum. The TSHE/GASH type corresponds to the [Eurhynchium] St o k e s i e l l a oregana - Mahonia [Berber i s ] nervosa - Gaulther ia  shallon - Pseudotsuga menziesi i - Tsuga heterophylla phytocoenosis (biogeocoenosis #27) of Krajina (1969). The Hylocomium splendens - St o k e s i e l l a oregana - Gaultheria shallon - Pseudotsuga menziesii association described by Kojima (1971) from Vancouver Island i s i d e n t i c a l to this habitat type, with s i m i l a r l y high abundance of Linnaea borealis and Chimaphila  umbellata. Orloci's (1961) orthic Gaultheria forest type i s similar except that Holodiscus discolor and Pteridium aquilinum are present in s i g n i f i c a n t amounts in his type. The TSHE/GASH habitat type of Franklin et a_l. (1979) is nearly i d e n t i c a l to th i s TSHE/GASH type except for the presence of Acer circinatum as a conspicuous shrub in the Mt. Rainier study. This i s also 97 F i g u r e 21. Tsuga h u t e A o p h y l M I GaulXhvuxx. shallon (TSHE/GASH) h a b i t a t type. F i g u r e 22. Tsuga hcttftophylla / Gaulthzxjjx shallon - ZoJibexls nznvosa / Achlys tAAjphylla (TSHE/GASH-BENE/ACTR) h a b i t a t type. 98 true for the Tsuga heterophylla - Acer circinatum - Gaultheria  shallon type of C o r l i s s and Dyrness (1965) for the Oregon Coast Range. In the central western Cascades of Oregon, Dyrness et a l . (1974) have described a Tsuga heterophylla - Rhododendron  macrophyllum - Gaultheria shallon association that occupies somewhat drier habitats than the TSHE/ACCI/GASH type. The abundance of Rhododendron macrophyllum distinguishes i t from types further north. This species becomes less common in Washington and rare in B. C. (Franklin and Dyrness 1973). Castanopsis chrysophylla is another conspicuous shrub present in the Oregon Cascades type that is absent in the present study. The Douglas-fir and Douglas-fir - western hemlock types of Fonda and B l i s s (1969) for the Olympic mountains and the PSME/GASH community type of del Moral and Long (1977) are also related. 5.4.2.2. Tsuga heterophylla / Gaultheria shallon - Berberis  nervosa / Achlys t r i p h y l l a (TSHE/GASH-BENE/ACTR) habitat type This habitat type was found only in the Oyster River val l e y . Samples range from 475 to 725 m in elevation on irregular concave mid-slope positions in the TSHE and TSHE-ABAM vegetation series. Parent material i s a thin layer of g l a c i a l t i l l (50-80 cm) over bedrock. S o i l texture i s generally loam to sandy clay loam with moderate . stoniness. Stands are a l l approximately 260 years old, with an average dominant canopy height of 37 m. The overstory i s dominated by Pseudotsuga menziesii, with a s i g n i f i c a n t amount of Tsuga heterophylla and Thuja p l i c a t a . Western hemlock i s very abundant in the lower canopy layers. 99 There is some disturbance from cedar pole cutting in most of these stands; consequently, the t o t a l volume and proportion of cedar in these samples i s underestimated, even though s i t e s were selected where cutting had been minimal. The shrub layer consists of variable cover of Gaultheria  shallon, Rosa gymnocarpa and Berberis nervosa (Figure 22). Vacc inium parvifolium i s a constant species and is the most abundant of the Vaccinium species present, which include: Vacc inium alaskaense, Vaccinium ovalifolium and a single occurrence of Vacc inium membranaceum. Symphoricarpos albus, Symphoricarpos mollis and Amelanchier a l n i f o l i a are also present on 50% of the s i t e s . A r i c h herb layer i s present, including several species that occur in a l l sampled stands (in order of decreasing abundance): Achlys tr i p h y l l a , Lathyrus nevadensis, Chimaphila  umbellata, Adenocaulon bicolor and Goodyera o b l o n g i f o l i a . Viola  orbiculata, Cornus canadensi s and Pedicular is racemosa are abundant in a l l but one sample. Other herbs of frequent occurrence include: Festuca occidentalis, Pteridium aquilinum, Osmorhiza c h i l e n s i s , and Chimaphila menziesi i . The high cover of Lathyrus nevadensis in a l l samples i s unique in that t h i s species was not encountered in great abundance or frequency elsewhere in the study area. Hylocomium splendens i s the most dominant moss species along with Rhytidiadelphus loreus. Rhytidiopsis robusta and Rhyt idiadelphus triquetrus are also common. The TSHE/GASH-BENE/ACTR habitat type has no equivalent in types previously described for B. C. It appears intermediate in 100 moisture regime to the Gaultheria- and Polystichum-dominated associations of Orloci (1961), Krajina (1969) and Kojima (1971). The types that these authors describe as intermediate are "moss" types dominated by Hylocomium splendens and Stokesiella oregana with Berberis nervosa in the shrub layer. These c h a r a c t e r i s t i c s are not shared by this habitat type. The most similar type i s the TSHE/ACTR habitat type described for Mt. Rainier National Park by Franklin et a_l. (1979). It has a r i c h herbaceous layer and low cover of Gaulther ia shallon, distinguishing i t from the same author's TSHE/GASH type. Because Franklin et a l . 's TSHE/ACTR type i s of limited areal extent, is mostly within 250 year old stands, and has not been described elsewhere, they question i t s status as a habitat type. The TSHE/GASH-BENE/ACTR habitat type is very similar in species composition and dominance, with the exception of Acer circinatum, and i s found on similar southerly exposures. It i s also within the same age class of stands (range 240 - 265 years), though th i s may be coinc idental. 5.4.2.3. Tsuga heterophylla / Polystichum munitum (TSHE/POMU) habitat type The TSHE/POMU habitat type was found on a variety of aspects on moderate to steep middle to lower slopes of the TSHE and TSHE-ABAM vegetation series. The lowest elevation samples are near Cameron Lake at 240 and 270 m in elevation. Other samples range from 500 to 680 m. Parent material i s of g l a c i o -f l u v i a l , morainal and c o l l u v i a l o r i g i n . S i l t loam s o i l s are most common, but textures range from sandy loam to clay loam. S o i l s 101 have moderate to high coarse fragment content and are moderately well- to imperfectly-drained. Stand age, tree height, canopy cover and basal area a l l varied considerably between samples. The overstory is dominated by Pseudotsuga menziesi i , with Tsuga heterophylla and Thuja p i i c a t a common in co-dominant to intermediate layers. The understory is dominated by Tsuga  heterophylla, as is the seedling layer. Berber i s nervosa and Vaccinium parvi folium are constant shrubs of variable cover. The herb layer is characterized by high cover of Polyst ichum mun i turn with the constant presence of Achlys t r i p h y l l a , T i a r e l l a tr i f o l i a t a and T r i l l i u m ovatum (Figure 23). T i a r e l l a lac in i a t a , Viola orbiculata, and Chimaphila umbellata are common. Species c h a r a c t e r i s t i c of wetter habitats occur in some samples. The moss layer commonly includes Stokesiella oregana, Plagiothec ium undulatum, Hylocomium splendens and Rhyt idiadelphus loreus. The TSHE/POMU habitat type is comparable to other swordfern dominated types described for B. C. and the P a c i f i c Northwestern U. S.: biogeocoenoses 2b and 24 (Krajina 1969), the Achlys t r i p h y l l a - Polyst ichum munitum association (Kojima 1971), the Polystichum forest type (Orloci 1961) and the TSHE/POMU habitat type (Franklin and Dyrness 1973, Franklin et a l . 1979). 5.4.2.4. Unc l a s s i f i e d samples Several stands were sampled that could not be c l a s s i f i e d in the preceding description of habitat types. Two s i t e s (plots 1131 and 1146) occur on steep (80%) c o l l u v i a l slopes on 102 Figure 23. Tsuga hctoAophylla / Polystichum m m i i & m (TSHE/POMU) habitat type. Figure 24. Tsuga heXoAophylla zone - u n c l a s s i f i e d sample with depauperate understory. 103 southwest aspects in the Cameron valley. Pseudotsuga menziesii, Tsuga heterophylla and Thuja p i i c a t a are present in the overstory. There are no shrubs present and only sparse herbaceous cover consisting of Chimaphila menziesi i , Linnaea  boreali s, Viola orbiculata, Achlys t r i p h y l l a , Campanula  scouler i , Pyrola pi c t a , Tr i e n t a l i s lat i f o l i a and several other species. Both si t e s have periodic disturbance from s l i d i n g surface materials preventing the establishment of herbs and shrubs (Figure 24). Some mosses are common on rock surfaces, including Stokesiella oregana, Rhyt i d i ops i s robusta and Hylocomium splendens. These sites resemble the "moss" type of Kojima (1971) and the moss-dominated phytocoenosis (biogeocoenosis #25) of Krajina (1969) that is used to characterize zonal ecosystems (Stokesiella oregana - Hylocomium  splendens - Douglas-fir - western hemlock) in the eastern Vancouver Island drier maritime CWH variant of Klinka e_t a l . (1979). One stand (1129) that i s unlike any other sample consists of a pure Pseudotsuga menziesi i overstory with an understory of Berberis nervosa (40%) and Linnaea borealis (20%). Rosa  gymnocarpa, Chimaphila umbellata and Campanula scouleri are also abundant. No Gaultheria shallon or Polyst ichum munitum is present. Hylocomium splendens (90%) and Sto k e s i e l l a oregana (15%) dominate the moss cover. This s i t e appears to represent intermediate moisture conditions between the TSHE/GASH and TSHE/GASH-BENE/ACTR habitat types. It resembles the [Eurhynchium] Stokesiella oregana - [Berberis] nervosa  Pseudotsuga menziesi i phytocoenosis (biogeocoenosis #28) of 104 Krajina (1969), though the high cover of Hylocomium splendens seems contrary to Krajina and Klinka's (1977) association of Stokesiella oregana with Berberis nervosa. The fourth unique stand (1134) is also completely dominated by Pseudotsuga menziesii. The shrub layer is characterized by high cover of Holodiscus discolor (20%) and Rosa gymnocarpa (12%). A r i c h herb layer includes Adenocaulon bic o l o r , Fragaria  vesca, Hierac ium albiflorum, Chimaphila menziesii, Gali um  t r i florum and Melica subulata. Some Berberis nervosa and Polyst ichum munitum are present. This stand most closely resembles c o l l u v i a l slopes in the PSME-TSHE zone's PSME/HODI/POMU habitat type. 5.4.3. Ordination results 5.4.3.1. DECORANA The most successful ordinations of samples in the Tsuga  heterophylla zone were obtained using 20 of the 25 samples, deleting o u t l i e r s and u n c l a s s i f i e d samples. Results of a DECORANA ordination of a l l species are presented in Figure 25 showing the d i s t r i b u t i o n of vegetation series and habitat types. It is d i f f i c u l t to assess the d i s t r i b u t i o n of series because so few samples are outside of the TSHE series. The three habitat types are separated along the f i r s t axis. The TSHE/GASH type occupies an intermediate position along this axis, but has been s p l i t in two along the second axis. Examination of species tables reveals that this s p l i t is due to differences in overstory composition. Two plots have no Tsuga heterophylla in Figure 25. DECORANA o r d i n a t i o n of samples i n the TSHE zone habitat types and s e r i e s ( a l l s p e c i e s ) . habitat types: is f ^St* ^ TSHE/GASH A A. ® TSHE/G/\SH/BENE/ACTR H TSHE/POMU s e r i e s : * TSHE-ABAM ** THPL-TSHE(PSME) TSHE (unmarked) axis 1 Figure 26. DECORANA or d i n a t i o n of samples i n the TSHE zone habi t a t types and s e r i e s (understory only) ,. 195 axis 1 290 106 the dominant, co-dominant or intermediate layers and l i t t l e in the understory. This contrasts with other samples in the type. Overstory i s almost exclusively Pseudotsuga menziesii in these samples, with some Chamaecyparis nootkatensis in one plo t . Samples in the TSHE/GASH-BENE/ACTR type, a l l found in the same valley, form the tightest grouping. The TSHE/POMU habitat type i s f a i r l y spread out along both axes, indicating variable species composition within the type. Further sampling might suggest s p l i t t i n g t h i s type into more than one unit. A DECORANA ordination of understory species alone i s presented in Figure 26. It produced a much better separation between types than the f u l l species ordination, though type separation i s not d i s t i n c t when considering either axis alone. The elimination of overstory has caused a s h i f t in positioning of types along the f i r s t axis. The PSME/GASH-BENE type i s now intermediate between the others. The order of the three types on the second axis is the same as the f u l l species f i r s t axis. Examination of overstory data revealed no obvious tree species differences between types that would make the overstory of the TSHE/GASH type more similar to the TSHE/POMU type than that of the TSHE/GASH-BENE/ACTR habitat type. The ov e r a l l sum of several small changes in dominance ( p a r t i c u l a r l y red cedar Al and A3 and western hemlock A2 layers) caused t h i s s h i f t . Samples in the TSHE/GASH type are not separated into two d i s t i n c t groups as when overstory was included, but two samples remain most unlike others in the type due to the presence of several species: Lactuca muralis, Polystichum munitum, T r i e n t a l i s l a t i f o l i a and Festuca o c c i d e n t a l i s . The TSHE/POMU type i s more clustered with 107 the removal of overstory species from the ordination. This is due to rare occurrences of Abies amabilis, Abies grandis, Pinus  monticola and Acer macrophyllum in some plots and the variable cover of Douglas-fir, cedar and hemlock in the three canopy layers. Understory composition is less variable than overstory, with the dominant influence of Polystichum muniturn, T i a r e l l a  t r i f o l i a t a , Achlys t r i p h y l l a and T r i l l i u m ovatum. The TSHE/GASH-BENE/ACTR type remaines in a f a i r l y tight cluster compared to other types, with the exception of one plot. This plot has no Cornus canadensis, Pedicular is racemosa, Vaccinium parvi folium or Viola orbiculata that are common to a l l other samples. It i s not noticeably di f f e r e n t from other plots in the cover of Gaultheria shallon, Berberis nervosa and Achlys t r i p h y l l a . The d i s t r i b u t i o n of the major indicator species on the DECORANA understory ordination of samples i s presented in Figure 27. The Polyst ichum munitum and Gaulther ia shallon habitat types are separated well on the basis of high cover of these two species. Achlys t r i p h y l l a and Berberis nervosa occupy similar ranges to each other. The absence of Polyst ichum munitum in half of this range serves to separate the TSHE/GASH-BENE/ACTR samples from the Polyst ichum mun i turn type. Several minor species are more s p e c i f i c to one or more of the types, but their cover and constancy are too sporadic for them to be used as character spec ies. The ordering of species having at least three occurrences in the samples along the f i r s t axis of a reciprocal averaging ordination ( i d e n t i c a l to DECORANA order) is presented in Figure 28. This axis can be subdivided into three segments on the basis Figure 27. D i s t r i b u t i o n of major understory species on a DECORANA understory ordination - TSHE zone. a. Gaultheria shallon '1 1 0 n ^ " 9 99 * ° ,'5 9 g 9 / i3 5 9 \ 4 b. Berberis nervosa / 4 \ '2 \ 0 1 2; 00 3 , + ' 0 0 \ / \ / \ 4 1 1/ s-3^ c. Polystichum munitum / o 5 rs 8 i \ 3/ 0 + s-^9 _----""+'" 0 0 + + 0 0 d. Achlys t r i p h y l l a / 3 5 \ + I I 1 \ 5 4 3/ \ 5 '' 00 e. T i a r e l l a t r i f o l i a t a 3\ H 00 The cover scale used f o r species i s given i n Appendix Table A23. -5^0-1 0 109 Figure 28. D i s t r i b u t i o n of species and habitat types along an RA ordination f i r s t axis - TSHE zone. SPECIES PLOT NO.: 008088521425141336243 21504043382547570838 2 TYPE; ****** + ]++-+++ SYAL SYMO PERA OSCH LANE CASC2 VAOV PTAQ ADBI ROGY AMAL COCA RUUR VAAL HIAL FEOC LIB02 VIOR2 THPL TSHE 2 PSME2 GOOB GASH ACTR CHUM VAPA PSME 3 THPL 3 LIC03 TSHE 3 TRLA2 PSME POGL4 THPL2 BENE TSHE CHME LAMU POLO 2 TITR TROV POMU TILA COMA 3 LICA3 868— 66-6-88486 6-44-4 848484 86 — -6-46 448-4 844444 898888444 -46-44|4 48868 66846 -464981 _ 4 _ -I 6 — !46-84-6-96— 8886861 88884 9—69986-88 •986-6-4 •64 h - 6 — 86 46-66-9-8— 868888J-66686-6 8-86-68686-4866-8 64446646446444-4-44-899886 99996999-69-8-888888-4—4 88-888 664868684648-8-8848-688864-94864-869-684 4-6-444-8—46 4-64 -84884-46666-8-44-64 —4-4-488644-8-4 666889446686-6^46688 - 4 - 4 66 99999999999999999999 4W-44 -4-68--68688 -8868 -4444 —44-418 - 4 - 4 — 4 -6 — 8 -- 4 — 4 B-844-— 8 4 4-48-8-9-6 16—6-6-41 -4-| I 4 — 4 1 -4-4-44 8-4-- 6 — 8—8-898668 -9-899 -68-84 8468 -448 86-688 44-444 988989 -6-688 See Appendix III for f u l l species names. Cover values are scaled by the matrix maximum. Numbers i n tree species abbreviations i n d i c a t e canopy layers (2 = A2; 3 = A3). TSHE/GASH/BENE/ACTR TSHE/GASH TSHE/POMU -4-8-110 of habitat types. One can see from this diagram that the TSHE/GASH-BENE/ACTR habitat type has many species . that distinguish i t from the other two types. Figure 29 shows an ordination using only overstory species. The resulting d i s t r i b u t i o n of samples shows no relat i o n s h i p to habitat types. Because of poor representation of di f f e r e n t series, i t i s hard to evaluate the ordination in t h i s regard; however, the three samples of the TSHE-ABAM series were a l l found on the l e f t half of axis 1. The use of data transformed to an octave scale produced a better spread of samples than raw basal area data because i t reduced the effect of species of high dominance. These results confirm the observation that ordination is a poor technique for data with poor species richness, e s p e c i a l l l y for samples with only a few common and several "rare" species. Overstory composition seems to be a poorer overall indicator of s i t e potential because i t i s more greatly affected by h i s t o r i c events ( i . e . f i r e , weather during seedling establishment, seed source, etc.) than understory, which has had time to s t a b i l i z e over the l i f e of the stand. 5.4.3.2. Polar ordination With some knowledge of the gradients represented in the data, a polar ordination was constructed with chosen endpoint samples representing opposite extremes of moisture and potential radiation gradients. Results are presented in Figure 30. Both overstory and understory species were included. Data were transformed to an octave scale and species with less than three occurrences were deleted. Because radiation and moisture are I l l Figure 29. DECORANA ordi n a t i o n of samples i n the TSHE zcne-with temperature p l o t t e d (overstory only) 180 o B P ** d AVERAGE MAY--SEPT. TEMPERATURE O 10.5 - 11.5° C @ 11.6 - 12 .5° C @ > 12.6°C # c o l d a i r i n f l u e n c e habitat types: TSHE/GASH ® TSHE/GASH/BENE/ACTR gg TSHE/POMU s e r i e s : * TSHE-ABAM ** THPL-TSHE(PSME) TSHE (unmarked) 5§ axis i 200 112 69 Figure 30. Polar o r d i n a t i o n of samples in the TSHE zone with TSHE/GASH endpoints chosen for high WSI,radiation. B low WSI, r a d i a t i o n O O O O TSHE/GASH O'.TSHE/GASH/BENE/ACTR O O high WSI, r a d i a t i o n TSHE/POMU axis 1 " Figure 31. Polar o r d i n a t i o n of samples i n the TSHE zone with TSHE/GASH/BENE/ACTR endpoints f o r high WSI.radiation. B low WSI, r a d i a t i o n 0 ° A 0 high WSI, r a d i a t i o n O o axis 1 113 in t e r r e l a t e d , the spread of plots forms a diagonal across the ordination axes from high to low values of WSI and potential radiation. A good separation of habitat types resulted. If one assumes, however, that the plot order represents a continuous gradient of these two properties, the TSHE/GASH-BENE/ACTR type is mistakenly given an intermediate position when in fact i t has the highest o v e r a l l WSI and radiation values. Had two TSHE/GASH-BENE/ACTR samples been chosen, the ordination results would have been quite d i f f e r e n t , as shown in Figure 31. This i l l u s t r a t e s the danger of interpreting ordinations in terms of pairs of environmental gradients when species composition is the actual parameter used for comparison. It also shows the importance of the correct choice of gradient endpoints i f one is to obtain satisfactory r esults. The complex moisture-topography influences in the TSHE/GASH-BENE/ACTR habitat type complicate comparison of these three ecosystems along simple gradients. In terms of separating the types on the basis of species composition, polar ordination produced good r e s u l t s . One exception to this favorable grouping occurred for a sample with high T i a r e l l a tr i f o l i a t a and low Gaulther ia shallon cover compared to other TSHE/GASH-BENE/ACTR plots, as well as several minor species common to the TSHE/POMU samples. It was not re-c l a s s i f i e d because i t has no Polyst ichum munitum. 5.4.3.3. Relationship of ordination axes to environmental gradients Calculated environmental parameters are plotted on the DECORANA understory ordination in Figures 32a through 32c. 114 F i g u r e 32. Environmental v a r i a b l e s p l o t t e d on a DECORANA understory o r d i n a t i o n ( F i g u r e 2 6 ) - TSHE zone. a. Water S t r e s s Index (WSI) 4D° • !• A A * y ± © s t r e s s < i growing season O no s t r e s s s t r e s s < i s t r e s s >i growing season b. P o t e n t i a l S o l a r R a d i a t i o n • A A A • • U 1 2 O > 1 9 0 xlO cal/cm /yr A © 1 7 0 - 1 9 0 • < 1 7 0 A c. Average p r e c i p i t a t i o n (May - September) O 14 - 18 cm A A A © 19 - 2 5 cm • # 2 6 - 31 cm o g ° o o a . 115 D e f i n i t e trends in Water Stress Index (WSI), potential solar radiation and p r e c i p i t a t i o n values were observed between the three types. No trends were observed between plots in comparisons of elevation, temperature and AWSC. Samples in the TSHE/GASH-BENE/ACTR type had the highest WSI and solar radiation values and the lowest growing season p r e c i p i t a t i o n . For samples in the TSHE/POMU type, no water stress was predicted and solar radiation input was low, with one exception in either case. P r e c i p i t a t i o n was intermediate with respect to other types. The TSHE/GASH type was intermediate in WSI and solar radiation, but was highest in r a i n f a l l . While these three environmental factors do d i s t i n g u i s h between types, they are not strongly related to a single ordination axis. It i s d i f f i c u l t to relate differences in vegetation composition to a single moisture gradient (as in the PSME zone) in this set of samples. The wettest type from the standpoint of available s o i l moisture i s the TSHE/POMU type. A l l samples are from si t e s on lower to lower middle slope positions with favorable texture and rooting depth receiving seepage from upslope. This favorable topographic moisture position i s aided by moderate to high p r e c i p i t a t i o n ( r e l a t i v e to the zone) and low potential solar radiation which, in combination, help to reduce evapotranspiration demand (and WSI values). The type that appears d r i e s t on the basis of an examination of environmental variables is the TSHE/GASH-BENE/ACTR habitat type. WSI i s highest in t h i s type because of shallow s o i l s , slopes of mainly southerly exposure and low p r e c i p i t a t i o n . R a i n f a l l is the lowest in the zone because a l l samples f a l l within the rainshadow area in the Oyster River v a l l e y . Many of 116 the species present in this type are more commonly associated with the PSME zone. Such species include: Symphoricarpos mollis and albus, Rosa qymnocarpa, Amelanchier a l n i f o l i a , Pachyst ima  myrsinites, Festuca occidentalis, Campanula s c o u l e r i , Arenaria  macrophylla, and T r i e n t a l i s l a t i f o l i a . The microtopography associated with these plots complicates this i n t e r p r e t a t i o n . The underlying bedrock is gently undulating and close to the surface, with an overlying veneer of morainal deposits of varying depth. Observations during sampling showed that the "dry s i t e " indicators were found on deeper t i l l deposits. Where s o i l i s only a few centimetres thick, seepage water from upslope becomes close to the surface. In areas with shallow s o i l , "wet s i t e " indicators, such as Achlys t r i p h y l l a , Adenocaulon b i c o l o r ,  Lactuca muralis, T i a r e l l a t r i f o l i a t a , and T r i l l i u m ovatum, were found and are apparently able to survive from seepage water, despite summer droughts. Because microtopography changed over short distances, these two "sets" of species were often found immediately adjacent to each other. This "mixed" group of indicator species complicates any mathematical treatment of the data; consequently, in the understory ordination, t h i s type' has species that cause i t to ordinate in an intermediate position on axis 1 and at the dry extreme on axis 2. It is d i f f i c u l t to place the TSHE/GASH type to the right or l e f t of the TSHE/GASH-BENE/ACTR because of the l a t t e r ' s complex microtopography. In terms of macroclimate the former is less dry, yet in topographic moisture i t appears to be d r i e s t of the three. Other unique species in the Oyster River samples include Pedicular is  racemosa, Cornus canadensi s, Gaultheria o v a t i f o l i a , and 117 Vacc inium alaskaense. Plots at the same elevation further south did not have these species that are generally found at higher elevations. The more northerly l a t i t u d e appears to account for this difference. While temperature showed no relationship to habitat type d i s t r i b u t i o n on either the f u l l species or understory ordination, i t did show a relationship to the f i r s t axis of the overstory ordination, as shown in Figure 29. Though weak, the trend i s from cooler temperature on the l e f t to warmer on the right. Two plots with d e f i n i t e cold a i r drainage influence, not accounted for by extrapolated temperature data, occupy the l e f t side of the f i r s t axis. This relationship is too weak to draw any meaningful conclusions. It i s s i g n i f i c a n t to note that WSI gave a better o v e r a l l c o r r e l a t i o n with types than AWSC, temperature, p r e c i p i t a t i o n or potential solar radiation alone. This suggests that i t may be a useful integrator of these factors for s i t e comparison. In summary, a l l ordinations except overstory alone produced a good separation of habitat types into more or less d i s t i n c t c lusters in the ordination diagrams, supporting their i d e n t i f i c a t i o n as separate c l a s s i f i c a t i o n units. Furthermore, these three units could be related to differences in moisture and solar radiation using measured and/or extrapolated data. WSI appeared to be a good integrator of moisture factors. Overstory ordinations indicated no r e l a t i o n s h i p to habitat types but suggested some relat i o n s h i p to temperature. The complex nature of the gradient relationships and species composition of types make correlations to single ordination axes d i f f i c u l t . Choice of 118 polar ordination endpoints and careful interpretation of the re s u l t s were shown to be of c r i t i c a l importance. 5.5. Description and analysis of the Abies amabilis - Tsuga  heterophylla (ABAM-TSHE) vegetation zone 5.5.1. Description of the vegetation series Five vegetation series are represented by the 32 samples in the ABAM-TSHE zone. They are distinguished from each other on the basis of the r e l a t i v e abundance of regeneration of four tree species: Abies amabilis, Chamaecyparis nootkatensi s, Thuja  p i i c a t a and Tsuga heterophylla. The two most widespread series are the ABAM and TSHE-ABAM. The ABAM series defines those habitats where Abies amabilis is nearly the sole tree species reproducing successfully. Seedlings are usually very abundant. This series occurs on well-drained s i t e s at middle and higher elevations within the zone, generally above 800 m. It i s more common and extends lower on cooler, northerly exposures than on southerly exposures and i s common in valley bottoms and areas with pockets of cold a i r drainage. Pseudotsuga menziesi i i s usually not abundant as a serai overstory dominant; The TSHE-ABAM series i s found on well-drained s i t e s at lower elevations within the zone. It i s more common on south aspects than the ABAM series. On these s i t e s , Tsuga heterophylla i s reproducing more successfully than Abies amabilis. This series indicates the warmest conditions within the zone. Pseudotsuga menziesii i s frequently found as the dominant serai 119 overstory species. Thuja p i i c a t a varies from minor to co-climax status. Another series common to warmer', lower elevation stands in the ABAM-TSHE zone is the ABAM-TSHE(THPL). On these s i t e s , Abies  amabi1i s and Tsuga heterophylla are both reproducing successfully but amabilis f i r i s in greater abundance. Thuja  p i i c a t a i s common as a minor or co-climax component. This series generally indicates moister habitats than the TSHE-ABAM series. The complementary series to the ABAM-TSHE(THPL) found at higher elevations in the zone i s the ABAM-TSHE(CHNO). It i s defined as an area supporting reproduction of Abies amabilis in greater abundance than Tsuga heterophylla, with Chamaecyparis  nootkatensis present with greater cover than Thuja p i i c a t a . This series occupies moister, less well-drained s i t e s than the ABAM series found at similar elevations. The CHNO series is defined where Chamaecyparis nootkatensis is the most abundant or nearly sole species reproducing successfully. Abies amabilis i s usually very poorly represented, as are other species. A common c h a r a c t e r i s t i c of t h i s series is a shallow s o i l layer with ponded water during some period of the year. This series also appears to be more common on higher elevation southerly exposures. Because of the shallow s o i l s and southerly exposures, this series may also experience moisture stress during summer droughts. 5.5.2. Description of habitat types Site c h a r a c t e r i s t i c s for samples in each habitat type of the ABAM-TSHE zone are presented in Appendix V, Tables A17 -120 A22. Association tables for each type are given in Appendix VI, Tables A33 - A38. Table VI summarizes the vegetation c h a r a c t e r i s t i c s for habitat types in thi s zone. 5.5.2.1. Chamaecyparis nootkatensis / Gaultheria shallon (CHNO/GASH) habitat type The CHNO/GASH type was found on upper elevation south-facing slopes. Sample plots range from 820-1060 m elevation in the south Nanaimo River v a l l e y . This type was also found on south-facing slopes at lower elevations (470-650 m) further north near the Tsable River. Parent material i s of c o l l u v i a l and morainal o r i g i n . S o i l s range in texture from gravelly loam to sandy clay and have moderately good to poor v e r t i c a l drainage. Bedrock within one metre or less of the s o i l surface i s common in several sample plots with one p i t having a s l i g h t l y compacted "C" layer at 40 cm. There i s a lack of a well-developed "H" organic layer on many s i t e s . Sampled stands are from 250-350 years old, yet the height of dominant trees i s less than 28 m, with one exception at 470 m in the Tsable River v a l l e y . Canopy density i s quite variable (67-93%), as i s basal area (59-113 m 2/ha). Tsuga heterophylla dominates the canopy layers with Pseudotsuga menziesi i . A si g n i f i c a n t component of Chamaecyparis nootkatensis is found in a l l layers. Thuja p l i c a t a i s also present in most stands. Abies  amabilis i s rare in the tree canopy, but occurs in variable amounts as seedlings. Chamaecyparis nootkatensis reproduction exceeds Thuja p l i c a t a on these s i t e s . The shrub layer is dominated by Gaultheria shallon (Figure 121 Table VI. Summary o f major species for habitat types i n the ABAM-TSHE vcoetatinn zone. TYPE: T R E E LAYER Abiu amabiZii Chamaecypa>iib nootkateniii Pieudotiuga me.nzi.UAA. Thuja plicata Tiuga heteAophylia Tiuga meAtemiana SHRUB LAYER BeAbeAi6 nenvoia GaultheAia ovatifiolia GaultheAia ihallon Oplopanax hoHAidum Rhododendron albi^lonxm RibeA lacuAtAe. P.ubu6 Apectabilii Vaccinium alaikacniz Vaccinium membAanaczum Vaccinium ovalifiolium Vaccinium paAvi^olium HERB LAYER Achlyb tAiphylla Ade.no caulon bicolon. AthynJjum {i&Lx-iemina Blzchnum ipicant Chimaph-ila menzieAii Clvimaphila umbellata Clintonia uni£lo>ia CoinuA canadensis VntjopteAiM au&tAiaca GoodyeAa dblongi&olia GymnocaApium aWyopteAij, Lactuca muAaliA Linnaea boizaliA LiAteAa cawiina LUteAa coidata Luzula ipp. Lycopodium clavatum Omohkiza chiZeniii Polyitichum munitum Vyhola izcunda Rubui ptdatiu, Stenant.hiim occidentals. StAeptoput, amplexi^oliuA Stneptoput KOiCUi StAeptopui AtAzptopoidte lixvieJUia lac-inXata TiaAeJLla tAi^oliata TniZtium ovatum VeAatAum veAide, Viola glabella Viola cnbicutata CHNO/ A3 AM/ SB AM/ ABAM/ ABAM/ ABAM/ GASH VAAL/ ACTR/ VAAL/ V A A L ( O V ) / OPHO VAPA T I T R STREPTOPUS RUPE P C 1 MC" PC MC PC MC PC MC PC MC PC MC I + . 0 I V 3. 8 IT 3. 7 V 4. 2 V 5. 3 V 8. 0 I V 4. .2 I I I 3. ,4 I +. 7 V 3. 4 V 5. 9 V 6. ,4 IV 7. 4 I V 5. 0 I I I 5. 0 I I I 2. .9 V 3. ,7 I I I . 3. .4 I I +. 9 I V 2. 6 V 4. .8 V 7. .4 I V 5. 6 V 7. 7 V 7. 8 V 6. 5 I I 2. ,0 I I 1. .6 V 5. 4 I +. ,0 I I I +, .7 I I I 1. ,0 I I I 2. 2 I +. .0 I I I 1. ,8 X +. .0 I +. 0 I + . 0 I I + . 7 V 8. .1 I I I .8 I +. 0 V 4. 7 I +. 0 I V 1. .7 I I + . ,0 I + . 0 I +. 0 I I I +. 0 I I I 1. .6 I I I 1. .0 I V 3. .5 I I 1. 2 V 4. 5 V 5. 8 I I 1. .7 I I I 2. .6 I + . 2 I I +. 0 I V 3. 0 I I 3. .1 V 2. .5 V 4. .0 I I 4. •0. V 3. .4 I I I 1. .9 V 2. .9 I I + . 1 V + . ,5 I I I 1. .6 V 4. .9 V 3. ,3 I 1. , 2 V 4. .9 I + . 0 I + . ,0 I I I 1. .0 I I + . 0 I + . 0 V 2. .9 I I 1. .6 I V .3 I 1. .2 I I 1. .2 V 1 .2 V 2. .3 I V +. .9 I I +. .0 V 3 .4 I V 3 .0 I I I 1. .0 I V 1. .1 I I I 2. .1 I I + .4 I I + , .0 I I 1. .7 I + . 0 I I I 1 .5 I I + .8 I V 1. .9 I I I 1. .1 • I +, .0 I +. .0 I I I 1. .0 r + .2 I V + .5 I I I 1 .0 I I +. .4 I .0' I + .0 I I I 4 .7 1 1 .5 I + .0 I V + .7 I + .0 I I I + .0 I V 1 .8 V 3 .4 I I I 1 .2 I V 2 .2 I I I 2 .2 11 1 .2 I I + .1 I I I + .4 I I I + .7 I I I + .0 I 1 .5 I I I + .7 I 1 .5 I I + .2 I I + .1 I + .0 I I I + .0 I + .0 I + .0 I I I 2 .1 I I I 1 .4 I + .0 I I + .2 V 1 .2 I I I + .0 I V 1 .3 I I I 1. .6 I I + . 0 V 3 .6 I I I + .0 I I I + .2 I I I + .5 V 2 .0 I V 1 .6 I V 1 .0 I I + .4 I V 2 .8 V 4 .1 I I I 1 .0 I + .O I I I 1 .1 I + .0 V 1 .2 I + .0 V 2 .0 I + .0 I V 1 .1 I + .0 V 2 .7 I + .0 I I I 3 .4 I I I 2 .0 I I I + .0 I I I + .2 V 2 .8 V 1 .2 I I + .0 V 1 .5 I I I 2 .2 V 3 .8 V 3 .2 'IV 1 .1 V 5 .8 I I + .0 I V + .7 V + .7 I + .0 V 1 .5 I + .0 I I + .1 I I I + .0 I + .2 I + .0 I + .0 I I I 1 .0 I + .0 I V 1 .3 I V 1 .4 I I + .6 I I + .7 Presence Class: I = 1-20%; II = 21-40%; III = 41-60%; IV - 61-80%; V = 81-100% Mean Coverage: Values given are averages of the following classes: + = trace; 1 = 0.1-1.0%; 2 = 1.1-2.2%; 3 = 2.3-5.0%; 4 = 5.1-10.0%; 5 = 10.1-25.0%; 6 = 25.1-33.0%; 7 = 33.1-50.0%; 8 = 50.1-75.0%; 9 = over 75.0% 122 33). Variable amounts of several Vacc inium species are present; the most constant being V. alaskaense and V. Parvifolium. V. membranaceum i s present in the three highest elevation stands. Chimaphila umbellata is the only constant species in the herbaceous layer. Linnaea borealis and Pyrola secunda have a presence of 67%. Gaultheria o v a t i f o l i a and Chimaphila menziesi i are in 50% of the samples. A l l other herbs are of scattered occurrence. Rhytidiopsis robusta i s the most common moss species present, generally covering from 15 to 30% of the ground surface. Hylocomium splendens is also common, with some Pleurozium shreberi. The lichen Alectoria sarmentosa i s a conspicuous epiphyte in a l l samples. Sample 1157 is most unlike the other samples, though i t appears to f i t best within t h i s type. The presence of several herbaceous species of s i g n i f i c a n t cover distinguishes i t from other samples: Campanula s c o u l e r i , Festuca o c c i d e n t a l i s , Lactuca  murali s, Madia sativa, Montia parvi f o l i a , and Polyst ichum l o n c h i t i s . This i s the lowest elevation sample within the Nanaimo va l l e y and borders the TSHE zone as these herbs, more common in warmer, drier areas, indicate. This type is most clos e l y related to the TSHE/GASH habitat type. The presence of Chamaecyparis nootkatensis, Vaccinium  alaskaense, Rubus pedatus and Rhytidiopsis robusta in the CHNO/GASH type indicate a cooler, wetter (poorer drained) environment than the warmer, lower elevation TSHE/GASH habitat. The evidence of poor v e r t i c a l drainage despite the otherwise dry nature of the CHNO/GASH s i t e s (thin s o i l , southerly aspect, steep slope) i s a peculiar a t t r i b u t e of this type which probably 123 F i g u r e 3 3 . Chamaecypafiis nootkatensis I GaultheJiMi shallon (CHNO/GASH) h a b i t a t type. F i g u r e 3 4 . Abies amabilis I VaccJjuum alask.ae.iuz - Vaccinium paAvi^olixm (ABAM/VAAL-VAPA) h a b i t a t type. 124 accounts for Chamaecyparis nootkatensis dominance. This species can tolerate these conditions and outcompetes Abies amabilis on these s i t e s . The CHNO/GASH type i s apparently of small areal extent within the study area. A similar type has not been described in the region. Franklin et a l . 's (1979) ABAM/GASH type occupies a similar topographic position with similar species composition except for substantially greater ABAM in the overstory and reproduction layers and very low (20%) Chamaecyparis nootkatensis presence. Their CHNO phase of the ABAM/VAAL habitat type does not appear comparable because i t has no Gaulther ia shallon, though i t may be related. 5.5.2.2. Abies amabilis / Vacc inium alaskaense - Vacc inium  parvifolium (ABAM/VAAL-VAPA) habitat type This habitat type was found on moderately-sloping, mid-slope, northerly exposures between 700 and 860 m in the Cameron, Nanaimo and Oyster River v a l l e y s . An additional stand was found in the Tsable River at 470 m. S o i l parent material i s mostly morainal with variable texture, moderate stoniness and moderate v e r t i c a l drainage. In the majority of samples, the s o i l layer i s less than 50 cm to bedrock. A l l s o i l s have a f a i r l y well-developed humified organic layer. The overstory dominants average 37 m in height in stands from 225 to 300 years old. Canopy closure i s approximately 95% in most samples. Tsuga heterophylla and Pseudotsuga menziesi i are constant overstory dominants. Western hemlock remains a constant in a l l canopy layers, but Douglas-fir i s present only in the upper positions. Thuja p l i c a t a i s present in a l l samples, 125 forming a s i g n i f i c a n t portion of the Al and A3 layers in several stands. Abies amabilis i s most frequently in the understory (A3) and seedling layers, where i t i s less abundant than western hemlock seedlings. The shrub layer i s dominated by Vacc inium alaskaense and Vaccinium parvifolium. Vaccinium membranaceum and Vacc inium  ovalifolium are common to rare. Berberis nervosa i s present in 85% of the samples but has very low cover. A l l other shrubs are of minor importance. Linnaea borealis is a constant in the herb layer throughout the type but varies in cover from a trace to 20%. Other constant herbs are: Chimaphila menziesii, Goodyera o b l o n g i f o l i a , and Vi o l a orbiculata. Chimaphila umbellata, L i stera cordata, Cornus canadensis, and Rubus pedatus are also common. Total cover by the herb layer i s generally small with many species of minor to incidental occurrence (Figure 34). The moss layer i s dominated by a dense carpet of Rhytidiopsis robusta that ranges from 35-90% cover for the sample p l o t s . This type characterizes zonal ecosystems and has an intermediate relationship to the CHNO/GASH and ABAM/ACTR-TITR types. The former is cooler and d r i e r , the l a t t e r warmer and wetter. An Abies amabilis - Vacc inium alaskaense association i s recognized as the zonal plant community for the Abies amabilis zone (wet CWH biogeoclimatic subzone) by a l l authors in t h i s region. Krajina's (1969) biogeocoenosis #34 i s included in the ABAM/VAAL-VAPA habitat type. The zonal ecosystem of the submontane wetter maritime CWH variant of Klinka e_t a l . (1979) is the Rhytidiadelphus - Red Huckleberry - Alaska Blueberry -126 Western Hemlock biogeocoenotic type. This type i s equivalent to the ABAM/VAAL-VAPA habitat type, as i s Kojima's (1971) Vaccinium  alaskaense association. The zonal types of Orloci (1961) appear to be somewhat d i f f e r e n t . Species used to characterize his two Vaccinium alaskaense forest types (Clintonia u n i f l o r a , Plagiothecium undulatum, and Acer circinatum) are either absent or not abundant in thi s habitat type. The ABAM/VAAL-VAPA type appears quite similar to the ABAM/VAAL, BENE phase of Franklin (1966) and Franklin et a_l. (1979). Some notable differences in species composition in these types include Xerophyllum tenax, T i a r e l l a u n i f o l i a t a and higher cover of Clintonia u n i f l o r a than in the present study. The Abies amabilis - Vaccinium alaskaense - Cornus canadensis association of Dyrness et. a_l. (1974) i s also s i m i l a r . 5.5.2.3. Abies amabilis / Achlys t r i p h y l l a - T i a r e l l a t r i f o l i a t a (ABAM/ACTR-TITR) habitat type The ABAM/ACTR-TITR habitat type was found on steep (50-80%) c o l l u v i a l slopes of variable exposure on lower middle to upper slope positions. Samples in the Tsable and Oyster River valleys range from 560 to 620 m in elevation. Stands are at higher elevations in the Cameron River valley (780-1085 m) on southwest aspects. S o i l s are loamy with a high coarse fragment content which, along with steep slope, make these s i t e s very well-drained. E f f e c t i v e rooting depth i s 70-120 cm. The humus layer i s generally not well-developed. Sample stands are between 250 and 350 years old, with an overstory dominated by Pseudotsuga menziesi i and Tsuga 127 heterophylla. Thuja p l i c a t a and Abies amabilis are present in lesser amounts in the overstory. Amabilis f i r and western hemlock dominate the reproduction, and vary in their r e l a t i v e abundance throughout the type. The most c h a r a c t e r i s t i c feature of the shrub and herb layers i s their sparseness (Figure 35). The steep, unstable slopes indicate that periodic s l i d i n g of the surface layer has prevented establishment of many plants. As a r e s u l t , l i t t e r covers most of the slope surface. Further evidence i s the frequency of exposed tree roots. The only constant shrub species is Vaccinium parvifolium, found only as a low shrub. Berberis  nervosa, Vacc inium alaskaense, Vaccinium membranaceum and Rosa  gymnocarpa have 40-60% presence in the samples. The herb layer includes Achlys t r i p h y l l a , T i a r e l l a  t r i f o l i a t a , T i a r e l l a l a c i n i a t a and Chimaphila menziesi i as constant species. V i o l a orbiculata, Polyst ichum munitum, Lactuca  muralis, and T r i l l i u m ovatum are also common (80% presence), but only in trace amounts. Even the moss layer i s depauperate compared to other types in the zone. Rhytidiopsis robusta, commonly blanketing other types, i s reduced to 1-15% cover. This type i s apparently common throughout the study area, yet i t s status as a genuine habitat type may be questionable because of the role of periodic disturbance of the s o i l surface in maintaining the depauperate f l o r a . It may be more appropriately considered as a steep phase of the ABAM/VAAL-VAPA habitat type. It i s quite similar to two of the u n c l a s s i f i e d samples (1131, 1146) with depauperate understories in the TSHE zone. 128 Figure 35. Abies a m a b i L U / Aattty-i tfvipkyZZa - TioJittta thifaotiata. (ABAM/ACTR-TITR) habitat type. 129 There i s no apparent equivalent to this type in the l i t e r a t u r e . Kojima's (1971) Achlys t r i p h y l l a variant of the Achlys t r i p h y l l a - Polyst ichum munitum association is found on gentler slopes and morainal parent material with substantial differences in dominant species. The ABAM/ACTR association of Dyrness et a l . (1974) occupies a similar moisture position but has a much more well-developed herb layer and many differences in species. 5.5.2.4. Abies amabi1i s / Vacc inium alaskaense / Streptopus spp. (ABAM/VAAL/STREPTOPUS) habitat type The ABAM/VAAL/STREPTOPUS habitat type was found in the Cameron, Tsable and Oyster River valleys on northeasterly exposures between 790 and 1040 m elevation. It also occurred at 500 m near the Cameron River under the influence of cold a i r drainage. S o i l s are derived from morainal, c o l l u v i a l and glacio-f l u v i a l parent materials. S o i l textures range from gravelly, sandy loams to clay loams with variable stoniness. Sites are moderately well- to imperfectly-drained. A l l samples have a well-developed and frequently thick (up to 17 cm) H organic horizon. The overstory dominants range from 240 to 300 years old and average 39 m in height. The Al layer i s dominated by Tsuga  heterophylla along with Abies amabi1is and Pseudotsuga menziesi i which vary in importance between samples. The lower canopy (A3) is dominated by amabilis f i r and western hemlock. Amabilis f i r seedling cover i s very abundant, and may number as high as 50,000 stems per hectare in some samples. 130 The shrub layer i s dominated by a mixture of Vacc inium  alaskaense, Vaccinium parvi folium and Vacc inium ovalifolium. Vacc inium membranaceum and Berberis nervosa are also present in half of the samples. Cover of shrubs varies from 4-45% (mean 21%) . The herb layer i s the most diagnostic feature of t h i s type, containing a r i c h mixture of species, though cover of a single species rarely exceeds 5% (Figure 36). Total herb cover varies between 20 and 59% (average 33%). Several species in the L i 1iaceae family are common: Streptopus amplexi f o l i u s , Streptopus roseus, Streptopus streptopoides, T r i l l i u m ovatum, and Stenanthium occidentale. Abundance of the three Streptopus spp. varies, so the genus was used in the naming of the habitat type rather than a single species. Constants include Achlys  t r i p h y l l a , T i a r e l l a t r i f o l i a t a , T i a r e l l a l a c i n i a t a , Pyrola  secunda and Streptopus amplexifolius. Rubus pedatus i s also common, but varies in cover. Species d i v e r s i t y in the herb layer can be quite high, as shown in plot 1173 with 35 species. The moss layer i s dominated by Rhytidiopsis robusta with cover from 10-60%. Rhytidiadelphus loreus and Dicranum spp. are also common. The ABAM/VAAL/STREPTOPUS habitat type occupies wetter and generally cooler sites than the t y p i c a l ABAM/VAAL type, and is found on lower elevation (less wet and cold) s i t e s than the ABAM/VAAL(OV)/RUPE type. A similar ecosystem is indicated by the Rhyt idiadelphus loreus - [Eurhynchium] S t o k e s i e l l a stokesi i -Blechnum spicant - Polystichum muni turn - Streptopus roseus  Vacc inium o v a l i folium - Vaccinium alaskaense - Pseudotsuga 131 menziesi i - Abies amabilis - Thuja p i i c a t a phytocoenosis (biogeocoenosis #32) of Krajina (1969). Thuja p i i c a t a , however, is a rare constituent in the ABAM/VAAL/STREPTOPUS habitat type. The Streptopus roseus - T i a r e l l a t r i f o l i a t a portion of the edatopic g r i d of Klinka (1977) describes similar habitats. There i s no apparent equivalent in Kojima's (1971) study. The orthic Blechnum forest type of Orloci (1961) is the closest r e l a t i v e in his c l a s s i f i c a t i o n , though i t has Rubus s p e c t a b i l i s and lacks the high cover of Achlys tr i p h y l l a found in the present study plo t s . The ABAM/STRO association of Franklin (1966) as described in Franklin and Dyrness (1973) i s e s s e n t i a l l y equivalent. The Abies amabilis - T i a r e l l a uni f ol iata association of Franklin e_t a l . (1979) and Dyrness et a l . (1974) i s also quite c l o s e l y related. Achlys t r i p h y l l a and T i a r e l l a dominate the herb-rich understories. Vacc inium membranaceum and Streptopus roseus are abundant as in the present type. The major difference in the Oregon and Washington types i s that they have a less developed shrub layer. The ABAM/TIUN association of Franklin (1966) i s more similar to th i s study than the same type of Dyrness e_t a l . (1974) because of greater Vaccinium parvi folium and less Acer  circinatum. The ABAM/ACTR association of the l a t t e r authors i s also related to the ABAM/VAAL/STREPTOPUS habitat type. 132 5.5.2.5. Abies amabi1i s / Vacc inium alaskaense - Vacc inium  ovalifolium / Rubus pedatus (ABAM/VAAL(OV)/RUPE) habitat type The ABAM/VAAL(OV)/RUPE habitat type occupies cold, wet upper elevation sites in the ABAM-TSHE zone and the adjacent ABAM-TSME vegetation zone. A l l samples are between 900 m and 1000 m elevation on northerly aspects. Parent material is of morainal or c o l l u v i a l o r i g i n occurring as a blanket or thin veneer over bedrock. Rooting depth and coarse fragement content are var i a b l e . Most s o i l s have s i l t loam to clay loam textures and are frequently imperfectly- to poorly-drained. Sampled stands are a l l between 240 and 290 years old, and are dominated by Tsuga heterophylla and Tsuga mertensiana. Western hemlock represents more than 50% of the Tsuga dominants and is also more abundant in lower canopy and seedling layers. Abies amabilis i s the most successfully reproducing species. Chamaecyparis nootkatensis i s also common in a l l canopy layers. Pseudotsuga menziesi i and Thuja p l i c a t a are generally rare. Shrub cover is variable, ranging from a high of 62% to as low as 2%. Vaccinium alaskaense and Vacc inium ovalifolium dominate th i s layer with common but not constant occurrence of Rhododendron a l b i florum and Vacc inium membranaceum (Figure 37). Menziesia ferruginea is present in thi s type with low constancy. Vacc inium parvifolium i s rare or absent in most samples. Rubus pedatus and Pyrola secunda are the most constantly occurring herb species, though with variable but usually sparse cover. Most other species occur only sporadically with low abundance. The moss layer i s d i s t i n c t l y dominated by Rhytidiopsis robusta with 30-90% cover on a l l p l o t s . 1 3 3 Figure 37. Abies amabilis I VaccsLniuni alaskaense - Vaccinium ovali^olium / Rubus pedatus (ABAM/VAAL (OV)/RUPE) habitat type. Figure 38. Abies amabiLis I Oplopanax kowiidum (ABAM/OPHO) habitat type. 134 This type i s cooler and wetter than any of the other habitat types in the zone. The most similar type i s the ABAM/VAAL/STREPTOPUS. This habitat type appears most similar to types described in B. C. for the Mountain Hemlock biogeoclimatic zone. Krajina's Rhytidiopsis robusta - Dicranum  fusescens - Rubus pedatus - Vaccinium alaskaense - Vaccinium  ovalifolium - Abies amabi1is - Tsuga mertensiana phytocoenosis (biogeocoenosis #45) i s c l o s e l y related. The zonal ecosystem for the eastern Vancouver Island montane forested Mountain Hemlock variant (Klinka e_t a_l. 1979) i s a Vacc inium o v a l i f olium  Vaccinium membranaceum - Abies amabi1is - Tsuga mertensiana association. The mesic type given by Klinka (1977) i s described as being dominated by Vacc in ium alaskaense or Vaccinium  ov a l i folium and Rubus pedatus. Both of these associations resemble the ABAM/VAAL(OV)/RUPE'habitat type. The Abies amabilis  Menziesia ferruginea habitat type of Franklin et al. (1979) appears most similar to t h i s type. In their association, Rubus  pedatus is abundant along with similar dominant shrubs and herbs, including Rhododendron a l b i florum. Franklin (1966) recognized a similar ABAM/MEFE type as t r a n s i t i o n a l between the Amabilis F i r and Mountain Hemlock vegetation zones because of the c h a r a c t e r i s t i c mixture of mountain and western hemlock, with the l a t t e r in greater abundance. Menziesia ferruginea i s common in the ABAM/VAAL(OV)/RUPE habitat type, though not a constant dominant. Other than t h i s difference, the habitats and species are very s i m i l a r . Other clo s e l y related types of Franklin et a l . (1979) include the ABAM/RHAL habitat type (at higher elevations) and the ABAM/VAAL habitat type, RUPE phase in cooler, moister 135 habitats than t y p i c a l Vacc inium alaskaense associations. 5.5.2.6. Abies amabilis / Oplopahax horridum (ABAM/OPHO) habitat type The ABAM/OPHO habitat type occurs on well-drained a l l u v i a l materials in valley bottoms with a high, free-flowing water table. It i s also found on lower slopes that have free-flowing seepage water where i t often forms fingers upslope in narrow ravines. The two stands sampled are at approximately 800 m elevation in the King Solomon basin of McQuillan Creek, a tributary of China Creek. Normally, two plots would not be considered s u f f i c i e n t for defining a separate type; however, th i s type was observed elsewhere in higher elevation valley bottoms and lower slopes and i s quite e a s i l y recognized. The two plots represent the two topographic positions that the ABAM/OPHO type occupies. Both have s o i l s with sandy texture and high coarse fragment content. Productivity on t h i s habitat type i s excellent. On both s i t e s (165 and 195 years old) the dominant trees are over 50 m t a l l with an average basal area of 115 m2/ha. The overstory i s dominated by Abies amabilis, with Tsuga heterophylla as a co-dominant. Regeneration i s solely amabilis f i r except for western hemlock on rotting wood. The shrub layer i s dominated by Oplopanax horridum, which occurs in large patches (Figure 38). Plot 1171 shows l i t t l e Oplopanax horridum in the vegetation table because the sample was taken adjacent to, but not in, one of these patches. Rubus  s p e c t a b i l i s and Ribes lacustre may also be present. 136 The herbaceous layer i s lush, and includes the following species in order of abundance: T i a r e l l a t r i f o l i a t a , Achlys  t r i p h y l l a , Polyst ichum munitum, Athyrium f i l i x - f e m i n a , Streptopus roseus, Streptopus amplexifolius, T i a r e l l a l a c i n i a t a ,  T r i l l i u m ovaturn, Montia s i b i r i c a , and Osmorhiza c h i l e n s i s . Gymnocarpium dryopteris and Trautvetteria c a r o l i n i e n s i s are also abundant on the a l l u v i a l s i t e . Constancy and abundance data are d i f f i c u l t to interpret with only two samples. Rhizomnium  glabrescens and Rhyt idiadelphus loreus are the most common mosses present. The ABAM/OPHO habitat type i s similar to the Plagiomnium  insigne - Leucolepsis menziesi i - Polystichum munitum - Rubus  sp e c t a b i l i s - Ribes bracteosum - Oplopanax horr idum - Abies  amabi1i s - Picea sitchensis - Thuja p i i c a t a phytocoenosis (biogeocoenosis #31) of Krajina (1969). The Oplopanax horridum -Adiantum pedatum type of Orloci (1961) and Kojima (1971) is also equivalent. Klinka (1977) does not l i s t an Oplopanax horridum association for the montane wet Coastal Western Hemlock zone, but has a similar Streptopus roseus - Oplopanax horr idum association for the Mountain Hemlock zone, on si t e s with imperfect to poor drainage and a mesotrophic to eutrophic nutrient regime. The ABAM/OPHO habitat type appears i d e n t i c a l to the ABAM/OPHO habitat type of Franklin et a l . (1979). These authors also found the type on both valley bottoms and lower slopes and note the high productivity of the type. A l l of the above authors l i s t Thuja p l i c a t a as a c h a r a c t e r i s t i c species. Cedar was not found in the present study plots, though i t i s evidently common elsewhere in t h i s type. A related community in 137 the western Oregon Cascades i s the CHNO/OPHO association of Dyrness et a l . (1974). 5.5.3. Ordination results 5.5.3.1. DECORANA A DECORANA ordination of 31 samples in the ABAM-TSHE zone is presented in Figure 39. Octave and downweighting data transformations were used. Similar results were obtained using a l l species and understory alone, with habitat types tending to form d i s t i n c t c l u s t e r s in both. The f i r s t axis has been interpreted as a moisture gradient. The CHNO/GASH and ABAM/OPHO habitat types occupy the dry and wet extremes of axis 1, respectively. The second axis has been interpreted as a temperature gradient. The highest elevation plots in the zone (above 900 m) from the ABAM/VAAL(OV)/RUPE habitat type occupy the upper extreme of thi s axis. At the opposite extreme, plots from the ABAM/ACTR-TITR type form a grouping. This type is common at lower elevations, though some exceptions are found on high elevation southerly exposures. A DECORANA ordination of tree species produced a poor separation of vegetation series (Figure 40). Tree data used for the ordination did not include the seedling layer, which i s used for defining the series; consequently, the ordination only shows the degree to which overstory in these samples is a r e f l e c t i o n of the potential climax tree species. Serai overstories were less common in the ABAM-TSHE zone than in the TSHE zone, where serai Douglas-fir predominates. There is a separation of the 1 3 8 F i g u r e 39 . DECORANA o r d i n a t i o n of samples i n the ABAM-TSHE zone - h a b i t a t types. 2 4 0 A A A A A CHNO/GASH O ABAM/VAAL-VAPA • ABAM/ACTR-TITR • ABAM/VAAL/STREPTOPUS A ABAM/VAAL(OV)/RUPE ^ ^7ABAM/0PH0 o a O • • • o o o o o ^ 7 • • a x i s 1 2 8 0 F i g u r e 40- DECORANA o r d i n a t i o n of samples i n the ABAM-TSHE zone (overstory only) - vege t a t i o n s e r i e s . 2 2 0 • • CHNO O • ABAM-TSHE(CHNO) O ABAM • • A TSHE-ABAM ^ ABAM-TSHE(THPL) O O • . O A O ^  A A o o o o a x i s 1 28° 139 ABAM and ABAM-TSHE series along the f i r s t axis. The three series with Chamaecyparis nootkatensis and Thuja p i i c a t a are somewhat separated along the second axis. When superimposed, these two separations become obscure. A clump of samples representing a l l of the series i s found in the centre of the axes; consequently, overstory composition appears to be a f a i r l y poor overal l indicator of series p o t e n t i a l . Environmental correlations with either axis were unsuccessful. 5.5.3.2. Polar Ordination A polar ordination was performed with endpoints chosen to represent moisture and temperature gradients based on DECORANA res u l t s . Similar data transformations were used. The o v e r a l l configuration of plots is quite similar to DECORANA, though the cl u s t e r i n g of types i s not as d i s t i n c t (Figure 41). 5.5.4. Relationship of ordination axes to environmental gradients Comparison of environmental data with the DECORANA ordination produced mixed r e s u l t s . None of the plotted variables showed any strong trends associated with either axis. The WSI values are highest for samples in the ABAM/VAAL-VAPA habitat type at the centre l e f t of the axis. Predicted stress i s rare in the CHNO/GASH type at the far l e f t ; however, a l l plots have high solar radiation values and three of the f i v e are on ridgetops with the drying influence of wind exposure. When a r b i t r a r i l y d i v i d i n g axis 1 in half, 59% of the samples on the l e f t show some stress compared to 21% on the right (Figure 42a). For t h i s 140 F i g u r e 41 . P o l a r o r d i n a t i o n of samples i n the ABAM-TSHE zone-h a b i t a t types. 70 ¥ CN IC •r-K • CHNO/GASH • O ABAM/VAAL/VAPA • ABAM/ACTR/TITR O • ABAM/VAAL/STREPTOPUS A ABAM/VAAL(OV)/RUPE ABAM/OPHO • • ' o o Q • o 8 • D o o • • \ 7 \ 7 • • • ± ^ a x i s 1 90 141 Figure 42 . Environmental variables plotted on a DECORANA ordination of samples i n the ABAM-TSHE zone. a. Water Stress Index (WSI) • some stress O no stress * high water table Q Q Q O O O 10/17=59% • • n • o o • o 3/14=21% b. Elevation O o o o o o o o o • • o o o • • > 9 0 0 m O < 9 0 0 in X7 Q • 142 same s p l i t , the average AWSC is 9.4 cm and 12.6 cm for the l e f t and righ t , respectively. The range, however, is approximately 5-20 cm for both halves, showing the v a r i a b i l i t y when considering t h i s single factor. The elevation trend associated with temperature along the second axis has several exceptions due to the complicating influences of exposure and cold a i r drainage (Figure 42b). Extrapolated average growing-season temperature showed no s i g n i f i c a n t trend along this axis. The extrapolation of temperature (and precipitation) data for high elevation s i t e s from low elevation c l i m a t i c stations is tenuous. In addition, average growing-season temperature may not be the best measurement for comparing temperature differences between these s i t e s . Maximum and minimum temperatures and/or temperatures during other portions of the year may be of greater s i g n i f i c a n c e . 5.6. Transect Comparison The establishment of transects sampled at regular elevational intervals allowed comparison of the d i s t r i b u t i o n of vegetation c l a s s i f i c a t i o n units in two r i v e r valleys with d i f f e r e n t climatic influences, yet similar aspect and elevational ranges. The South Nanaimo River valley i s in a pronounced rainshadow, making i t quite dry. The Cameron River valley i s farther north and west and is under the wet,west coast influence extended inland by the Alberni i n l e t . The Cameron valley bottom is also much broader at the point of sampling than the South Nanaimo, as i l l u s t r a t e d in Figures 43 and 44 (also see Figure 4 3 . Distribution of vegetation in the South Nanaimo River valley, NANAIMO RIVER TRANSECT a i e i a t l e e ( m l 1 1 0 0 — 950 — 800 650 500 <! ABAM-TSME ABAM-TSHElCHNO r ._.„ _-„_ " 11 M*X"JSHE"QHPQ ~ ^ ~ ~ ^ ^ J * I o n t TSHE-ABAM ~ ~ ~ ~ TSHE SW~ » p l o t l o c a t i o n r e p e t a t l o n z o n e s v e g e t a t i o n s e r i e s '»• h a b i t a t t y p e s 1 J CHNO ABAM- TSHE M ^ - ^ ABAM:TSHE [C_HNO)_ CHNO i « , — . TSHE > ^ TSHE ABAM-TSHE(THPL| NE s c a l e - 1cm = 200m 1 s e e T a b l e VII U) F i g u r e 44. D i s t r i b u t i o n of v e g e t a t i o n i n the Cameron River v a l l e y . CAMERON RIVER TRANSECT ' plot location — vegetation zones vegetation series S W ' N E scale- 1cm = 200m 1soe Table VII 145 photos in Figures 2 and 3). Table VII l i s t s the habitat types occurring in these two transects. 5.6.1. Environmental c h a r a c t e r i s t i c s of the S. Nanaimo and Cameron ri v e r valleys Extrapolated temperature and p r e c i p i t a t i o n data, potential solar radiation, s o i l texture and calculated available water storage capacity (AWSC) and Water Stress Index (WSI) data were compared for the two va l l e y s . Average growing-season p r e c i p i t a t i o n i s greater in the Cameron valley than in the Nanaimo v a l l e y . WSI values were calculated for each sample. V i r t u a l l y no stress was predicted for the Cameron valley except on three plots, two of which showed signs of poor drainage. The t h i r d was on an upper southeast-facing slope with thin s o i l . In contrast, 2 to 3 months of moderate stress were predicted for four plots on middle to lower southwest slopes in the Nanaimo vall e y , with an additional 1 month of severe stress for a s i t e on thin c o l l u v i a l material. Stress was also predicted for two Chamaecyparis nootkatensis series plots where a compacted "C" layer or bedrock was found at 40 to 70 cm. There was also evidence of poor v e r t i c a l drainage on these plots; consequently, the predicted WSI values are questionable. Available water storage capacity (AWSC) varied considerably within both valleys as a result of s o i l texture, rooting depth and coarse fragment content. In the Nanaimo transect, lower elevation plots were on gravelly sandy loams to loamy sands. Higher elevation plots were on s i l t loams to loams with moderate stoniness, though often shallow (50-60 cm); consequently, they 146 Table VII. Habitat types present i n the Cameron and Nanaimo transects. No. Habitat type 6 T&VLQO. heAteJiophytta / GojjJLthoJxJjx i,haZton 8 Ti>uga hoXzAophylLa. / Poly&tichwm munitum 9 ChamazcyptvuJ) nootkatzvUAj, / GaaltheAxa. AkaJULon 10 Ab-cei amabAJM, / Vacctnium aZcu>kazn6Q. -Vaccintum paAvifaoltum 11 Abiej, amablUJi I Ac.hyJLi> PU.ph.ylla -JiaKoJJjx. tnJ-lolAjjuba. 12 AbieJ> amabiliXM / VaccAjuum a&ufcaettie / Stn.2.ptopuA spp. 13 Ab<Lej> amabttu, / Vaccuyitum alaAkaznAe. -V. ovalAifiolAjum / RubuA pudaJiu, 14 AbJJLb amabAjJJi / Optopanax koAAAidum 15 Abioj, ama.bJJUj, - TAuga nvmXQ.n6ia.na. / Rhododendron albi^lo/um 16 Unclassified samples i n the TSHE zone with depauperate understory on unstable surface materials Abbrev'n TSHE/GASH TSHE/POMU CHNO/GASH ABAM/VAAL-VAPA ABAM/ACTR-TITR ABAM/VAAL/STREPTOPUS ABAM/VAAL(OV)/RUPE ABAM/OPHO ABAM-TSME/RHAL 147 had higher AWSC (14-19 cm) compared to lower elevation plots (3-15 cm). In contrast, s o i l s in the Cameron transect were f i n e r -textured o v e r a l l . Higher elevation plots ranged from sandy clay loams to clay loams; lower elevation plots were loams to s i l t loams. The highest elevation plots on northeast aspects had shallow s o i l s with high coarse fragment content; therefore, there was no general trend in AWSC with elevation as in the Nanaimo valley transect. Potential annual solar radiation varies with l a t i t u d e , slope angle and aspect. Being "potential", i t does not take into account cloudiness differences between the two valleys, that also a f f e c t actual radiation. A comparison of nearby climatic stations shows that the Nanaimo valley receives greater radiation input than the Cameron. Total annual hours of bright sunshine for the Alberni Lupsi Cupsi station (AES) are 1599; the Nanaimo airport station (AES) has 1846 hours. Average d a i l y (calculated) solar radiation for May through September i s 421 and 428 cal/cm 2 for these two stations, respectively. While actual radiation in the valleys themselves w i l l be d i f f e r e n t , the general relationship should be sim i l a r . Average growing-season temperature is s l i g h t l y lower for the Cameron va l l e y . Combined with greater cloudiness, greater p r e c i p i t a t i o n and finer-textured s o i l s , the Cameron valley has a much cooler, moister environment with less potential evapotranspiration than the Nanaimo valle y . 148 5.6.2. Di s t r i b u t i o n of vegetation zones and series The TSHE, ABAM-TSHE and ABAM-TSME vegetation zones are represented in the two v a l l e y s . The TSHE zone occurs below approximately 700 m on the north-facing slope of the Nanaimo valley and approximately 730 m in the Cameron v a l l e y . It occurs about 80 to 100 m higher on south aspects in both v a l l e y s . The ABAM-TSHE zone occurs above the TSHE zone in both v a l l e y s . The ABAM-TSME zone occurs above the ABAM-TSHE zone on north aspects above approximately 1000 m in both transects. The presence of Tsuga mertensiana on south slopes i s uncommon in either v a l l e y . The d i s t r i b u t i o n of vegetation series has been plotted and reveals some important differences between the two va l l e y s . The TSHE series i s more common in the Nanaimo transect where i t . was found on lower, northeast aspects (<600 m) and on southwest aspects as high as 800 m (Figure 43). In the Cameron transect, t h i s series occurs on southwest aspects up to 860 m, but i s not found on north aspects, where i t i s replaced by the TSHE-ABAM series (Figure 44). This series i s found from the valley bottom (500 m) to 780 m on northeast-facing slopes in the Cameron, and occurs on the southwest-facing slope above the TSHE series at 940 m. It i s found only as a narrow band at 740 m on the northeast aspect of the Nanaimo transect. The ABAM-TSHE(THPL) series i s found above the TSHE-ABAM series on north aspects in both the Cameron (860 m) and Nanaimo (820 m) va l l e y s . In the ABAM-TSHE zone, the series begins at 820 and 860 m on northeast aspects in the Nanaimo and Cameron transects, respectively. The d i s t r i b u t i o n of series in the two valleys shows that the elevation range of Abies amabilis extends 149 much lower in the Cameron valle y . Both valleys have vegetation in the valley bottoms affected by cold a i r drainage. The ABAM-TSHE(THPL) series occupies t h i s position in the Nanaimo valley and the TSHE-ABAM series occupies the bottom of the Cameron va l l e y . Another s i g n i f i c a n t difference between the two transects i s in the presence of Chamaecyparis nootkatensis. The two series, CHNO and ABAM-TSHE(CHNO), are present in the Nanaimo transect on the southwest, facing slope above 800 m, and at 900 m on the northeast-facing slope, but are absent in the Cameron transect. Their occurrence coincides with shallow s o i l having r e s t r i c t e d v e r t i c a l drainage. At the same elevation with deeper, well-drained s o i l , Abies amabilis i s more abundant. Another possible explanation for the abundance of Chamaecyparis nootkatensis on south-facing slopes i s suggested in observations by Krajina (1969). Heavy snow accumulations may insulate the ground, keeping i t unfrozen in winter. This, he claims, promotes Abies  amabilis regeneration. The s o i l may be colder in winter on south aspects where snowpack i s reduced from solar melting and thus provide conditions under which yellow cypress is more able to compete. Chamaecypar i s nootkatensis i s three times as abundant in the Nanaimo valley as in the Cameron, while Thuja p i i c a t a i s equally abundant in both v a l l e y s . 5.6.3. Di s t r i b u t i o n of habitat types The two river valleys are quite d i f f e r e n t in the plant communities they support. The most s t r i k i n g difference i s in the 150 abundance of Gaultheria shal'lon. The Nanaimo transect has vigorous Gaultheria shallon habitat types along the entire southwesterly slope and the lower northeasterly slope (Figure 43). In contrast, no Gaultheria-dominated communities are found in the Cameron transect; only trace amounts are present in three of the samples (Figure 44). A second contrasting feature is the abundance of Polyst ichum munitum. The TSHE/POMU habitat type i s present on the lower slopes of both aspects in the Cameron valley transect, represented by three samples. This type was found only in a small area (sample 1141) at the base of the north-facing slope in the Nanaimo transect. A t h i r d difference i s in the presence of the ABAM/ACTR-TITR habitat type. This type i s not found in the Nanaimo transect but occurs at mid-slope on north aspects and on upper south-facing slopes in the Cameron v a l l e y . The drier climate and coarser-textured s o i l s of the Nanaimo valley most l i k e l y account for the abundance of s a l a l and lesser amounts of more mesophytic species l i k e swordfern and v a n i l l a l e a f . The fourth major contrast i s caused by differences in slope. The south-facing slope of the Cameron transect is much steeper than the south-facing slope of the Nanaimo transect (Figures 43 and 44). Materials are of c o l l u v i a l o r i g i n and there is some evidence of active s l i d i n g of surface materials; consequently, the understory i s f a i r l y depauperate throughout t h i s slope. Canopy density i s not great, so shading i s apparently not a factor. This contrasts with the abundant s a l a l cover on the southerly exposure of the Nanaimo transect. 151 The broader valley bottom of the Cameron river at the point of sampling supports two wet habitat types not found in the narrow portion of the Nanaimo va l l e y : ABAM/VAAL/STREPTOPUS and ABAM/OPHO. The two valleys support similar associations on middle and upper north aspects. The ABAM/VAAL-VAPA habitat type i s found at mid-slope positions, the ABAM/VAAL(OV)/RUPE type above t h i s , and an Abies amabi1is - Tsuga mertensiana - Rhododendron albiflorum association at the apex. The l a t t e r community is in the ABAM-TSME (Mountain Hemlock) vegetation zone, so i t has not been described for t h i s study. 1 5 It i s characterized by a generally dense .shrub layer dominated by Rhododendron a l b i florum, Vaccinium alaskaense, Vaccinium ovalifolium and Vacc inium  membranaceum, with a carpet of Rubus pedatus and Rhytidiopsis  robusta. 5.6.4. Ordination results from grouped transect data A series of ordinations was completed combining a l l 31 plots (16 Nanaimo, 15 Cameron). 1 6 Figure 45 shows the results of a DECORANA ordination of a l l samples. Samples have been l a b e l l e d according to vegetation series designations as determined using a f i e l d key (Packee 1979). A f a i r l y good separation was made between samples in the TSHE, TSHE-ABAM, and ABAM serie s . Samples 1 5Appendix VI, Table A39 gives vegetation c h a r a c t e r i s t i c s for samples in the ABAM-TSME vegetation zone that were not included in the c l a s s i f i c a t i o n . l sFour samples from the Cameron valley were taken from data c o l l e c t e d for MacMillan Bloedel Limited in 1978. 152 Figure 45. DECORANA ordi n a t i o n of samples i n the Cameron and Nanaimo transects - vegetation s e r i e s . • o • v A • A V • TSHE TSHE-ABAM ABAM-TSHE(THPL) ABAM-TSHE(CHNO) CHNO ABAM • • • • A 1979 p l o t s only axis 1 360 Figure 46. DECORANA o r d i n a t i o n of samples in the Cameron and Nanaimo transects (understory only) - habitat types. 320 9 * • O TSHE/GASH © TSHE/POMU A CHNO/GASH A ABAM/ACTR-TITR • ABAM/VAAL-VAPA B ABAM/VAAL(OV)/RUPE T ABAM/VAAL/STREPTOPUS ^ ABAM-TSME/RHAL * u n c l a s s i f i e d A O o o • • • o o o axis 1 450 153 of the ABAM-TSHE(THPL) and ABAM-TSHE(CHNO) series ordinated in the centre of the previous three series. A d i s t i n c t separation was made between the one TSHE-ABAM series plot in the Nanaimo valley and the TSHE-ABAM series plots in the Cameron v a l l e y . This plot borders the ABAM-TSHE(THPL) series in i t s reproduction and has understory differences that separate i t from the Cameron plo t s . Another separation occurred between plots in the TSHE series because of differences in the abundance of Gaultheria  shallon and the associated d i v e r s i t y of herbs. The f i r s t ordination axis in Figure 45 appears to r e f l e c t topographic moisture differences caused by elevation and aspect. Plots on dry, middle to lower southwest aspects were clustered on the l e f t , and high elevation NE aspects on the r i g h t . Samples on high elevation southwest aspects, valley bottoms and lower north slopes occupy an intermediate position. The r e l a t i o n of the second axis to any single environmental gradient i s not c l e a r . A DECORANA ordination of understory species alone i s presented in Figure 46. An octave data transformation was used with downweighting of rare species. The four 1978 plots were not included. The d i s t r i b u t i o n of habitat types has been plotted on the ordination diagram. The Gaultheria-dominated TSHE/GASH and CHNO/GASH habitat types are clustered together in two groups at the l e f t end of axis 1 and the lower end of axis 2. The high cover of s a l a l masks other species differences between the two types, es p e c i a l l y with overstory removed. One group includes three samples that occupy steep, southwest facing slopes with moderately high Gaulther ia cover and many herbs, of which 154 T r i e n t a l i s l a t i f o l i a , Festuca occidentalis, Lactuca muralis and trace amounts of Polyst ichum munitum are c h a r a c t e r i s t i c . In the second cluster of these two types are the TSHE/GASH samples from f l a t and northerly aspects and the highest elevation sample in the type. These plots have very high and vigorous Gaultheria cover to the exclusion of most herbs. Vaccinium parvifolium, absent in the f i r s t group, i s also common. Also included with th i s second group are three CHNO/GASH samples on upper elevation (900 - 1060 m) south aspects. These samples ordinated furthest to the right. The opposite extreme of axis 1 i s occupied by the two ABAM-TSME zone samples (Abies amabilis - Tsuga mertensiana -Rhododendron albiflorum association) and the c l o s e l y related ABAM/VAAL(OV)/RUPE habitat type. Axis 2 separates the Gaultheria-dominated samples from the TSHE/POMU and ABAM/ACTR-TITR samples. The grouping of the l a t t e r two habitat types shows their close s i m i l a r i t y in understory composition. The ABAM/VAAL-VAPA type ordinated in the centre of both axes, as one might expect from i t s intermediate moisture and temperature relationship to other p l o t s . The two u n c l a s s i f i e d samples with sparse understories are most similar to the TSHE/POMU and ABAM/ACTR-TITR habitat types. Axis 1 represents a gradient from d r i e r , warmer and/or more southerly exposures to cooler, wetter, northerly exposures ( l e f t to r i g h t ) . Figure 47 presents a species-by-sample matrix arranged in f i r s t axis order from which one can see the d i s t r i b u t i o n of species (<2 occurrences deleted) along this gradient. Axis 2 separates wetter and d r i e r habitats. A polar ordination using only understory species was 155 Figure 47. Species-by-sample'matrix in RA f i r s t axis order for understory species in the Cameron and Nanaimo transects. PLOT Not : 3554633—-53443443-655453554545 8746013 52739897-169086021123 2 SPECIES: FEOC -51-32 MOPA -5-2—1 CABU2 1-1 HIAL — 1 1 — 1 7 LAMU 4551411 31 ROGY — 1 1 ATFI 2 3 CASC -542-12 3—1 OPL02 4411-1 11 1 POGL4 —113-1 1 — 1 PYPI 1-23 1 1 NONE 1—2 1 BROMU 14 1 ARMA3 1 1 TRLA2 -131421—1 1 1-POMU 8-1-414343—43—7 CHME 1322443-11251422221—41-21-1 ACTR 5—2-1414—8-43—6 6 BENE 4—141—1 32118431-1 GASH -89-91 9-94-912—8748 . COMA 3 3—122 TROV 1 1—1-11—11 1 LIC03 —11 1-4—221-1 2 TITR 5 134—6-53-13-3 5-1-11-GOOB —11-12—11111-1111 11 CHUM -421511—15-542255165633 21--TILA l l l _ _ 6 _ 4 3 _ _ 2 - 3 1-1-1— LIB02 -3-2334—12-1-2212481-2182 VAPA 1—2-231341245255764664-75112— VIOR2 2-22 2-31-12 — 1 2-1—3-COME — 11 COCA 1 4-4—2-4 GAOV 1 14-2 1 LICA3 -4—1 4— 2 11-42 BLSP 1—6 4 1 STRO 1 1 1-LUZUL 1 1-VAME 2 2 25-852-2-52 LYCL 4 2-5-1-4-1 CLUN 1 4 2-HYMO 1-11 11-PYSE 1 1 11-12-1312-VAAL • 2—1625324428597 STAM 2-1—1-RUPE — — — — 1 — 1 3 — 1 - 6 1 2 5 1 7 2 VAOV : — i — _ — - 4 1 - 1 4 - 5 CLPY — 1 3 VEVI 2-111 SOSI 21 RHAL 11167 ^Blank plot numbers are for samples collected by MacMillan Bloedel i n 1978. 2 See Appendix III for f u l l species names. Cover values are scaled from the matrix maximum. 156 performed using computer-chosen endpoints which represent maximum compositional d i s s i m i l a r i t y (Figure 48). Axes are similar to the DECORANA ordination. The l e f t and right endpoints of axis 1 could be labelled warm and cool, respectively. The TSHE/POMU and ABAM-TSME/RHAL types occupy the warm and cool ends of this axis, respectively. The second axis endpoints could be termed wet and dry, with the ABAM/VAAL/STREPTOPUS and TSHE/GASH types occupying the respective extremes. This ordination shows a similar grouping of the two Gaultheria-dominated types as in DECORANA ordinations, with most of the CHNO/GASH plots ordinating closer to the cool end of axis 1. The TSHE/POMU and ABAM/ACTR-TITR types are again grouped together. The ABAM/VAAL-VAPA habitat type occupies a central position, but i s more spread out than in the DECORANA ordination. This is due to the fact that PO compares these samples to endpoints, spreading out intermediate samples unless they are unlike a l l endpoint samples. This ordination includes the four 1978 plots that include an ABAM/OPHO habitat type. This sample ordinated within the TSHE/POMU group. Species and sample ordinations of overstory alone are presented in Figures 49 and 50. The f i r s t axis DECORANA ordination revealed the follwing sequence of understory (A3) tree species from the warm-dry to the wet-cool portions of the interpreted gradient: Pseudotsuga menziesii, Pinus monticola, Thuja p i i c a t a , Tsuga heterophylla, Abies amabilis, Chamaecyparis  nootkatensis, Tsuga mertensiana. The sequence of overstory dominant and codominant (Al) and intermediate (A2) layers was s l i g h t l y d i f f e r e n t . A f i r s t axis RA arrangement of a species-by-Figure 48. Polar o r d i n a t i o n of samples i n the Cameron and Nanaimo transects (understory only) - habitat types. o Q O TSHE/GASH O O © TSHE/POMU A A CHNO/GASH O A ABAM/ACTR-TITR A • ABAM/VAAL-VAPA A B ABAM/VAAL(0V)/RUPE • T ABAM/VAAL/STREPTOPUS * A • ABAM-TSME/RHAL * . ABAM/OPHO * u n c l a s s i f i e d A , • O • O • m 1978 and • 1979 p l o t s axis 1 100 Figure 49. DECORANA or d i n a t i o n of samples i n the Cameron and Nanaimo transects using tree basal area - s e r i e s . 180 • TSHE ® « TSHE-ABAM • ABAM-TSHE(THPL) © A ^ ABAM-TSHE(CHNO) © v CHNO D ABAM • • • • • -axis 1 seo 158 Figure 50. DECORANA ordination of overstory species-'-in the Cameron and Nanaimo transects. 300 TSHE3 *PSME *ABAM , D „ n , *TSME3 ABAM3* ABAM2* *TSME2 *CHN03 *PIM03 *PSME2 *TSHE2 PSME3* *THPL2 *PIC0 *THPL3 *TSHE *CHNO *PIMO *THPL *TSME " axis 1 480 See accoimoanying sample ordination in Figure 49. Numbers in tree species abbreviations i d e n t i f y the canopy layer (unmarked = A l ; 2 = A2; 3 = A3). 159 sample matrix shows the d i s t r i b u t i o n of tree species in th i s set of samples (Figure 51). The sequence of the understory tree layer follows the moisture and temperature preferences of these species as reported in the l i t e r a t u r e (Minore 1979). Comparison was made of ordinations of transect data using basal area data versus estimated percent canopy cover data for tree species. Results show no s i g n i f i c a n t advantage in using measured basal area over cover estimates. The same general trends were observed with both, though s l i g h t differences not s i g n i f i c a n t enough to affe c t the interpretation did re s u l t . 5.6.5. Interpretation of environmental gradients Environmental data for each sample were plotted on ordination scattergrams to analyze patterns. Results are summarized in Figures 52a and 52b for the understory polar ordination. The trend in habitat types along axis 1 suggests a temperature gradient. Extrapolated temperature data support t h i s interpretation. Axis 1 was divided in half along a "natural" break in plot groupings. Average growing-season temperature for plots on the l e f t half i s 11.3° C; average temperature for the right half is 10.6° C. Temperature ranges from 9.3 to 12.4° C for a l l samples. The second axis i s interpreted as a moisture gradient. Calculated WSI and potential solar radiation values support t h i s interpretation. The axis was divided in half along a "natural" break. Eight of thirteen samples (62%) in the upper "dry" half showed some predicted water stress. Only 3 of 14 samples- (21%) 160 Figure 51. Species-by-sample matrix i n RA f i r s t axis order f o r tree species i n the Cameron and Nanaimo transects. PLOT NO. : 4553455-365655443353-34—445345 2322119-608160099773-88—375164 SPECIES: TSME2 675-5 ABAM 66-55 TSME3 75-51 3 1 ABAM2 6645 7 4 TSME 888—54 4 3 CHN02 5 5 — 5 - 2 — 3 — 2 CHNO 675—55—5 5 ABAM3 867675-77 — 5165111—4 CHN03 144-214—45-744 TSHE 778-99877-86898899-5-6766866 TSHE 3 -6575—575567265555—77-6758561 TSHE2 —6-66746-6461666658-1677-6715-THPL3 —1-52 445644211-6-16-517672-PSME 86-7-78777667779989888889889 THPL 6—45-6666-6-5-667—655 PIM03 4 4--' 4 THPL2 545—6-55-6—4667576-PSME2 721-5 6 6 — 5 — 5 5 1 7 6 PSME3 1 4 5 5-5142 "Blank p l o t numbers are for samples c o l l e c t e d by MacMillan Bloedel i n 1978. 'See Appendix I II for f u l l species names. Numbers i n species abbreviations indicate canopy layers (2 = A2; 3 = A3). Cover values are scaled from the matrix maximum. 161 Figure 52. Environmental variables plotted on a polar ordination for the Cameron and Nanaimo transects. a. WSI and potential solar radiation. O ave. 181M ave. 138M A O O A O A A • • o • WSI s o l i d = some water stress open = no water stress Radiation 0 > 190 M cal/cm 2/yr •170-190 A < 170 b. Average growing season temperature. Temperature ^ < 11.0 • ^ • 1 • > 11.0 •w 1 V (range = 9.3 V 1 •v ave.=11.3 | ave. = 10.6 162 in the lower "wet" half showed any stress. WSI values averaged 2 months of moderate stress for plots where stress was predicted. Potential annual solar radiation averaged 180.7 M cal/cm 2/year for the upper half of axis 2 and 137.5 M cal/cm 2/year for the lower h a l f . Solar radiation for a l l samples ranged from 88 to 200 M cal/cm 2/year. Because temperature and p r e c i p i t a t i o n data r e l i e d upon extrapolation with elevation, p r e c i p i t a t i o n and elevation also showed a trend associated with the f i r s t ordination axis from lower elevation, warmer, drier plots ( l e f t ) to higher elevation, cooler, wetter plots ( r i g h t ) . AWSC showed no relationship to either axis. Although the two axes were evaluated in terms of two separate properties, they are both related to temperature and moisture gradients and, as ' such, cannot be considered independent. The f i r s t axis most strongly r e f l e c t s elevational differences between these properties; the second axis most strongly r e f l e c t s topographic and location differences. Vegetation in these two valleys appears to be influenced most by temperature and moisture regime. Further investigation of n u t r i t i o n a l and other ecological factors would contribute to an even better understanding of vegetation patterns in these two locations. 163 CHAPTER 6 DISCUSSION 6.1. Comparison of habitat types to plant communities in previous c l a s s i f i c a t i o n s The habitat types described in th i s study were broadly comparable to the plant communities described in existing c l a s s i f i c a t i o n s in B. C. and the P a c i f i c Northwestern U. S. A summary of the comparison with other c l a s s i f i c a t i o n s is presented in Table VIII. This is not intended as an equivalency table, but should be interpreted as showing broad ecological s i m i l a r i t i e s . For instance, some of the Oregon and Washington types are d i f f i c u l t to c a l l "equivalent" because of several differences in species composition. A few of the types i d e n t i f i e d in this study did not appear in other c l a s s i f i c a t i o n s , but they were of limited areal extent and may have been included in broader types of other authors. Some of the habitat types described in t h i s study had considerable v a r i a b i l i t y in their species composition, p a r t i c u l a r l y some of the types in the ABAM-TSHE zone. This suggests that subdivisions might have been indicated had further sampling been undertaken. The habitat type c l a s s i f i c a t i o n presented here i s by no means a complete c l a s s i f i c a t i o n of a l l of the plant communities found on eastern Vancouver Island, as shown by the u n c l a s s i f i e d samples. Comparison with other c l a s s i f i c a t i o n s , however, suggests that a l l of the major ecosystem types recognized in previous c l a s s i f i c a t i o n s were sampled. This study revealed some s i g n i f i c a n t differences between T a b l e V I I I . R e l a t i o n s h i p of h a b i t a t t y p e s t o o t h e r p l a n t community c l a s s i f i c a t i o n s i n B.C. and the P a c i f i c N o r t h w e s t e r n U.S. P S M E and PSME-TSHE zones K r a j i n a ( 1 9 6 9 ) * K r a j i n a S S p i l s b u r y ( 1 9 5 3 ) ;McMinn ( 1 9 S 7 ) ; Muel ler-Dombois (1959) O r l o c i (1961) Kojima (1971) F r a n k l i n and F r a n k l i n e t . a l . Dyrness ( 1 9 7 3 ) ( 1 9 7 9 ) 1. P S M E / A R M E / G A S K 2. P S M E / G A S H / B : N E 3 . P S M E / H O D I / P O M U 4. A B G P . / P O M U 5 . T H P L / L Y A M TS HE zone #19 #S # 1, l a and 2 a 11 # 1 6 S a l a l - l i c h e n D o u g l a s - f i r - s a l a l D o u g l a s - f i r - w. r e d ce d a r - s w o r d f e r n W. r e d cedar - skunk-cabbage N/A N/A D - f i r - A r b u t u s -s a l a l - F e s t u c a P S M E / G A S H PSME/HODI/GASH 1 PSME/HODI 5 N/A 6. TSHE/GASH 7. "SHE/GASH/BENE/ ACTR 3 . TSHE/POMU #27 #2b and 24 N/A O r t h i c G a u l - G a u l t h e r i a assoc.TSHE/ACCI/GASH 2 TSHE/GASH t h e r i a f . t . TSHE/RHMA/GASH 5 TSHE/ACTR P o l y s t i c h u m f . t . A c h l y s t r i p h y l 1 a -P o l y s t i c h u m a s s o c . TSHE/POMU 2'' 3 , / S TSHE/POMU ABAM-TSHE zone 9. CHNO/GASH 10. ABAM/VAAL/VAPA 11. ASAM/ACTP./TITR 1 2 . ABAM/VAAL/STREPTOPUS 13. ABAM/VAAL(OV)/RUPE 1 4 . ABAM/OPHO #34 # 3 2 #4S #31 N/A ( V a c c i r , ium-G a u l t h e r i a f . t . ) (Vaccinium-moss f7E75 ( O r t h i c Blechnum f . t . ) Oplopanax -Adi a n turn f . t . (ABAM/GASH) 1 V a c c i n i u m ABAM/VAAL, BENE a l a s k a e n s e a s s o c . p h a s e ABAM/VAAL/COCA 5 (ABAM/ACTR) 5 ( p a r t o f V. a l a s k a e n s e a s s o c ) ABAM/STRO* (ABAM/TIUN) 5 ABAM/MEFE 1 Op 1opanax -Adiantum a s s o c . ABAM/OPHO (ABAM/GA5H) ABAM/VAAL, BENE phase (ABAM/ACTR) (ABAM/TIUN) ABAM/MEFE (ABAM/VAAL,RUPE phase) (ABAM/RHAL) ABAM/OPHO ( ) » sor.e r e l a t i o n s h i p e v i d e n t f . t . - f o r e s t t y p e N/A = not a p p l i c a b l e * See text for descriptions I F r a n k l i n 1 9 6 6 ,/ C o r l i s s and Dyrness 1 9 6 5 ./ M e u r i s s e and Youngberg 1 9 7 1 J B a i l e y 1 9 6 6 / D yrness e_t.al_. 1 9 7 4 165 plant communities on Vancouver Island and mainland B. C , as shown in a comparison with the types described by Orloci (1961) for the Coastal Western Hemlock (CWH) biogeoclimatic zone. Two major differences are: 1. The rare occurrence of Acer circinatum on Vancouver Island, which i s common on the mainland (and in coastal Oregon and Washington). 2. Plagiothecium undulatum, used as a c h a r a c t e r i s t i c moss for modal ecosystems in the CWH zone on the mainland, was less abundant in samples than i s evident in Orloci's work. Rhyt idiadelphus loreus and Hylocomium splendens appear to be more common and ch a r a c t e r i s t i c on Vancouver Island. This study also revealed the need for a complete description of a l l the plant communities on eastern Vancouver Island in one volume. At this time, there i s no single document containing such a description. Community c l a s s i f i c a t i o n s are scattered in several University of B r i t i s h Columbia theses that are unavailable to many people. Published c l a s s i f i c a t i o n work for the central eastern portion of the island i s limited. Publications by the B. C. Forest Service for the study area treat the higher l e v e l s of biogeoclimatic c l a s s i f i c a t i o n : zones, subzones and variants (Klinka e_t a_l. 1979). Only zonal ecosystems are d e s c r i b e d . 1 7 Klinka's (1977) species selection 1 7The f i r s t detailed description of ecosystems by the BCFS other than zonal ecosystems for Vancouver Island i s given in the Koprino River Watershed Study on the northwestern coast (Klinka et a l . 1980b). 166 guide includes interpretations of the ecosystem c l a s s i f i c a t i o n in t h i s area, but i t i s not intended as a f u l l description of the ecosystems of the region. Again, only zonal ("mesic") ecosystems are provided with written descriptions. In addition, i t i s based primarily on work from the mainland and needs some modifications for Vancouver Island. Ecosystems in the Douglas-f i r biogeoclimatic zone (PSME and PSME-TSHE vegetation zones) have been described in publications by several authors (Krajina and Spilsbury 1953, McMinn 1959, Mueller-Dombois 1965) based on work concentrated in a single r i v e r valley (Nanaimo). The Coastal Western Hemlock biogeoclimatic zone (TSHE and ABAM-TSHE vegetation zones) has been treated in a single study by Kojima (1971) in Strathcona P r o v i n c i a l Park. Kojima's communities are unique because of the high base status of the volcanic parent material in his study area and, consequently, his c l a s s i f i c a t i o n is not t y p i c a l of the central eastern portion of the island. One can see, therefore, that the area covered by this thesis f i l l s a gap in the existing descriptions of ecosystems. Unfortunately, t h i s thesis was unable to treat ecosystems in the Mountain Hemlock (ABAM-TSME) zone, which has had no published study for Vancouver Island. A future publication treating a l l of the ecosystems of eastern Vancouver Island i s a much needed item. The application of the habitat type system in t h i s thesis was d i f f e r e n t from previous approaches in two ways: 1) both vegetation zones and series were used, and 2) habitat types were named using more than a single overstory and understory dominant. The reason for the f i r s t difference i s that the upper level s of the c l a s s i f i c a t i o n were accepted as given by Packee 167 (1979). Use of both the zone and series concepts presents some problems in hierarchy, as discussed in Chapter 1, but has the advantage of recognizing both a broad area of similar macroclimate (zone) and the more detailed mosaic of landscape units with the same edaphic and/or climatic climax tree species ( s e r i e s ) . The l a t t e r i s a useful aggregation of habitat types with similar tree species potential for forestry applications. The reason for the second difference is that the plant associations encountered on the P a c i f i c coast are not adequately described by a single dominant tree and understory species, as is possible in the drier mountainous regions further inland where the c l a s s i f i c a t i o n system originated. This difference does not affect the fundamental basis of the c l a s s i f i c a t i o n , but recognizes the greater v a r i a b i l i t y and mixture of dominant species in the coastal environment. Some comment should also be made with regard to the key to the vegetation series presented in Appendix I. This key i s tentative and includes some series not encountered in the course of the f i e l d work. Some of these series appear questionable within some vegetation zones. For example, a PSME or ABGR series within the wet, west coast TSHE-ABAM zone seems unl i k e l y or rare. Some problems occur in the use of the key in areas where dense shade or heavy l i t t e r accumulations exclude a l l regeneration, and where Abies amabilis is temporarily excluded from an area because of f i r e history. These and other d i f f i c u l t i e s must be resolved by the subjective judgement of the user. Questions concerning the usefulness of some of the series for i d e n t i f y i n g s i g n i f i c a n t l y d i f f e r e n t environments arrived 168 from the ordinations conducted in thi s study. 6.2. Methodology The results obtained using DECORANA were far superior to those of basic reciprocal averaging. A l l indications from t h i s study substantiate claims that DECORANA is perhaps the best indirect ordination technique available. In a l l cases, i t produced a better clustering of samples within habitat types and a better separation between types than polar ordinations using the same data input and transformations. The methodology of DECORANA i s generally more advantageous than polar ordination (PO) because a l l samples are compared to each other rather than to selected endpoints. Results obtained with PO are t o t a l l y dependent upon the choice of four benchmark samples. This requires that the user have some knowledge of the data set prior to analysis, and that a good choice of endpoints be made. When these conditions are met, PO can give very good results with the added advantage of being easy to inte r p r e t . In addition, PO is not subject to the d i s t o r t i o n of results from deviant samples, unless chosen as endpoints, that a f f e c t s DECORANA. Polar ordination is not suited to ordinating data from several vegetation zones because of the high beta (between sample) d i v e r s i t y in such data. Samples in the middle of the data set w i l l have few species in common with either endpoint and form a cluster in the centre of the axes. DECORANA, on the other hand, works well for data with f a i r l y high beta d i v e r s i t y . In recommending use of these two techniques, DECORANA i s 169 suggested for the f i r s t ordination of data, followed by PO once gradient relationships are more evident. The use of both methods is preferable to either technique alone. By combining two or more ordination methods, the user i s assured that interpretation is based upon e c o l o g i c a l l y - s i g n i f i c a n t differences between samples and not upon exaggerated differences caused by the idiosyncrasies of a single method. It also protects against over-interpretation of a single ordination. The form of the data input can have as much or more of an e f f e c t upon ordination results than the choice of methodology. The elimination of o u t l i e r samples is important when using DECORANA because their inclusion may lead' to distorted results ( H i l l and Gauch 1980). Deletion of rare species also improves ordinations. The downweighting option in DECORANA is equally e f f e c t i v e as editing, yet allows these species to have some effe c t upon the r e s u l t s . For ordination of forest association data, the use of understory species alone gives superior results to combining understory and overstory species in most instances. Tree canopy is a less e f f e c t i v e indicator of s i t e potential than understory because of h i s t o r i c a l factors a f f e c t i n g forest cover. Habitat types, because of the way they are defined, are more e a s i l y i d e n t i f i e d on ordination diagrams of understory species; whereas, vegetation series are based on reproduction of tree species. An error was made in not analyzing the tree seedling data in ordinations of series. This omission was discovered too late to be incorporated into the computer-coded data; consequently, series correlations with axes are based 170 s t r i c t l y on c h a r a c t e r i s t i c s of their understory and overstory composition. When using percent cover data for herbs and shrubs, more favorable results are obtained when data are transformed to a cover scale. This reduces the overpowering ef f e c t of species with high dominance. Testing the differences between actual cover and scaled data is useful in analyzing the variation in dominance of certain species. This i s p a r t i c u l a r l y applicable in the case of the Polystichum- and Gaultheria-dominated habitat types. It is advisable to record, the best estimate of actual percentages in the f i e l d . Data can then be transformed to any cover scale desired without being r e s t r i c t e d by i n i t i a l choice. The author agrees with P f i s t e r et a l . (1977) that, with minimal tr a i n i n g , most people are able to estimate cover with s u f f i c i e n t accuracy. Comparison of the author's estimates on a few test plots with cover obtained from 20 1-m2 microplots taken on the same 500-m2 area have substantiated t h i s . The choice of plot location has a greater effect upon cover than the technique for obtaining cover. As in choice of ordination methods, i t is best not to rely on a single use of data transformations and editings. Much of the interpretive value of ordination l i e s in the a b i l i t y to test the effects of data manipulations and alternative techniques. 6.3. Environmental gradients The most r e l i a b l e interpretations of environmental gradients associated with DECORANA results were obtained for the 171 f i r s t ordination axes. As cautioned by Gauch et a l . (1977), "second and higher axes may be ec o l o g i c a l l y - meaningless, c u r v i l i n e a r functions of lower axes" with RA and similar techniques. This makes interpretation d i f f i c u l t for a l l but a single major gradient. Another factor that complicates interpretation from the outset i s that variation in species composition i s associated with many interrelated factors which cannot be considered independently; consequently, i t i s d i f f i c u l t to evaluate two factors on two perpendicular axes and assume they are independent. In this study, a reasonably clear separation between moisture and temperature gradients was obtained in the ordination of understory data from the ABAM-TSHE zone. In ordination diagrams,, inferred gradients are sometimes expressed obliquely to the two axes when interaction between axes occurs. Because of t h i s , an attempt to test for s t a t i s t i c a l c o r r e l a t i o n between ordination scores and quantitative environmental data was abandoned. The climatic and s o i l s data used in thi s study to evaluate environmental gradients have certa i n l i m i t a t i o n s . A l l climatic data were extrapolated from the nearest .short- or long-term cl i m a t o l o g i c a l station. One of the reasons for using such information was to see just how useful i t can be for s i t e comparisons. Calculated temperature and p r e c i p i t a t i o n figures are based on equations that rely on assumed lapse rates. While the equations are based on clim a t i c transect data for Vancouver Island, i t should be stressed that they are only pr o v i s i o n a l , and are intended only for general comparisons. In addition, 172 c l i m a t i c data that could be extrapolated are averages; the extremes of these factors may be e c o l o g i c a l l y more important. Extrapolated information cannot take into account such phenomena as temperature inversions, l o c a l i z e d rainshadows and cloud patterns, that complicate the generalized temperature/elevation and prec i p i t a t i o n / e l e v a t i o n relationships. For instance, cloud patterns were found to 'have a s i g n i f i c a n t e f f e c t upon tent c a t e r p i l l a r d i s t r i b u t i o n on the Saanich Peninsula of Vancouver Island (Wellington 1965). In the present study, the cold a i r drainage frequently associated with valley bottoms and deep ravines made extrapolated temperature data of l i t t l e value for such s i t e s . Few of the ordinations of individual zones showed any r e l a t i o n s h i p to plotted temperature or p r e c i p i t a t i o n data. Because of the complex elevational patterns of vegetation caused by the interaction of topography and climate, discussed in a recent paper by Daubenmire (1980), ind i r e c t ordinations of the vegetation give better indications of temperature gradients than extrapolated data or direct comparisons with elevation. The extrapolated data were, however, s u f f i c i e n t to evaluate o v e r a l l v a r i a t i o n between zones. S o i l temperature is an important factor that was not evaluated in t h i s study. The s o i l temperature environment may be d i f f e r e n t from that indicated by a i r temperature. For example, finer-textured s o i l s w i l l be colder in the spring than coarser s o i l s because of their higher water retention capacity, which makes them slower to heat up (Bannister 1976). Consequently, two s i t e s with i d e n t i c a l a i r temperature may have very d i f f e r e n t s o i l temperature environments. 173 Potential solar radiation provided a general comparison between s i t e s , but could not account for such factors as shading from adjacent mountains and cloudiness. Aspect may also determine the influence of other climatic factors. For example, pr e v a i l i n g winds can influence evapotranspiratioh rates and p r e c i p i t a t i o n d i s t r i b u t i o n on d i f f e r e n t facing slopes. Attempts to measure and map radiation environments and correlate them with vegetation are complicated by the i n t e r r e l a t i o n s h i p s with other factors, p a r t i c u l a r l y s o i l moisture (Granger and Schulze 1977). Despite these complicating factors, potential radiation proved to be a better measurement for comparison of s i t e s than aspect alone because slope and latitude were also taken into account. A moisture gradient had the most pronounced effect upon species d i s t r i b u t i o n in ordinations of a l l zones. This finding i s consistent with a study of similar communities near Vancouver. Eis (1962) found that s o i l moisture, groundwater and s o i l permeability combined accounted for 79% of the v a r i a b i l i t y in plant communities from a s t a t i s t i c a l analysis of numerous s i t e variables. Kojima (1971) reached similar conclusions for ecosystems in Strathcona Park on Vancouver Island. The Water Stress Index (WSI) showed a better correlation with axes interpreted as complex moisture gradients than Available Water Storage Capacity (AWSC) alone. The effectiveness of the WSI was limited by the accuracy of the climatic data. It was also limited by the AWSC calculations, which were based on estimates of s o i l texture, coarse fragment content and rooting depth. There i s a certain error factor in these estimates, added 174 to the fact that the s o i l may be quite variable within the 500-m2 plot area that the single s o i l p i t is intended to represent. Add to t h i s the inherent i n a b i l i t y of the index to account for seepage and high water table and i t is surprising that i t provided any useful information. Whether or not the WSI gives accurate predictions as to the actual amount of moisture stress occurring on a given s i t e is questionable. It does, however, serve as a useful tool for evaluating the relat ive moisture differences between s i t e s . WSI values indicated that, despite general differences between types, there can be s i g n i f i c a n t v a r i a b l i l i t y within a type. WSI predictions showed that moderate and severe moisture stress i s common in the PSME and PSME-TSHE zones; stress in the TSHE zone is moderate and less common; and stress in the ABAM-TSHE i s infrequent. The physiological response of plants to moisture d e f i c i t s and other parameters l i m i t i n g plant growth i s the underlying factor to be considered when evaluating a species or community relationship to environment. Generalized values such as "permanent w i l t i n g point" may be used in estimating moisture stress, as in the WSI; however, the actual stress encountered by a plant i s dependent upon the physiological mechanisms i t has developed for dealing with such d e f i c i t s . Consequently, the same levels of moisture in the s o i l w i l l affect d i f f e r e n t tree or understory species to d i f f e r e n t degrees. There may also be moisture stress associated with excess water that leads to similar e f f e c t s as drought. Measurement of environmental gradients in terms of plant 175 physiological response may be the best evaluation method. Based on work by Waring and Cleary (1967) and Cleary and Waring (1969), several later studies (Waring e_t a l . 1972, Zobel et a l . 1976) have used measured plant moisture stress by the pressure bomb technique (Scholander et a l . 1965) and a Temperature Growth Index (TGI), based on dry weight increases in Douglas-fir seedlings linked to s o i l and a i r temperatures, to examine vegetation d i s t r i b u t i o n on a two-dimensional g r i d . Waring et a l . (1972) used these data to develop indices for predicting environment from species composition. Additional studies of thi s nature are necessary to establish the indicator value of certain plants in predicting s i t e c h a r a c t e r i s t i c s . More integration of phytosociological studies and physiological studies has been suggested (Bannister 1976) . The generalized data available for the present study are not s u f f i c i e n t l y precise to make an accurate q u a n t i f i c a t i o n of environment associated with habitat types or individual species, though some general relationships could be evaluated. Nutrient differences between s i t e s were not evaluated in this study because laboratory analysis of s o i l samples has not been completed. This additional factor could help to explain some of the results not interpretable in terms of moisture, temperature or solar radiation gradients. There are many other factors that complicate the prediction of s i t e c h a r a c t e r i s t i c s from plant indicators. The role of competion and chance cannot be overlooked. Sampling technique is another important factor introducing "noise" into a data set. It is often tempting to try to explain the occurrence of every 176 plant on a s i t e by seeking some causal environmental factor. This temptation i s even greater, and more dangerous, when carri e d into the mathematical analysis of data. It i s , therefore, important to keep one's perspective in interpreting ordinations and other s t a t i s t i c a l techniques to allow for a certain amount of unexplained v a r i a t i o n . 6.4. Ordination as an aid in c l a s s i f i c a t i o n Ordination proved to be a valuable aid in the c l a s s i f i c a t i o n of habitat types for thi s study. This is not a new discovery by any means; ordination has been used for this purpose in studies too numerous to mention. A l l of the recent work in habitat type c l a s s i f i c a t i o n in the P a c i f i c Northwestern U. S. has u t i l i z e d some ordination techniques, usually polar ordination ( P f i s t e r and Arno 1980, Franklin et a l . 1979). It has also been used in computer modeling of forest ecosystem attr i b u t e s for f i r e management predictions (Kessell 1979). Ordination, however, has not been widely used in B r i t i s h Columbia to date. The biogeoclimatic c l a s s i f i c a t i o n e f f o r t s in B. C. have r e l i e d largely upon t r a d i t i o n a l Braun-Blanquet tabular analysis for the segregation of taxonomic units. There are several ways in which ordination can help improve c l a s s i f i c a t i o n and provide additional information once a c l a s s i f i c a t i o n system has been developed. Ordination diagrams provide a v i s u a l way of evaluating the degree of homogeneity within types. There i s often too much emphasis placed on the average c h a r a c t e r i s t i c s of a vegetation type, rather than the v a r i a b i l i t y within a type. Ordination can help in the evaluation 177 of t h i s v a r i a b i l i t y . I t can a l s o a i d i n i d e n t i f y i n g p o s s i b l y m i s - c l a s s i f i e d samples. The degree of d i f f e r e n c e or s i m i l a r i t y between c l a s s i f i c a t i o n u n i t s can be evaluated v i s u a l l y and may lea d to f u r t h e r segregation or r e d e f i n i t i o n of types when s i g n i f i c a n t v e g e t a t i v e or environmental a t t r i b u t e s are i n v o l v e d . F i n a l l y , o r d i n a t i o n can be a u s e f u l t o o l i n the a n a l y s i s of environmental gra d i e n t r e l a t i o n s h i p s among c l a s s i f i c a t i o n u n i t s . Perhaps the most important c o n s i d e r a t i o n f o r s u c c e s s f u l use of o r d i n a t i o n i s to avoid o v e r - i n t e r p r e t a t i o n . C a r e f u l a n a l y s i s of r e s u l t s and a thorough f a m i l i a r i t y with the methodology are e s s e n t i a l . A s s o c i a t i o n t a b l e s are a necessary companion i n using o r d i n a t i o n f o r c l a s s i f i c a t i o n . They are needed to c h a r a c t e r i z e types and he l p to i n t e r p r e t the p o s i t i o n i n g of a given sample with r e s p e c t to others i n the same or d i f f e r e n t type. The B. C. Forest S e r v i c e t a b l e p r e p a r a t i o n program i s h e l p f u l f o r t h i s purpose. O r d i n a t i o n does not a c t u a l l y perform c l a s s i f i c a t i o n - - i t only a i d s i n t h i s p r o c e s s . The judgment amd s u b j e c t i v e i n t e r p r e t a t i o n of the user i s the f i n a l word i n c l a s s i f i c a t i o n . As in f o r e s t r y i t s e l f , c l a s s i f i c a t i o n i s an a r t as w e l l as a s c i e n c e . Numerical techniques can save time and r e v e a l obscure r e l a t i o n s h i p s , but there i s s t i l l no s u b s t i t u t e f o r f i e l d experience and i n t e g r a t i n g a b i l i t y of the human mind. 178 CONCLUSIONS Both DECORANA and polar ordination are e f f e c t i v e tools for the c l a s s i f i c a t i o n and gradient analysis of forest habitat types. DECORANA produces superior results to reciprocal averaging and polar ordination in the majority of cases. Ordinations of understory percent cover data transformed to an octave scale produce the most favorable clustering of samples representing habitat types. Ordinations of tree species alone or tree and understory species combined are less successful. Vegetation series are not readily segregated using ordination of tree and understory species. Ordination of vegetation zones i s most successful using DECORANA and produces a continuous d i s t r i b u t i o n of samples along ordination axes with no sharp separation between zones. Environmental gradients can be related to the vegetation variation i d e n t i f i e d by ordination axes. A moisture gradient accounts for most of the variation between habitat types within any single vegetation zone. A Water Stress Index is a useful integrator of s i t e moisture factors for general comparisons, but has several l i m i t a t i o n s due to extrapolated climatic data input and i t s i n a b i l i t y to account for seepage water and microclimate. Potential solar radiation and temperature gradients also influence the d i s t r i b u t i o n of habitat types. The combination of ordination techniques and t r a d i t i o n a l association table synthesis i s a useful procedure for the c l a s s i f i c a t i o n of forest communities in coastal B r i t i s h Columbia. 179 LITERATURE CITED Armstrong, J. E., D. R. Crandell, D. J. Easterbrook and J . B. Noble. 1965. Late Pleistocene stratigraphy and chronology in southwestern B r i t i s h Columbia and northwestern Washington. Geol. Soc. Amer. B u l l . 76: 321-330. Arno, S. F. 1979. Forest regions of Montana. USDA For. Serv., Res. Pap. INT-128, 39p. Intermountain For. and Range Exp. Stn., Ogden, Utah. Atmospheric Environment Service. 1973. Canadian Normals. Volume 1. Temperature. 1941-1970. Environ. Can. Atmos. Environ. Serv., 170p. Bailey, A. W. 1966. Forest associations and secondary plant succession in the southern Oregon Coast Range. Ph.D. thesis, Oregon State Univ. 164p. Bailey, A. W., and W. W. Hines. 1971. A vegetation-soil survey of a w i l d l i f e - f o r e s t r y research area and i t s application to management in northwest Oregon. Oregon State Game Comm. Res. Div. Game Rept. No. 2, 36p. Bailey, A. W.,• and C. E. Poulton. 1968. Plant communities and environmental in t e r r e l a t i o n s h i p s in a portion of the Tillamook Burn, northwestern Oregon. Ecology 49: 1-13. Bailey, R. G., R. D. P f i s t e r , and J . A. Henderson. 1978. Nature of land and resource c l a s s i f i c a t i o n - a review. J. Forestry 76(10): 650-655. Ballard, T. M. 1974. A water stress index for f i e l d use in south coastal B r i t i s h Columbia. A section of the Fi n a l Report to the Pa c i f i c Forest Research Centre, CFS, Res. Contract OSP3-0397 "Evaluation of Forest Site Moisture Regimes", 48p. Bannister, P. 1976. Introduction to Physiological Plant Ecology, John Wiley and.Sons, New York, 273p. Barrett, J. W. (ed.) 1962. Regional S i l v i c u l t u r e of the United States. Ronald Press Co., New York, 610p. B e i l , C. E., R. L. Taylor, and G. A. Guppy. 1976. The biogeoclimatic zones of B r i t i s h Columbia. Davidsonia 7: 45-55. Benzecri, J. P. 1969. S t a t i s t i c a l analysis as a tool to make patterns emerge from data. Methodologies of Pattern Recognition (Ed. By S. Watanabe), Academic Press, New York pp. 35-60. Bernier, B. 1968. Descriptive outline of forest humus-form c l a s s i f i c a t i o n . Proc. 7th Meeting of the Nat. S o i l Surv. Comm. Of Canada, Univ. of Alberta, Edmonton, pp. 139-154. 180 Black, T. A., K. G. McNaughton, and P. A. Tang. 1973. Studies of forest evapotranspiration and tree stem diameter growth. Final contract research report to the Canadian Forestry Service (DOE), March, 1973. 78p. Black, V. L. 1973. The estimation of net radiation to a Douglas-f i r forest using standard climatological data. B.S. thesis, S o i l Science, Univ. of B. C , Vancouver, 18p. Braun-Blanquet, J. 1932. Plant Sociology. McGraw-Hill Book Co. Inc., New York 439p. (Translated from the German by H. S. Conrad and G. D. F u l l e r . ) Bray, J. R. 1956. A study of the mutual occurrence of plant species. Ecology 37: 21-28. Bray, J. R. 1960. The composition of the savanna vegetation of Wisconsin. Ecology 41: 721-732. Bray, J. R. 1961. A test for estimating the r e l a t i v e informativeness of vegetation gradients. J. Ecology 49: 631-642. Bray, J. R., and J. T. Curtis. 1957. An ordination of the upland forest communities of southern Wisconsin. Ecol. Monogr. 27: 325-349. Brooke, R. C , E. B. Peterson, and V. J. Krajina. 1970. The subalpine Mountain Hemlock Zone. I_n V. J. Krajina (ed.), Ecology of western North America. Univ. of B. C. Dept. of Botany, Vol. 2(2): 153-349. Buffo, J. L., j . Fritschen, and J. L. Murphy. 1972. Direct solar radiation on various slopes from 0° to 60° north l a t i t u d e . USDA For. Serv., Res. Pap. PNW-142, 74p. P a c i f i c Northwest For. and Range Exp. Stn., Portland, Ore. Campbell, G. W. 1977. An Introduction to Environmental Biophysics. Springer-Verlag, New York. Canada S o i l Survey Committee, Subcommittee on S o i l C l a s s i f i c a t i o n . 1978. The Canadian system of s o i l c l a s s i f i c a t i o n . Can. Dept. Agric. Publ. 1646, 164p. Chapman, J. D. 1952. The Climate of B r i t i s h Columbia. A paper presented to the F i f t h B r i t i s h Columbia Natural Resources Conference, Univ. of B. C , Vancouver, February 27, 1952. 47p. Cleary, B. D., and R. H. Waring. 1969. Temperature: c o l l e c t i o n of data and i t s analysis for the interpretation of plant growth and d i s t r i b u t i o n . Can. J. Bot. 47: 167-173. Cordes, L. D. 1972. An ecological study in the Sitka spruce forest of the West Coast of Vancouver Island. Ph.D. thesis, Univ. of B. C , Vancouver, 452p. C o r l i s s , J . F., and C. T. Dyrness. 1965. A detailed s o i l -181 vegetation survey of the Alsea area in the Oregon Coast Range. In C. T. Youngberg (ed.), F o r e s t - s o i l relationships in North America, Oregon State Univ. Press, C o r v a l l i s , pp. 457-483. Curtis, J. T., and R. P. Mcintosh. 1951. An upland forest continuum in the p r a i r i e - f o r e s t border region of Wisconsin. Ecology 32: 476-496. Daubenmire, • R. 1952. Forest vegetation of northern Idaho and adjacent Washington, and i t s bearing on concepts of vegetation c l a s s i f i c a t i o n . Ecol. Monogr. 22: 301-330. Daubenmire, R. 1968. Plant Communities: a textbook of plant synecology. Harper and Row, New York 300p. Daubenmire, R. 1980. Mountain topography and vegetation patterns. Northwest S c i . 54(2): 146-152. Daubenmire, R., and J . B. Daubenmire. 1968. Forest vegetation of eastern Washington and northern Idaho. Wash. State Univ. Agr. Exp. Sta. Tech. B u l l . 60, 104p. Day, J. H., L. Farstad, and D. G. L a i r d . 1959. S o i l survey of southeast Vancouver Island and Gulf Islands, B r i t i s h Columbia. B. C. S o i l Survey, Report No. 6. Del Moral, R., and J. N. Long. 1977. C l a s s i f i c a t i o n of montane forest community types in the Cedar River drainage of western Washington, U. S. A. Can. J . For. Res. 7(2): 217-225. Dick-Peddie, W. A., and W. H. Moir. 1970. Vegetation of the Organ mountains, New Mexico. S c i . Ser. No. 4, Range S c i . Dept., Colorado State Univ., 28p. Dyrness, C. T., J. F. Franklin, and W. H. Moir. 1974. A preliminary c l a s s i f i c a t i o n of forest communities in the central portion of the western Cascades in Oregon. Coniferous Forest Biome B u l l . No. 4, Univ. of Washington, College of For. Res., Seattle, 123p. E i s , S. 1962. S t a t i s t i c a l analysis of several methods for estimatin of forest habitat and tree growth near Vancouver. Univ. of B. C. Faculty of Forestry B u l l . 4, 76p. Ellenburg, H. 1948. Unkrautgesellschaften als Mass fur den Sauregrad, die Verdichtung und andere Eigenschaften des Ackerbodens. Ber. Landtech. 4: 130-146. Fasham, M. J. R. 1977. A comparison of non-metric multidimensional scaling, p r i n c i p a l components and reciprocal averaging for the ordination of simulated coenoclines, and coenoplanes. Ecology 58: 551-561. Fonda, R. W., and L. C. B l i s s . 1969. Forest vegetation of the montane and subalpine zones, Olympic mountains, Washington. Ecol. Monogr. 39: 271-301. 182 Franklin, J . F. 1966. Vegetation and s o i l s in the subalpine forests of the southern Washington Cascade Range. Ph.D. thesis, Wash. State Univ., Pullman, 132p. Franklin, J. F., and C. T. Dyrness. 1973. Natural vegetation of Oregon and Washington, USDA For. Serv., Gen. Tech. Rept. PNW-8, 417p. P a c i f i c Northwest For. and Range Exp. Stn., Portland, Ore. Franklin, J. F., W. H. Moir, M. A. Hemstrom, and S. Greene. 1979. Forest ecosystems of Mount Rainier National Park. Review draft of a proposed PNW For. and Range Exp. Stn. Res. Pap., USDA For. Serv., 221p. de l a Fuente, C, 1977. Water stress in forest s o i l s : a FORTRAN program to compute WSI values. Bs.F. thesis, Fac. of Forestry, Univ. Of B. C , Vancouver, lOp. + Appendices. Furnival, T. G., E. Wyler, W. Reifsnyder, and T. G. Succaina. 1969. Computer program for computation of solar radiation at the top of the atmosphere and the day length. Unpublished program, Yale School of Forestry. Fyles, J. G. 1959. S u r f i c i a l geology. Oyster River, Comox, Nanaimo and Sayward D i s t r i c t s , B r i t i s h Columbia. Geol. Surv. of Canada, map 49-1959 (1:63000).. Fyles, J. G. 1960. S u r f i c i a l geology. Courtenay, Comox, Nelson, Nanaimo and Newcastle D i s t r i c t s , Vancouver Island, B r i t i s h Columbia. Geol. Surv. of Canada, map 32-1960 (1:63000). Fyles, J . G. 1963a. S u r f i c i a l geology of Home Lake and Par k s v i l l e map areas, Vancouver Island, B r i t i s h Columbia. Geol. Surv. of Canada, memoir 318, 142p. Fyles, J . G. 1963b. S u r f i c i a l geology. Nanaimo, B r i t i s h Columbia. Geol. Surv. of Canada, map 27-1963 (1:63000). Fyles, J . T. 1955. Geology of the Cowichan Lake area, Vancouver Island, B r i t i s h Columbia, B. C. Dept. Mines B u l l . 37, 73p. Garrison, G. A., J . M. Skovlin, C. E. Poulton, and A. H. Winward. 1976. Northwest plant names and symbols for ecosystem inventory and analysis. 4th Ed i t i o n . USDA For. Serv., Gen. Tech. Rept. PNW-46, 263p. P a c i f i c Northwest For. and Range Exp. Stn., Portland, Ore. Gauch, H. G. J r . 1977. ORDIFLEX — A f l e x i b l e computer program for four ordination techniques: weighted averages, polar ordination, p r i n c i p a l components analysis, and reciprocal averaging, Release B. Ecol. and Systematics, Cornell Univ., Ithaca, New York 185p. Gauch, H. G. J r . , and R. H. Whittaker. 1972. ordination techniques. Ecology 53: 868-875. Compar i son of 183 Gauch, H. G. J r . , R. H. Whittaker and T. R. Wentworth. 1977. A comparison of reciprocal averaging and other ordination techniques. J. Ecol. 65: 157-174. G i t t i n s , R. 1969. The application of ordination techniques. I_n I. H. Rorison (ed.) Ecological Aspects of the Mineral Nutrition of Plants. Symp. B r i t . Ecol. Soc. 1968, 9: 37-66. Goodall, D. W. 1954. Objective methods for the c l a s s i f i c a t i o n of vegetation. I l l An essay in the use of factor analysis. Aust. J. Bot. 2:304-324. Goodall, D. W. 1973. Sample s i m i l a r i t y and species c o r r e l a t i o n . I n R. H. Whittaker (ed.), Handbook of Vegetation Science 5: Ordination and c l a s s i f i c a t i o n of communities. Junk, The Hague, pp. 105-156. Granger, J. E., and R. E. Schulze. 1977. Incoming solar radiation patterns and vegetation response: examples from the Natal Drakensberg. Vegetatio 35(1): 47-54. G r i f f i n , J. R. 1967. S o i l moisture and vegetation patterns in northern C a l i f . Forests. USDA For. Serv., Res. Pap. PSW-46, 22p. P a c i f i c Southwest For. and Range Exp. Stn., Berkeley, C a l i f . G r i f f i t h , B. G. 1960. Growth of Douglas-fir at the University of B r i t i s h Columbia Research Forest as related to climate and s o i l . Univ. of B. C. Fac. of Forestry, For. B u l l . No. 2, 58p. Guinochet, M. 1973. Phytosociologie. Masson, Paris. Henderson, J. A., R. L. Mauk, D. L. Anderson, T. A. Davis, and T. J . Keck. 1977. Preliminary forest habitat types of the Uinta mountains, Utah. Dept of For. and Outdoor Recr., Utah State Univ., Logan. 94p. Heusser, C. J. 1960. Late-Pleistocene environments of North P a c i f i c North America. Amer. Geogr. Soc. Special Publ. No. 308, 372p. H i l l , M. 0. 1973. Reciprocal averaging: an eigenvector method of ordinaion. J . Ecol. 61: 237-249. H i l l , M. 0. 1979. DECORANA — A FORTRAN program for detrended correspondence analysis and reciprocal averaging. Ecol. and Systematics, Cornell Univ., Ithaca, New York 52p. H i l l , M. 0., and H. G. Gauch, J r . 1980. Detrended correspondence analysis, an improved ordination technique. Paper submitted to Vegetatio, 12p. Hines, W. W. 1971. Plant communities in the old-growth forests of north coastal Oregon. M.S. thesis, Oregon State Univ. 146p. Hitchcock, C. L., and A. Cronquist. 1973. Flora of the P a c i f i c Northwest. Univ. of Washington Press, Seattle, 730p. 184 Hoffman, G. R., and R. R. Alexander. 1976. Forest vegetation of the Bighorn mountains, Wyoming: a habitat type c l a s s i f i c a t i o n . USDA For. Serv., Res. Pap. RM-170, 38p. Rocky Mtn. For. and Range Exp. Stn., Ft. C o l l i n s , Colo. Holland, S. S. 1964. Landforms of B r i t i s h Columbia: a physiographic outline. B. C. Dept. Mines and Petr. Res. B u l l . 48, 138p. Jones, R. K. 1978. The numerical c l a s s i f i c a t i o n and mapping of vegetation in two mountainous watersheds of southeastern B r i t i s h Columbia. M.S. thesis, Fac. of Forestry, Univ. of B. C , Vancouver, 229p. Jones, R. K., and R. Annas. 1978. Vegetation. I_n K. W., G. Valentine et a l . (eds.), S o i l Landscapes of B r i t i s h Columbia, Resource Analysis Branch, Min. of Environ., V i c t o r i a , B. C , pp. 35-45. Keser, N., and D. St. Pierre. 1973. S o i l s of Vancouver Island: a compendium. B. C. Min. of Forests, Res. Div., Res. Note No. 56, unpaged. Kessell , S. R. 1979. Gradient Modeling: resource and f i r e management. Vol. 1. Springer Series on Environmental Management. Springer-Verlag, New York 433p. Kessell, S. R., and R. H. Whittaker. 1976. Comparisons of three ordination techniques. Vegetatio 32: 21-29. Kimmins, J..P. 1977. The need for ecological c l a s s i f i c a t i o n in B r i t i s h Columbia. B. C. Forestry, Assoc. Of B. C. Prof. Foresters, Vol. 2 No. 1, 2p. ( r e p r i n t ) . Klinka, K. 1976. Ecosystem units, their c l a s s i f i c a t i o n , interpretation, and mapping in the University of B r i t i s h Columbia Research Forest. Ph.D. thesis, Univ. of B. C , Vancouver, 622p. Klinka, K. 1977. Guide for the tree species selection and prescribed burning in the Vancouver Forest D i s t r i c t , 2nd Approx. B. C. Min. of For., Res. Div., 42p. + Appendices. Klinka, K., F. C. Nuszdorfer, and L. Skoda. 1979. Biogeoclimatic units of central and southern Vancouver Island. B. C. Min. of Forests, V i c t o r i a , B. C , 120p. + map. Klinka, K., and S. Phelps. 1979. Environment-vegetation tables by a computer program. Univ. of B. C. Fac. of Forestry, 24p. (unpublished). Klinka, K., R. N. Green, R. L. Trowbridge, and L. E. Lowe.1981a. Taxonomic c l a s s i f i c a t i o n of humus forms in ecosystems of B r i t i s h Columbia. F i r s t approximation. Province of B r i t i s h Columbia, Ministry of Forests. In Press. 185 Klinka, K., W. D. van der Horst, F. C. Nuszdorfer, and R. G. Harding. 1980b. An ecosystematic approach to a subunit plan Koprino River watershed study. B. C. Min. of Forests, V i c t o r i a , B. C., 118p. + maps. Kojima, S. 1971. Phytogeocoenosis of the Coastal Western Hemlock Zone in Strathcona Provincial Park, B. C , Canada. Ph.D. thesis, Dept. of Botany, Univ. of B. C , Vancouver, 321p. Krajina, V. J. 1959. Biogeoclimatic zones in B r i t i s h Columbia. Univ. of B. C. Botanical Series, No. 1, 47p. Krajina, V. J. 1960. Ecosystem c l a s s i f i c a t i o n of forests. S i l v a Fennica 105: 107-110, 123-138. Krajina, V. J. 1965. Biogeoclimatic zones and c l a s s i f i c a t i o n of B r i t i s h Columbia. Ecol. Western N. A. I: 1-17. Krajina, V. J. 1969. Ecology of forest trees in B r i t i s h Columbia. Ecol. Western N. A. 2(1): 1-146. Krajina, V. J. 1972. Ecosystem perspectives in forestry. The H. R. MacMillan Lectureship in Forestry, Univ. of B. C , Vancouver, 31p. Krajina, V. J., and H. Spilsbury. 1953. Forest associations on the east coast of Vancouver Island. I_n Forestry Handbook for B r i t i s h Columbia, 1st Edition., Univ. of B. C. Forestry Club, pp.142-145. Kuramoto, R. T. 1965. Plant associations and succession in the vegetation of the sand dunes of Long Beach, Vancouver Island. M.Sc. thesis, Dept. of Botany, Univ. of B. C , Vancouver, 87p. Lawton, E. 1971. Moss Flora of the P a c i f i c Northwest. The Hattori Botanical Laboratory, Nichinan, Miyazaki, Japan, 362p. + i l l u s . Layser, E. F., and G. H. Schubert. 1979. Preliminary c l a s s i f i c a t i o n for the coniferous forest and woodland series of Arizona and New Mexico. USDA For. Serv., Res. Pap. RM-208, 27p. Rocky Mtn. For. and Range Exp. Stn., Ft. C o l l i n s , Colo. Lesko, G. 1961. Ecological study of s o i l s in the Coastal Western Hemlock zone. M.Sc. thesis, Dept. of B i o l , and Botany, Univ. of B. C , Vancouver, 141p. Lewis, T. 1976. The t i l l - d e r i v e d Podzols of Vancouver Island. Ph.D. thesis, Dept. of S o i l Science, Univ. of B. C , Vancouver, 158p. L i s t , R. J. 1949. Smithsonian Meterological Tables, 6th e d i t i o n . Smithsonian I n s t i t u t i o n Press, Washington, D. C , 527p. (5th reprint - 1971). Maas, E. F. 1972. The organic s o i l s of Vancouver Island. Agric. Canada Res. Stn., Sidney, B. C. Contr. No. 231, 35p. 186 MacMillan Bloedel Limited. 1974. The biogeoclimatic zones of Vancouver Island and the adjacent mainland based on climax vegetation (3rd Approx.). MacMillan Bloedel Limited, Forestry D i v i s i o n , Nanaimo, B. C. (map). Mathews, W. H. 1947. Calcareous deposits of the Georgia S t r a i t area. B. C. Dept. of Mines and Petr. Res. B u l l . No. 23, 113p. McLean, A. 1970. Plant communities of the Similkameen Valley, B r i t i s h Columbia and their relationships to s o i l s . Ecol. Monogr. 20: 403-424. McMinn, R. G. 1957. Water relations in the Douglas-fir region on Vancouver Island. Ph.D. thesis, Dept. of B i o l , and Botany, Univ. of B. C , Vancouver. McMinn, R. G. 1959. Water relations in the Douglas-fir region on Vancouver Island. Can. Dept. Agric. Publ. No. 1091., 71p. McNaughton, K. G. 1974. A study of the energy balance of a Douglas-fir forest. Ph.D. thesis, Dept. of S o i l Science, Univ. of B. C , Vancouver, 97p. Meurisse, R. T., and C. T. Youngberg. 1971. Soil-vegetation survey and s i t e c l a s s i f i c a t i o n report for Tillamook and Munson F a l l s Tree Farms. Oregon State Univ. Dept. S o i l s , Report to Publishers Paper Co., 116p. Minore, D. 1979. Comparative autecological c h a r a c t e r i s t i c s of northwestern tree species - a l i t e r a t u r e review. USDA For. Serv., Gen. Tech. Rept. PNW-87, 72p. P a c i f i c Northwest For. and Range Exp. Stn., Portland, Ore. Mi t c h e l l , R., and W. H. Moir. 1976. Vegetation of the Abbott Creek Research Natural Area, Oregon. Northwest S c i . 50: 42-58. Moir, W. H., and J. A. Ludwig. 1979. A c l a s s i f i c a t i o n of spruce-fir and mixed conifer habitat types of Arizona and New Mexico. USDA For. Serv., Res. Pap. RM-207, 47p. Rocky Mtn. For. and Range Exp. Stn., Ft. C o l l i n s , Colo. Mueller-Dombois, D. 1959. The Douglas-fir associaions on Vancouver Island in their i n i t i a l stages of secondary succession. Ph.D. thesis, Dept. of B i o l , and Botany, Univ. of B. C , Vancouver, 570p. Mueller-Dombois, D. 1965. I n i t i a l stages of secondary succession in the Coastal Douglas-fir zone of B r i t i s h Columbia. Ecol. Western N. A. 1: 35-37. Mueller-Dombois, D., and H. Ellenberg. 1974. Aims and Methods of Vegetation Ecology. John Wiley and Sons, New York 547p. Muller, J. E. 1965. Geology. Comox Lake area. Geol. Surv. Can., Dept. of Mines and Tech. Surv., Map 2-1965 (1:126720). 187 Muller, J. E. 1971. Geological reconnaissance map of Vancouver Island and Gulf Islands. Geol. Surv. Can. Open F i l e , Map 62. Muller, J. E., and J. G. T. Carson. 1969. Geology and mineral deposits of Alberni map-area, B. C , Geol. Surv. Can. Paper 68-50, 52p. Muller, J. E., and J. A. Jeletzky. 1970. Geology of the Upper Cretaceous Nanaimo Group, Vancouver Island and Gulf Islands, B. C , Geol. Surv. Can. Paper 69-25, 77p. Northcote, K. E. 1973. The bedrock geology of Vancouver Island. In Keser, N. and St. Pierre. S o i l s of Vancouver Island: a compendium. B. C. Min. of Forests, Res. Div., Res. Note No. 56, (unpaged). Northcote, K. E., and J. E. Muller. 1972. Volcanism, Plutonism and Mineralization: Vancouver Island, Can. Inst, of Min. and Met. B u l l . , October, 1972. Noy-Meir, I., and R. H. Whittaker. 1977. Continuous multivariate methods in community analysis: some problems and developments. Vegetatio 33: 79-98. Odum, E. P. 1950. Bird populations of the Highlands (N. Carolina) Plateau in relation to plant succession and avian invasion. Ecology 31: 587-605. Ogil v i e , R. T. 1963. Ecology of spruce forests on the east slope of the Rocky mountains in Alberta. Ph.D. thesis, Wash. State Univ., Pullman, 189p. O r l o c i , L. 1961. Forest types of the Coastal Western Hemlock zone. M.Sc. thesis, Dept. of B i o l , and Botany, Univ. of B. C , Vancouver, 206p. O r l o c i , L. 1964. Vegetation and environmental variation in the ecosystems of the Coastal Western Hemlock zone. Ph.D. thesis, Dept. of B i o l , and Botany, Univ. of B. C , Vancouver, 199p. O r l o c i , L. 1978. Multivariate Analysis in Vegetation Research. 2nd e d i t i o n . Junk, The Hague, Netherlands, 451p. Packee, E. C. 1976. An ecological approach toward y i e l d optimization through species a l l o c a t i o n . Ph.D. thesis, Faculty of Graduate School, Univ. of Minnesota, 740p. + Appendices. Packee, E. C. 1979. Keys to the Vegetation Zones and Vegetation Series of south coastal B r i t i s h Columbia. MacMillan Bloedel Limited, unpublished internal report, 19p. Packee, E. C. 1981. Coastal B r i t i s h Columbia and southeast Alaska - vascular plant names and symbols for ecosystem inventory and analysis. MacMillan Bloedel Limited For. Res. Note No. 4, February, 1981., 70p. 188 Peterson, E. B. 1964. Plant associations in the subalpine Mountain Hemlock zone in southern B. C. Ph.D. thesis, Univ. of B. C., Vancouver, 171p. P f i s t e r , R. D. 1977. Ecological c l a s s i f i c a t i o n of forest land in Idaho and Montana. Ijn Proc. Ecol. C l a s s i f . of For. Land in Canada and Northwestern U. S. A., Univ. of B. C , Vancouver, pp. 329-358. P f i s t e r , R. D., B. L. Kovalchik, S. F. Arno, and R. C. Presby. 1977. Forest habitat types of Montana. USDA For. Serv., Gen. Tech. Rept. INT-34, 174p. Intermountain For. and Range Exp. Stn., Ogden, Utah. P f i s t e r , R. D., and S. F. Arno. 1980. C l a s s i f y i n g forest habitat types based on potential climax vegetation. For. S c i . 26(1): 52-70. Pielou, E. C. 1977. Mathematical Ecology. Wiley-Interscience, New York. Resource Analysis Branch. 1976. Vegetation data form, 2nd Draft, January, 1976. B. C. Min. of Environ., Res. Anal. Br., 4p. Resource Analysis Branch. 1979. Climatic data for short term clima t o l o g i c a l stations, (unpublished). Roehmer, H. 1972. Forest vegetation and environments on the Saanich peninsula. Ph.D. thesis, Dept. of Biology, Univ. of V i c t o r i a , 405p. Rowe, J. S. 1972. Forest regions of Canada. Can. Dept. Environ., Can. For. Serv. Publ. No. 1300, 172p. Schofield, W. B. 1979. Nanaimo bryophytes: a preliminary che c k l i s t and key to hepatics and mosses. Univ. of B. C , Dept. of Botany, 22p. (unpublished). Scholander, P. F., H. T. Hammel, E. D. Bradstreet, and E. A. Hemmingsen. 1965. Sap pressure in vascular plants. Science 148: 339-346. Shimwell, D. W. 1971. The description and c l a s s i f i c a t i o n of vegetation. Univ. of Wash. Press, Seattle, 322p. Smith, D. M. 1962. The Practice of S i l v i c u l t u r e , 7th E d i t i o n . John Wiley and Sons, Inc., New York 578p. Society of American Foresters. 1967. Forest cover types of North America (exclusive of Mexico). Soc. of Amer. For., Washington, D. C., 67p. S o i l Survey St a f f . 1975. S o i l Taxonomy. U. S. Dept. Agric. Handbook No. 436. U. S. Govt. Print. Office, Washington, D. C , 754p. 189 Sorensen, T. A. 1948. A method of establishing groups of equal amplitude in plant sociology based on s i m i l a r i t y of species content. K. Danske Vidensk Selsk. B i o l . Skr. 5(4): 1-34. Stevenson, J. S. 1945. Geology and ore deposits of the China Creek area, Vancouver Island, B r i t i s h Columbia. B. C. Min. of Mines Annual Report, 1944, pp. 142-161. Sukachev, V. N. 1944. Pr i n c i p l e s of genetic c l a s s i f i c a t i o n in biogeocoenology. Zh. Obshch. B i o l . 5(4): 213-227. Moskva. Sukachev, V. N., and H. D y l i s , (eds.). 1964. Fundamentals of Forest Biogeocoenology. Oliver and Boyd Ltd., Edinburgh and London, 672p. (Translated from the Russian by J . M. MacLennan.) Szczawinski, A. F. 1953. Corticolous and lig n i c o l o u s plant communities in the forest associations of the Douglas-fir forest on Vancouver Island. Ph.D. thesis, Dept. B i o l , and Botany, Univ. of B. C , Vancouver, 283p. Tansley, A. G. 1935. The use and abuse of vegetational concepts and terms. Ecology 16: 284-307. Tansley, A. G., and T. F. Chipp. 1926. Aims and Methods in the Study of Vegetation. The B r i t i s h Empire Vegetation Committee. Whitefriars Press, London. 383p. Taylor, R. L., and B. MacBryde. 1977. Vascular plants of B r i t i s h Columbia. The Univ. of B. C. Press, Vancouver, 488p. Trewartha, G. T. 1968. An introduction to climate, 4th Ed i t i o n . McGraw-Hill Book Co., New York 408p. Utzig, G., D. MacDonald, and P. Comeau. 1978. Ecological c l a s s i f i c a t i o n for the Nelson Forest D i s t r i c t , 2nd Approximation, B. C. Min. of Forests, July, 1978, 33p. + Appendices. Valentine, K. W. G., and L. M. Lavkulich. 1978. The s o i l orders of B r i t i s h Columbia. I_n Valentine, K. W. G. et al.. (eds.). The S o i l Landscapes of B r i t i s h Columbia. B. C. Min. of Environ., Res. Anal. Br., V i c t o r i a , pp. 67-95. Valentine, K. W. G., P. N. Sprout, T. E. Baker, and L. M. Lavkulich. 1978. The S o i l Landscapes of B r i t i s h Columbia. B. C. Min. of Environ., Res. Anal. Br., V i c t o r i a , 197p. Wade, L. K. 1965. Vegetation and history of the Sphagnum bogs of the Tofino area, Vancouver Island. M.S. thesis, Univ. of B. C , Vancouver, 125p. Waring, R. H. 1969. Forest plants of the eastern Siskiyous: their environmental and vegetational d i s t r i b u t i o n . Northwest S c i . 43: 1-17. Waring, R. H., and J. Major. 1964. Some vegetation of the 190 C a l i f o r n i a redwood region in re l a t i o n to gradients of moisture, nutrients, l i g h t and temperature. Ecol. Monogr. 34: 167-215. Waring, R. H., and B. D. Cleary. 1967. Plant moisture stress: evaluation by pressure bomb. Science 155: 1248-1254. Waring, R. H., K. L. Reed, and W. H. Emmingham. 1972. An environmental grid for c l a s s i f y i n g coniferous forest ecosystems. Proc: Res. on Coniferous Forest Ecosystems - a symposium. Bellingham, Wash. March 23-24, 1972. Wellington, W. G. 1965. The use of cloud patterns to outline areas with d i f f e r n t climates during population studies. Can. Ent. 97(6): 617-631. Whittaker, R. H. 1948. A vegetation analysis of the Great Smoky mountains. Ph.D. thesis, Univ. of I l l i n o i s , Urbana. Whittaker, R. H. 1956. Vegetation of the Great Smoky mountains. Ecol. Monogr. 26: 1-80. Whittaker, R. H. 1960. Vegetation of the Siskiyou mountains, Oregon and C a l i f o r n i a . Ecol. Monogr. 30: 279-338. Whittaker, R. H. 1962. C l a s s i f i c a t i o n of natural communities. Bot. Rev. 28: 1-239. Whittaker, R. H. 1967. Gradient analysis of vegetation. B i o l . Rev. 42: 209-264. Whittaker, R. H. (ed.). 1973. Ordination and C l a s s i f i c a t o n of Communities. Handbook of Vegetation Science 5: 1-737. The Hague: W. Junk. Whittaker, R. H. 1975. Communities and Ecosystems. MacMillan Publ. Co., Inc. New York 385p. Whittaker, R. H., and W. A. Niering. 1964. Vegetation of the Santa Catalina mountains, Arizona. I. Ecological c l a s s i f i c a t i o n and d i s t r i b u t i o n of species. J. Arizona Acad. S c i . 3: 9-34. Whittaker, R. H., and W. A. Niering. 1965. Vegetation of the Santa Catalina mountains, Arizona. I I . A gradient analysis of the south slope. Ecology 46: 429-452. Zobel, D. B., A. McKee, G. M. Hawk, and C. T. Dyrness. 1976. Relationships of environment to composition, structure, and d i v e r s i t y of forest communities of the central western Cascades of Oregon. Ecol. Monogr. 46: 135-156. 190a APPENDICES 191 APPENDIX I'. Keys to the Vegetation Zones and Vegetation Series of south coastal B r i t i s h Columbia (Packee 1 9 7 9 ) . Key 2 Vegetation Zones of South Coast B r i t i s h Columbia THIS. KEY'IS APPLICABLE TO CLIMATIC CLIMAX OR NEAR CLIMATIC CLIMAX STANDS FOUND ON MODAL SITES OVER A BROAD GEOGRAPHIC AREA. The key i s based strongly on diagnostic c o n i f e r s : Abam, Abgr, Abla, Pisi , Pico, Psme, Tshe and Tsme. Be sure that f i r e has not eliminated a diagnostic species. A. Psme and/or Abgr, Pico and Thpl present and reproducing s u c c e s s f u l l y (>25 stems/ha); Psme common overstory domi-nant ... B B. Tshe r a r e l y present; Psme and/or Abgr reproducing suc-c e s s f u l l y ; Psme and/or Pico common overstory dominants . . . . PSME t/Z (Key 5) BB. Tshe reproduction abundant, however, trees only occas-s i o n a l l y reach a codominant canopy p o s i t i o n ; often spike-topped Abgr reproducing s u c c e s s f u l l y ; s u c c e s s f u l reproduction of Psme rare to marginal; Psme and/or Abgr common overstory dominants... P S M E - T S H E VI (Key 5) AA. Psme not reproducing s u c c e s s f u l l y (<25 stems/ha); Tshe and/or Tsme and/or Abam reproducing s u c c e s s f u l l y ; Psme may or may not be present i n overstory...C C. Tshe reproducing s u c c e s s f u l l y ; Tsme may be present, but.of l e s s e r importance ...D D. Tshe reproduction dominant; Abgr may be present; Thpl present, reproduction v a r i a b l e ; Abam- absent or rare, reproduction l a r g e l y unsuccessful; Psme, Tshe and Thpl common overstory dominants...TSHE VI {Key 6) DD. Tshe and Abam reproducing success f u l l y . . . E E. Abam reproduction (unless f i r e h i s t o r y ) that of Tshe; Thpl and/or Clmo present; heavy win-ter snowpack l i n g e r i n g i n t o mid-spring... A B A M - T S H E VI [Key 3) EE. Abam reproduction < that of Tshe except i n areas of cold a i r drainage and along some streams; Thpl common; Chno rare or absent; l i g h t winter snowpack seldom l i n g e r i n g i n t o spring. . . F 192 APPENDIX I (continued) F. Pi si- and Pico infrequent except on certain edaphic situations; summer fog uncommon (burns off by late morning)...TSHE-ABAM VI (Key 2) FF. Pisi and Pico common; Pico and Thpl repro-duction frequently successful; Pisi common pioneer, but only marginally successful; cool summer, onshore flow of marine a i r and summer fog common... TSHE-PISI VI CC. Tsme reproducing successfully; Tshe may be present but of lesser importance...G G. Tshe and Abam reproducing successfully; reproduction of Tshe=Tsme; Tshe trees overtop Tsme; winter snowpack lingers only u n t i l late spring... ABAM-TSHE VI (Key 3) GG. Tshe absent or dis t i n c t l y < Tsme; Tsme reproducing successfully; Abam present or absent; heavy winter snowpack com-monly lingers into early or mid summer . . .H H. Abam reproduction dominant; Abam over-tops Tsme; Tsme more important than Tshe (numbers and size); closed canopy forest... ABAM-TSME VI {K&y 4) HH. Tsme dominant; Abam d i s t i n c t l y minor i n importance or absent; Abla common pioneer; Pial may be present; not closed canopy forest - forest clumps and meadows, approaches tree l i n e . . . TSME-ABLA VI (Key 9) 1979 APPENDIX I (continuedL. Key 3 Vegetation Series of the Abam-Tshe Vegetation Zone A. Habitats confined to avalanche areas; Alsi dominates s i t e ; conifer reproduction may be present, but unsuc-cessful in gaining dominance... ALSI VS AA. Habitats not confined to avalanche runs; conifers pre-sent and capable of dominating...B B. Tsme reproducing more successfully than Tshe...C C. Abam may be present and regenerating, but rele-gated to understory; Chno may be present, but limited in quantity (less than 5% of stems in stand) and relegated to understory... TSME VS CC. Abam reproducing successfully; Tshe may be pre-sent but reproducing less successfully than Tsme ABAM-TSME VS BB. Tsme reproducing less successfully than Tshe or ab-sent .. . D D. Abam present and reproducing successfully...E E. Abam nearly sole tree species, reproducing successfully... ABAM VS EE. Other conifers reproducing successfully in association with Abam...F F. Abam reproduction more successful than Tshe; Chno cover class di s t i n c t l y greater than Thpl cover class... ABAM-TSHE- {CHNO) VS FF. Both Abam and Tshe reproduction success-f u l ; Chno absent or poorly represented; Thpl commonly present...G G. Abam reproduction more successful than Tshe; Thpl minor to co-climax... ABAM-TSHE-(THPL) VS GG. Abam reproduction less successful than Tshe; Thpl minor to co-climax... TSHE-ABAM VS DD. Abam poorly represented and reproducing unsuc-cessfully. . .H APPENDIX I (continued) Chno sole conifer reproducing suc-cessfully; other conifers dis-t i n c t l y i n f e r i o r .. . CHNO VS Chno reproducing unsuccessfully . . .1 I. Tshe reproducing successfully • • • J J. Tshe major dominant, may share climax with Thpl...TSHE VS JJ. Tshe reproducing successfully, but commonly damaged severely by drought; Thpl present and frequently reproducing suc-cessfully; Psme a late serai species frequently dominat-ing overstory, but reproduc-tion only marginally success-f u l ; Abgr may be present... THPL-TSHE-IPSUE) VS K. Thpl dominant, reproducing successfully to poorly; no other conifer reproducing successfully... THPL VS KK. Pico and/or Psme reproduc-ing. . .L L. Pico reproducing suc-cessfully; Thpl and/or Psme reproducing, but rarely reaching the overstory canopy... PICO VS LL. Psme reproducing suc-cessully; warm slopes; Pico may be present... PSME VS H. HH. 1979 APPENDIX I (continued!. Key 4 Vegetation Series of the Abam-Tsme Vegetation Zone A. Habitats confined to avalanche runs; Alsi dominates sit e ; conifer reproduction may be present, but unsuccessful in gaining dominance... ALSI VS AA. Habitats not confined to avalanche runs; conifers present and capable of dominating...B B. Abla and Tsme reproducing s u c c e s s f u l l y . . . T S M E - A B L A VS BB. Abla may be present but reproducing unsuccessfully; other conifers reproducing successfully...C C. Chno sole conifer reproducing successfully; other conifers di s t i n c t l y i n f e r i o r . . . CHUO VS CC. Other conifer reproduction > that of Chno...D D. Tsme reproducing successfully; Abla reproduction absent to marginally successful; Abam relegated to understory; Chno may be present in limited quantity (less than 5% of stems in stand) and relegated to understory... T S M E VS DD. Abam reproducing successfully...E E. Abam nearly sole tree species; reproducing successfully... A B A M VS EE. Other conifers reproducing successfuly in association with Abam. ..7 F. Abam and Tsme reproducing successfully; Abam dominant; Tshe reproducing less suc-cessfully than Tsme... A B A M - T S M E VS FF. Abam and Tshe reproducing successfully; Abam dominant; Tsme reproducing less suc-cessfully than Tshe or absent... A B A M - T S H E - ( C H N O ) VS 19 79 APPENDIX I (continued) G. Psme r e p r o d u c t i o n m a r g i n a l l y succe s s -f u l ; s tands dominated by Tshe and Thpl r e p r o d u c t i o n ; Abgr r e p r o d u c t i o n u s u a l l y p r e s e n t . . . TH?L-TSHE-(PSME) VS GG. Psme and Abgr not r e p r o d u c i n g succe s s -f u l l y ; Abgr f r e q u e n t l y not p r e s e n t ; Tshe o b v i o u s l y c l i m a x ; Thpl p re sent as c o - c l i m a x . . . TSHE VS 1979 APPENDIX I .(continued) 197 Key 5 Vegetation Series of the Psme and Psme-Tshe Vegetation Zones A. Habitats dominated by hardwoods; conifer reproduction may be present, but unsuccessful in gaining dominance . . .B B. Alru dominates s i t e ; reproduction may be only margin-ally successful; along lowest terraces or flood plains of streams and some Aquic (or Peraquic) SMR's. . . ALRU VS BB. Quga present and reproducing; Psme seedlings present, but unsuccessful i n gaining dominance; confined to Gulf Islands, southeast coastline Vancouver Island, Saanich Peninsula... QUGA VS AA. Habitats not dominated by hardwoods; conifer reproduction successful...G C. Pico nearly sole dominant, reproducing successfully, Thpl and/or Psme largely reproducing unsuccessfully, rarely gain codominant position; on Xeric, Aquic or Peraquic SMR's or Lit h i c s o i l s . . . PICO VS CC. Pico not sole dominant; not reproducing successfully . .. D D. Psme reproducing successfully, nearly sole domi-nant; Abgr rare or absent; Thpl present or absent PSME VS DD. Psme reproducing marginally successfully to unsuc-cessfully; greater coverage by reproduction of other species. . . E E. Abgr reproducing successfully; Thpl may be present as co-climax; Tshe reproduction ab-sent or unsuccessful, coverage di s t i n c t l y less than Abgr... ABGR US EE. Abgr not reproducing successfully, or i f i t i s , greater coverage by more shade-tolerant spe-cies .. . F F. Thpl sole dominant, no other conifer re-producing successfully; Thpl reproduction marginally successful... THPL VS FF. Thpl not sole dominant; Tshe Abgr and/or Psme present...G 198 APPENDIX J; (continued). GG. Psme not part of the climax...H H. Abam not present and not reproduc-ing; Tshe obviously climax; Fnpl present as co-climax (occasionally replaced by Chno at higher eleva-tions... TSHE VS HH. Abam reproducing successfully...I I. Abam reproduction less successful than Tshe; Thpl present in climax TSHE-ABAM VS II. Abam reproduction more successful than Tshe; Thpl or Chno present .. .J J. Thpl minor to co-climax, cover class equal to or greater than Chno; Chno commonly absent... A B A M - T S H E - [THPL] VS JJ. Chno cover class distinctly greater than Thpl cover class A B A M - T S H E - ( C H M O ) VS 1979 199 APPENDIX I (continued) Key 6 Vegetation Series of the Tshe Vegetation Zone A. Habitats dominated by hardwoods ; conifer reproduction may be present, but unsuccessful i n gaining dominance...B B. Alsi dominates sit e ; habitats confined to avalanche runs/associated talus... ALSI VS BB. Aim dominates sit e ; reproduction may only be margin-all y successful; along lowest terraces or floodplains of streams and some Aquic SMR's... A L R U VS AA. Habitats not dominated by hardwoods; conifer reproduction successful...C C. Pico nearly sole dominant, reproducing successfully; Thpl and/or Psme largely reproducing unsuccessfully; rarely gain codominant position; on Xeric, Aquic or Peraquic SMR's or Lithic s o i l s . . . P I C O VS CC. Pico not sole dominant, not reproducing successfully . . . D D. Psme reproducing successfully, nearly sole domi-nant; Abgr rare or absent; Thpl -present or ab-sent... P S M E VS DD. Psme reproducing marginally successfully to un-successfully; greater coverage by reproduction of other species...E E. Abgr reproducing successfully; Thpl may be present as co-climax; Tshe reproduction un-successful, coverage di s t i n c t l y less than Abgr... A B G R VS EE. Abgr not reproducing successfully, or i f i t i s , greater coverage by more shade tolerant species ...F F. Thpl sole dominant, no other conifer repro-ducing successfully; Thpl reproduction marginally successful... THPL VS FF. Thpl not sole dominant; Tshe, Abam and/or Psme reproducing...G G. Psme reproduction marginally successful, stands dominated by Tshe and Thpl repro-duction; Abgr reproduction usually present... THPL-TSHE-(PSME) VS 2 0 0 APPENDIX I (continued) GG. Thpl not sole dominant; Tshe, Abam and/or Psme reproducing successfully . . . H H. Psme reproduction marginally success-f u l , stands dominated by Tshe and Thpl reproduction; Abgr reproduction present or absent...THPL-TSHE-(PSME) VS HH. Psme not part of the climax...I I. Abam not present and not reproducing; Tshe major species reproducing suc-cessfully; Thpl present or absent T S H E VS II. Tshe always present; Abam present or absent...J J. Pico lingers into climax; repro-duction marginally successful to unsuccessful; confined to Aquic or Peraquic SMR1 s; Thpl most successful conifer; Psme absent THPL-TSHE-[FTCO) VS JJ. Pico not common in climax stand, Abam present...K K. Abam reproduction coverage greater than that of Tshe; Thpl present in climax... ABAM-TSHE-[THPL] VS KK. Abam reproduction coverage less than that of Tshe; Thpl present i n climax... T S H E - A B A M VS 1979 APPENDIX I (continued! Key 7 Vegetation Series of the Tshe-Abam Vegetation Zone A. Habitats dominated by hardwoods; conifer reproduction may be present, but unsuccessful in gaining dominance ...B B. Alsi dominates s i t e ; habitats confined to avalanche runs/associated talus... A L S I VS BB. Aim dominates s i t e ; reproduction may be only margin-al l y successful; along lowest terraces or floodplains of streams and some Aquic SMR's... A L R U VS AA. Habitats not dominated by hardwoods; conifer reproduction successful...C C. Pico reproducing successfully, at least equal to associates...D D. . Pico nearly sole dominant and reproducing success-f u l l y ; Thpl or Psme largely reproducing unsuccess-f u l l y , rarely gain codominant position; on Xeric, Aquic or Peraquic SMR's or L i t h i c s o i l s . . . PI CO VS DD. Pico not sole dominant, at least equal to associ-ates {Thpl, Abam, Tshe); Tshe almost completely lacking i n overstory... P1C0-THPL VS CC. Pico not reproducing successfully or i f i t i s , dis-t i n c t l y i n f e r i o r to i t s associates and on Aquic SMR • • • E E. Psme reproducing successfully, nearly sole domi-nant; Abgr rare or absent; Thpl present or ab-sent... P S M E VS EE. Psme reproducing marginally successfully to unsuccessfully; greater coverage by reproduc-tion of other species...F F. Abgr reproducing successfully; Thpl may be present as co-climax; Tshe reproduction un-successful, coverage dist i n c t l y less than Abgr... A B G R VS FF. Abgr not reproducing successfully or i f i t i s , greater coverage by more shade-tolerant species...G G. Thpl sole dominant, no other conifer re-producing successfully; Thpl reproduction marginally successful... T H P L i^S APPENDIX I (continued) Key 8 Vegetation Series of the Tshe-Pisi Vegetation Zone A. Alvu overstory dominant, reproduction may only be mar-ginally successful; conifer reproduction may be present, but unsuccessful in gaining dominance; along lowest ter-races or floodplains of streams and some Aquic (or Per-aquic) SMR's... A L R U VS AA. Alvu not climax; conifer reproduction successful...B B. Pico reproducing successfully to marginally success-f u l l y . .. C C. Pico nearly sole dominant and reproducing success-f u l l y ; Thpl and occasionally Chno reproducing but rarely gain dominant position; on Aquic or Peraquic SMR's or Lithic s o i l s . . . P I C O VS CC. Pico not sole dominant, reproducing successfully to marginally successfully in association with Thpl, Tshe and Abam.. .D D. Pico regeneration at least equal to associates; Tshe and Abam almost completely lacking in over-story... PICO-THPL VS DD. Pico lingers into climax; reproduction marginally successful to unsuccessful; Thpl most successful and dominant conifer; Tshe always present; Abam present or absent... THPL-TSHE-[PICO] VS BB. Pico not present i n climax stand, reproducing unsuc-cessfully. .. '£ E. Pisi nearly sole dominant and reproducing successfully to marginally successfully; confined to outer coast... P I S I VS EE. Pisi not sole dominant, reproducing unsuccess-f u l l y ; reproduction dominated byThpl, Tshe and/or Abam...F F. Thpl dominant, reproducing successfully to poorly; no other conifer reproducing suc-cessfully... T H P L VS FF. Tshe and Abam reproducing successfully...G G. Abam reproduction coverage greater than that of Tshe; Thpl present in climax A B A M - T S H E - ( T H P L ) VS GG. Abam reproduction coverage less than that of Tshe; Thvl present i n climax T S H E - A B A M VS APPENDIX I (continued) Key 9 Vegetation Series of the Tsme-Abla Vegetation Zone A. Habitats confined to avalanche runs; Alsi dominates s i t e ; conifer reproduction may be present, but unsuc-cessful i n gaining dominance... A L S I VS AA. Habitats not confined to avalanche runs; conifers present and capable to dominating...B B. Vial sole tree species present and reproducing; main-land only.. . P I A L VS BB. Vial may be present but reproducing unsuccessfully; other conifers reproducing successfully...C C. Abla reproducing successfully; Tsme absent, scarce, or incapable of dominating; commonly parkland... A B L A VS CC. Tsme reproducing successfully...D D. Tsme and Abla both reproducing successfully... T S M E - A B L A VS DD. Abla absent or infrequent; reproducing unsuccess-f u l l y to marginally successfully... E E. Tsme reproducing successfully; Abla reproduc-tion absent to marginally successful; Abam, i f present, relegated to understory; Chno may be present in limited quantity (less than 5% of stems in stand) and relegated to under-story... T S M E VS EE. Abam and Tsme present and reproducing success-f u l l y . . . A B A M - T S M E VS 1979 2 0 4 APPENDIX I I : Canopy layers and v i g o r , abundance and s o c i a b i l i t y c l a s s e s V = V e t e r a n s — t r e e s above the general l e v e l of crown cover remaining from a previous stand. D = Dominants—trees with crowns extending above the general l e v e l of crown cover. C = Codominants—trees forming the general l e v e l of crown cover. I = Intermediates—trees shorter than dominants and codominants, but with ' crowns extending i n t o the canopy cover formed by these l a y e r s . S = Supressed—trees with crowns e n t i r e l y below the general l e v e l of crown cover, showing signs of reduced vigor. U = U n d e r s t o r y — t r e e s below the main canopy layers (including reproduction greater than 1.5 meters) showing no signs of reduced v i g o r . Table A2. Understory and tree canopy layers unsed i n f i e l d t a l l y and used i n f i e l d observations and data a n a l y s i s . Table A l . Tree canopy layers used i n f i e l d t a l l y (after Smith 1962). data a n a l y s i s (Brooke et. a l . 1970). A layer Dominant and codominant trees (includes veterans). Intermediate t r e e s . Suppressed trees over 9 m. i n height. B layer Saplings* and shrubs between 1.8 m. and 9 m. i n height. Shrubs and woody plants 15 cm. to 1.8 m. i n height. C layer Small woody plants l e s s herbaceous p l a n t s . than 15 cm. i n height and a l l D la y e r Bryophytes, lichens and seedlings. *saplings were t a l l i e d i n the A l a y e r . 20 APPENDIX II (continued). Table A3. Vigor, s o c i a b i l i t y and abundance scales used i n f i e l d observations. 1. Vigor r a t i n g s (Peterson 1964) 0 Species dead + Vigor poor 1 Vigor f a i r 2 Vigor good 3 Vigor e x c e l l e n t 2.' Abundance scale (Tansley and Chipp 1926, as c i t e d i n Shimwell 1971) 1 Rare 2 Occasional 3 Frequent 4 Abundant 5 Very abundant 3. S o c i a b i l i t y classes (Shimwell 1971). 1 Growing i n one place, s i n g l y 2 Grouped or t u f t e d 3 In troops, small patches or cushions 4 In small colonies, i n extensive patches or forming carpets 5 In great crowds or pure populations 206 Appendix I I I : L a t i n and common names of species found on sample p l o t s . ABAM kbZeA amabZtZA (Dougl.) Forbes amabilis f i r ABGR kbZzA ghXXndZA (Dougl.) Forbes grand f i r ABLA2 AbZeJ, ZoAZocaApa (Hook.) Nutt. alpine f i r ACGL AcQA. gZabAum Torr. Rocky Mountain maple ACMA AceJi macAOphyZZum Pursh b i g l e a f maple ACMI AchZZZe.0. mZlZe.^oZZum L. common yarrow ACTR Achlys tnlphyUiCL (Smith) D.C. v a n i l l a leaf ACRU Aetata hxxhhja. (Ait.) W i l l d . W. red baneberry ADBI Acfe.noCOuZon bZcoZoh. Hook. t r a i l - p l a n t ADPE AdZantum po.daXxxm L. northern maidenhair fern AGROP AgK.opyh.OVi spp. Gaertn wheatgrass AGTE kgh,0AtZA te.nUAJ> S i l o t h c o l o n i a l bent grass ALRU kZnuA hxibha. Bong. red alder ALVI AZZotAopa VAAgata T.& G. candy s t i c k AMAL AmeIxX.YiC.hlQA. oZnZ&oZZa Nutt. Saskatoon ANLY2 Anemone ZyaZZZ B r i t t . L y a l l ' s anemone ARME AAbutuA me.nzZeJ>ZZ Pursh P a c i f i c madrone ARC03 kuctOAtaphyZoA columbZana Piper b r i s t l y manzanita ARXM AActo<AtaphyZoA me.dZa Greene intermediate manzanita ARUV Afldt0istxx.phyl.06 UVCL-UAAZ (L.) Spreng kinnikinnick ARMA AAQJWAAJX macAophytta Hook, bigleaved sandwort ARLA khnZca. ZatZ&oZZa Bong. broad-leaved arnica ARSY khxincuA &ytvQJstOA Kostel. Sylvan goat's beard ATFI kthyAixxm ^ Ztix-^QjnZna (L.) Roth lady fern BEAQ BeAbejiZi, aquZ&oZZum Pursh t a l l Oregon grape BENE BeAbeAAA neAVOAa Pursh d u l l Oregon grape BLSP BZzchnum ApZcant (L.) Roth deerfern BOHO BoAchnZakZa hookeAZ Walpers Vancouver ground cone BRIN BKOmuA ZneAmZi Leys Hungarian brome grass BRPA BnomUA pacZ^ZcuA Shear P a c i f i c brome grass BRVU BtiomuA vuZgaAAj> (Hook.) Shear Columbia brome grass BROMU BKOmuA spp. (L.) brome grass CABU2 CaZypAO buZboAa (L.) Oakes f a i r y s l i p p e r CASC2 Campanula ACOuZeAZ Hook. Scouler's harebell CAREX CaAQX spp. L. sedge CHNO Chama.ZcypaAZ& nOOtkaXenAZi, (D.Don) Spach. yellow cypress CHME ChAjnaphZZtx. me.nzZeJ>ZZ (R.Br.) Spreng Menzies'pipissewa CHUM ChZmaphZZa umbeJLZxxXa (L.) Bart common pipsissewa CLPY CZa.dotha.mnuM pyh.oZae.kZoh.UA Bong, copperbush CLUN CZZntonZa unZ^Zoha. (Schult.) Kunth. blue-bead c l i n t o n i a COHE CoZZomZd heXeAophyZta Hook diverse-leaved collomia COAS Coptic OApZcnZ^oZZa S a l i s b . spleenwort-leaved gold thread COMA3 CohxxZZotihZza. macuZata Raf. spotted c o r a l root COME CoKaZZohhZza. meAte.nAZa.na Bong, western c o r a l root CORAL CoH.aZZon.hZza spp. Chat. c o r a l root COCA CoAnUA can.ade.nAZ6 L. western bunchberry CONU CoA.na6 nuXtxxZZZZ Aud. western flowering dogwood CYSC CytZisUA ACOpaAJJXA (L.) Link scotch broom DASP VanthonZa ApZcaXa (L.)Beauv. poverty oatgrass DEEL VeAchampAZo. eZongata (Hook.) Munro slender hairgrass DRAU2 VfiyopteAZA auAthAXtca Adans. spiny shieldfern ELGL EZymuA gZaucuA Buckl. blue wild-rye 207 APPENDIX I I I (continued). ELYMU EPAN EQUIS FEOC FRVE FRCA GATR GALIU GAOV GASH GERAN GOOB GYDR HEGL2 HEMI HIALY HOD I HYRA HYMO JUNCU JUC04 LAMU LANE LIC04 LIB02 LICA3 LIC03 LISTE LOCI LOHI LOUT2 LONIC LUPE LUPA LUZUL LYCL LYAM MASA MADI2 MESU MEFE MOUN2 MOPA MOPE MOSI NONE OPHO OSCH PAMY PERA PHEM PISI PICO ElymuA spp. L. wildrye Epitobium anguAtifiollum L. fireweed Equisztum spp. L. h o r s e t a i l FzAtuca OCCA.d2.ni/xJLU> Hook, western fescue FhJXgaAixx. VZAca L. wood strawberry F K i i i l l o A i a COMchai.C2.nAi!> (L.) Ker-Gawl r i c e r o o t f r i t i l l a r y Galium tAl^lohxxm Michx. sweet-scented bedstraw Galium spp. L. bedstraw GaulihzAia OVOti&olia Gray oregon wintergreen GaulthzAia Ahallon Pursh s a l a l GzAanium spp. L. geranium Goody2Aa oblongi&olia Raf. large-leaved rattlesnake orchid GymnocaApixxm OAyoptzAiA (L.) Newm. oakfern WzuchzAa glabfia W i l l d . smooth alumroot HzucheAa micAantka Dougl. small-flowered alumroot HizAacium albifalohum Hook, white hawkweed HolodiACUA discoloh. (Pursh) Maxim, oceanspray HypochacAiA ftadicata L. common cat's ear HypopityA monotAopa Crantz. fringed pinesap JuncuA spp. L. rush lunipzAUA communis L. common juniper Lactuca muAalii, (L.) Fresen. wall l e t t u c e LothyhUA n2.vad2.nAlA Wats, purple nevada peavine LlJUdim columbianum Hanson Columbia l i l y Linnaea boficaliA L. northern twinflower LiiteAa cauAina Piper northwestern twayblade LiiteAa COHdata. (L.) R.Br, heart-leaved twayblade LiAt2Aa spp. R.Br, twayblade LoniceAa cilioAa (Pursh) D.C. western trumpet honeysuckle Lonic2Aa kiipidula (Lindl.) Dougl. hairy honeysuckle LoniceAa utahznAiA Wats. Utah honeysuckle LoniceAa spp. L.. honeysuckle Luztkza pzctinata (Pursh) Kuntz. pa r t r i d g e f o o t Luzula paAvi^lohJX (Enrh.) Desv. small-flowered woodrush Luzula spp. D.C. woodrush Lycopodium clavatum L. running clubmoss LyAlchitum amZAicanum Hulten S St. John skunk cabbage Madia AOtlva Mol. tarweed Maianthzmum diiaiatum (Wood) Nels. & Macbr. f a l s e l i l y - o f - t h e - v a l l e y Mzlica Aubulata (Griseb.) Scribn. oniongrass MznzizAia ^ZAAuginza Smith P a c i f i c menziesia MonotAopa uni^lonja L. indianpipe MonOa paAvifiolia (Moc.) Greene small-leaved montia Montia p2A£oliata (Bonn) Howell miner's lettuce Montia Aibitica (L.) Howell S i b e r i a n spring beauty Nothochzlonz nzmoHOAa (Dougl.) Straw, woodland beard-tongue Oplopanax hohAidum (Smith) Miq. d e v i l ' s club OAmOhhiza cliilznAiA H. & A. mountain sweet c i c e l y PachyAtima myfiAinitZA (Pursh) Raf. oregon boxwood PzdijCulaAJLA HaczmoAa Dougl. s i c k l e t o p lousewort Phyllodocz 2mpztAlfaoHmli> (Sw.) D. Don red mountain heather P.tce.a AltchznAlA (Bong.) Carr s i t k a spruce PinuA COntOAta Dougl. lodgepole pine 208 APPENDIX III (continued). PIMO PZnuA montZcoZa Dougl. western white pine POA Poa spp. L. bluegrass POGL4 PoZypodZum gZyc.yM.hZza D.C. Eat. l i c o r i c e fern POL02 PoZyAtZchum ZonchZtZA (L.) Roth mountain h o l l y fern POMU PoZyAtZchum munZtum (Kaulf.) P r e s l . swordfern POTR2 PopuZuA thZchocaApa T. S G. western black cottonwood PREM PhunuA zmaAgZnata (Dougl.) Walp. b i t t e r cherry PSME PA2.lldot6U.ga mznzZzAZZ (Marbel) Franco. Douglas-fir PTAQ PtZhA.dZum aquZlZnum (L.) Kuhn. bracken fern PTAN PtzAoApoha andn.ome.dZa Nutt. pinedrops PYAS PyK-OZa, OAOAZ&oZZa Michx. common pink pyrola PYPI Pyh.oZa pZcta Smith white-veined pyrola PYSE Pyh-OZa. AZCUnda L. one-sided wintergreen RAUN2 RanunCuZuA uncZnatuA D. Don l i t t l e - f l o w e r e d buttercup RHPU RhamnuA puAAhZana D.C. cascara RHAL Rhodode.ndh.on aLbZ^Zonxxm Hook, white rhododendron RILA RZbZA ZacuAtAZ (Pers.) Poir black swamp gooseberry RIBES RZbZA spp. L. currant ROGY R06a gymnOCOApa Nutt. baldhip rose RUNI Rubo6 nZvaZZb Dougl. snow dewberry RUPA RubuA paAvZ^Zoh-UA Nutt. thimbleberry RUPE RubuA pzdaZuA J . E. Smith creeping raspberry RUSP RubuA ApZCtabZZZA Pursh salmonberry RUUR RubuA UAAZmiA Cham. & Schlecht p a c i f i c t r a i l i n g blackberry SARA SambucuA hacemoAa L. red elderberry SAMBU SambucuA spp. L. elderberry SETR Se.ne.cZo tAZanguZahAA Hook. arrowleaved ragwort SMRA SmZZacZna ha.ce.m06a (L.) Desf. f a l s e Solomon's seal SOSI Soh.buA AZtchznAZi Roemer s i t k a mountain ash STC04 Stachy6 COOtcyae. Heller Cooley's hedge-nettle STOC2 StcnanthZum OCcZdzntaZz Gray western mountain b e l l s STAM StAZptopUA ampZzxZ^oZZuA (L.) D.C. cucumber-root twisted-stalk STRO StAZptopuA h.0AZUA Michx. simple-stemmed twisted-stalk STST StAZptopuA AtAzptopoZdzA (Ledeb.) Frye & Rigg small twisted-stalk SYAL SymphohZcaApoA aZbuA (L.) Blake common snowberry SYMO SymphohA.cah.poA moZZZt, Nutt. t r a i l i n g snowberry TABR Tax.UA bh.ZvZkoZZa Nutt. western yew THOC JhaZZctAum OCcZdzntaZz Gray wester meadow-rue THPL Thuja pZZcata Donn. western red cedar TILA TZaAzZZa ZacZnZata (Hook.) Wheel cut-leaved foamflower TITR TZaAzZZa tAi^oZZata L. t r i f o l i a t e - l e a v e d foamflower TIUN TZaAzZZa unZfioZZata (Hook.) Kurtz. u n i f o l i a t e - l e a v e d foamflower TRCA3 ThJXJuZveXZeAZa caAoZZnZznAZA (Walt.) B a i l . f a l s e bugbane TRLA2 ThAizntaZZA ZatZ^oZZa Hook, broad-leaved starflower TROV ThJJLZZum ovatum Pursh western white t r i l l i u m TSHE TAuga hzteAophyZZa (Raf.) Sarg. western hemlock TSME TAuga mZAtznAZana (Bong.) Carr mountain hemlock VAAL VaccZnZum aZaAkaznAZ Howell Alaska blueberry VAME VaccZnZum mzmbh.ana.czum Dougl. black blueberry VAOV VaccZnZum OVOtZ^oZZum Smith oval-leaved blueberry VAPA VaccZnZum paAvZfioZZum Smith red huckleberry VASC2 VaZeAZana ACOUZZAZ Rydb. Scouler's v a l e r i a n VALER VoLzhAXXniX. spp. L. v a l e r i a n APPENDIX III (continued). VEVI VcAalAum vJAide A i t . f a l s e hellebore VEAM VoAOnlca amQAX.ca.na Schwein. American speedwell VIAM Vlcla amoAA-cana Muhl. American vetch VISA Vlcla 6atlva L. common vetch VIGL Viola glabella Nutt. yellow wood v i o l e t VIOR2 Viola otlblculaXa Geyer evergreen yellow v i o l e t VIOLA Viola spp. L. v i o l e t 1 / Nomenclature follows Hitchcock and Cronquist 1973. Abbreviations follow Garrison e t . a l . 1976 and Packee 1981. 2 1 0 APPENDIX IV: Cl i m a t i c s t a t i o n s , r a d i a t i o n t a b l e s , a v a i l a b l e water storage capacity tables and environmental data used f or input i n the Water Stress Index (WSI) model. Table A4. AES and RAB c l i m a t o l o g i c a l stations used for data extra p o l a t i o n . Map Code Station Elements Elev. No. Station Name No. Observed (m) A spec 1 Hollow (RAB)* 102511 P 411 E 2 Wowo (RAB) 102411 P,T 691 E 3 Campbell River A i r p o r t (AES) 1021261 P,T,S 105 f l a t 4 Cumberland (AES)@ 1022250 P,T 159 N 5 Tsable River FP (RAB) 102027 P,T 236 S 6 2E2500 (RAB) 102404 P,T 762 E 7 2E1400 (RAB) 102403 P,T 427 E 8 Cathedral (RAB) 102501 P 248 f l a t 9 A l b e r n i Lupsi Cupsi (AES) 1030210 P,T,S 9 W 10 F r a n k l i n River (RAB) 103403 P,T 177 SW 11 P a r k s v i l l e (AES) 1025970 P,T 82 NE 12 Sandy (RAB) 102506 P 58 W 13 Nanaimo Departure Bay (AES) 1025070' P,T,R 27. SE 14 Wellington (RAB) 102508 P 84 NE 15 Nanaimo A i r p o r t (AES) 1025370 P,T,S 30 f l a t 16 McKay Lake (RAB) 102004 P,T 290 N 17 Cassidy (RAB) 102001 P,T 201 N 18 Lookout High (RAB) 102128 P,T 1113 E 19 Lookout Low (RAB) 102127 P,T 823 NE 20 Cowichan Lake Forestry (AES) 1012040 P,T,S 177 f l a t * RAB @ AES P T S R Resource Analysis Branch short-term s t a t i o n Atmospheric Environment Service s t a t i o n P r e c i p i t a t i o n Temperature Bright Sunshine Total Solar Radiation 211 APPENDIX IV (continued) Table A5. Radiation Index table f o r Water Stress Index (WSI) c a l c u l a t i o n s (Ballard 1974). Radiation Radiation , Aspect Slope (%) Index Factor North 0 - 1 2 A 1.0 12-32 B 0.9 32-5 1 C 0.8 51-70 D 0.7 70+ E 0.6 Northeast, 0 - 1 8 A . 1 . 0 Northwest 18-40 B 0.9 40-60 C 0.8 60-84 D 0.7 84+ E 0.6 East, 0 - 3 6 A 1.0 West 36+ B 0.9 South, 0+ A 1.0 Southeast, Southwest ^ I n r e l a t i o n to r a d i a t i o n f o r 0% slope. 212 APPENDIX IV (continued) Table A6. Available Water Storage i n s o i l layers making up the root zone (Estimate thickness of organic layers or e f f e c t i v e thickness of mineral layers of various textures.) (from B a l l a r d 1974) . , — T e x t u r a l c l a s s e s — Tri xc Jens s s ^ . E f f e c t i v e thickness (cm) of mineral s o i l layers of organic AWSC layer (cm) s L S S L f S L L s i L C L c 1cm 4 10 8 6 4 4 3 3 3 2 8 20 15 13 9 7 6 6 6 3 12 30 23 19 13 11 9 9 9 4 16 40 31 25 17 14 11 12 12 5 20 50 38 31 22 18 14 15 15 6 ' 24 60 46 38 26 21 17 18 18 7 28 70 54 44 30 25 20 21 21 8 32 80 62 50 35 29 23 24 24 9 36 90 69 56 39 32 26 27 27 10 40 100 77 63 43 36 29 30 30 11 44 110 85 69 48 39 31 33 33 12 48 120 92 75 52 43 34 36 36 13 52 130 100 81 57 46 37 39 39 14 56 140 108 88 61 50 40 42 42 15 60 150 115 94 65 54 43 45 45 16 64 160 123 100 70 57 46 48 48 17 68 170 131 106 74 61 49 52 52 18 72 180 138 113 78 64 51 55 55 19 76 190 146 119 83 68 54 "58 58 20 80 200 153 125 87 71 57 61 61 21 84 162 131 91 75 60 64 62 22 88 169 138 96 79 63 67 67 23 92 177 144 100 82 66 70 70 24 96 185 150 104 86 69 73 73 25 100 192 156 109 89 71 76 76 26 200 163 113 93 74 79 79 27 169 117 96 77 82 82 28 175 122 100 80 85 85 29 181 126 104 83 88 88 APPENDIX IV (continued) Table A7. Environmental data f o r i n d i v i d u a l sample p l o t s . 213 AVERAGE WATER AVERAGE POT.ANNUAL CLIMATE STATIONS PLOT ELEV. MAY-SEP AWSC STRESS MAY-SEP SOLAR RADN. USED IN EXTRAPOL. NO. (m) PRECIP. (cm) INDEX TEMP(°C) 10 3cal/cm 2/yr P p t . d i s t . / s o l a r 1109 130 19.3 4.0 4.0+1.0 14.3 170 15l NA2 1110 260 21.8 6.5 1.5+2.5 13.5 128 15 NA 1111 185 21.8 6.5 1.0+3.5* 15.1 170 11 ND 3 1112 110 19.1 2.5 4.0+1.0 14.4 170 15 NA 1113 640 29.5 5.0 0.0+5.0 12.7 128 11 ND 1114 580 28.4 6.5 0.0+4.0 13.0 151 11 ND 1115 275 23.4 17.0 0.0+1.5 14.6 170 11 ND 1116 535 27.7 5.0 1.0+4.0 13.2 204 11 ND 1117 560 30.5 21.0 0.0+0.0 13.8 137 7 AL4 1118 650 31.8 7.0 0.5+3.5 13.3 170 7 AL 1119 300 24.6 11.0 0.0+0.5 13.5 105 . 8 AL 1120 430 26.7 17.5 0.0+0.0 12.8 105 8 AL 1121 60 19.1 11.0 1.5+2.5* 15.8 170 11 ND 1122 500 27.7 25.0 0.0+0.0* 12.4 181 8 AL 1123 580 29.0 8.0 0.5+2.5 12.0 170 8 AL 1124 365 24.9 3.0 5.0+0.0 14.2 204 11 ND 1125 185 21.8 10.0 1.5+2.0* 15.1 170 11 ND 1126 280 24.4 9.0 0.5+2.0 13.6 166 8 AL 1127 440 26.9 3.5 3.0+2.0 12.8 182 8 AL 1128 270 24.4 10.0 0.0+2.0 13.7 151 8 AL 1129 470 27.4 9.0 0.0+2.0 12.6 206 8 AL 1130 760 31.8 5.0 1.0+3.0 11.0 166 8 AL 1131 755 31.5 18.5 0.0+0.0 11.1 206 10 AL 1132 1085 36.6 8.0 0.0+1.0 9.3 206 10 AL 1133 940 34.5 10.0 0.0+0.0 10.1 159 . 10 AL 1134 660 30.2 7.5 0.5+2.0 11.6 206 8 AL 1135 240 23.9 12.5 0.0+0.0 13.9 131 8 AL 1136 510 27.9 15.5 0.0+0.0 12.4 170 10 AL 1137 680 30.5 10.0 0.0+0.0 11.5 105 10 AL 1138 610 29.5 22.0 0.0+0.0* 11.8 88 10 AL 1139 780 32.0 9.0 0.0+0.0 10.9 105 10 AL 1140 860 33.3 13.0 0.0+0.0 10.5 151 10 AL 1141 965 34.8 4.0 1.0+4.0* 9.9 137 10 AL 1142 1070 36.6 8.0 0.0+1.0* 9.3 137 10 AL 1143 500 26.4 18.5 0.0+0.0* 12.1 125 16 NA 1144 1180 39.1 10.5 0.0+0.0* 8.7 193 8 AL 1145 210 23.4 12.5 0.0+2.0 14.0 170 8 AL 1146 860 33.3 15.0 0.0+0.0 10.5 206 10 AL 1147 660 29.5 9.5 0.0+0.5 11.7 125 19 NA 1148 580 27.9 3.0 4.0+1.0* 12.1 128 19 NA 1149 740 31.0 7.5 0.0+1.0 11.3 128 19 NA 1150 820 32.5 19.0 0.0+0.0 10.8 170 19 NA 1/ See c l i m a t i c s t a t i o n numbers i n Table A4. 2/ Nanaimo A i r p o r t (AES) 3/ Nanaimo Departure Bay (AES) 4/ A l b e r n i Lupsi Cupsi (AES) * imperfect drainage APPENDIX 3V (continued) Table A7. Environmental data f o r i n d i v i d u a l sample p l o t s (continued). AVERAGE WATER AVERAGE POT.ANNUAL CLIMATE STATIONS PLOT ELEV. MAY-SEP AWSC STRESS MAY-SEP SOLAR RADN. USED IN EXTRAPOL. NO. (m) PRECIP. (cm) INDEX TEMP(°C) 10 3cal/cm 2/yr P p t . d i s t . / s o l a r 1151 900 34.0 19.0 0.0+0.0 11.4 128 18 NA 1152 980 35.6 14.0 0.0+0.0* 10.9 88 18 NA 1153 1050 36.8 18.5 0.0+0.0* 10.5 137 18 NA 1154 660 • 29.5 10.5 0.0+2.0 11.7 220 19 NA 1155 740 31.0 15.0 0.0+0.0 11.3 193 19 NA 1156 900 34.0 7.0 0.0+4.0* 11.4 193 18 NA 1157 820 32.5 5.0 1.0+3.0 10.8 193 19 NA 1158 1060 36.8 9.5 0.0+1.0 10.5 193 18 NA 1159 980 35.6 12.0 0.0+0.0 10.9 193 18 NA 1160 580 27.9 8.5 0.0+3.0 12.1 220 19 NA 1161 440 25.1 11.0 0.0+2.0 12.4 170 16 NA 1162 1000 37.1 15.5 0.0+0.0* 10.1 137 11 ND 1163 1110 36.1 8.5 0.0+3.0 10.5 193 11 ND 1164 1045 37.6 11.5 0.0+0.0 9.5 201 10 AL 1165 1350 41.1 7.0 0.0+2.0* 7.8 165 10 AL 1166 1235 39.9 11.5 0.0+0.0 8.4 151 10 AL 1167 1140 38.9 9.0 0.0+0.0 9.0 105 10 AL 1168 1075 38.1 11.5 0.0+0.0* 9.3 170 10 AL 1169 1000 37.3 6.5 0.0+4.0* 9.7 170 10 AL 1170 1040 37.6 15.0 0.0+0.0* 9.5 151 10 AL 1171 830 35.3 10.0 0.0+0.0 10.6 151 10 AL 1172 800 35.1 6.0 0.0+4.0 10.8 170 ... io AL 1173 940 36.6 9.0 0.0+0.0 10.1 131 10 AL 1174 790 35.1 19.0 0.0+0.0 10.9 137 10 AL 1175 120 19.1 10.5 1.0+1.5 14.3 166 15 NA 1176 85 17.5 15.5 0.0+2.0 15.9 170 20 CL 1177 95 17.5 4.5 2.5+1.5 15.8 170 20 CL 1178 170 20.1 16 .'5 0.0+2.0 14.0 170 15 NA 1179 620 17.8 9.0 0.0+2.5 11.6 159 2 CR 1180 570 16.5 8.5 1.5+2.0 11.8 166 2 CR 1181 880 24.1 6.5 0.5+3.5 10.2 170 2 CR 1182 915 25.1 15.0 0.0+0.0 10.0 170 2 CR 1183 810 22.3 15.5 0.0+0.0 10.6 170 2 CR 1184 725 20.3 13.0 0.0+1.5 11.0 170 2 CR 1185 635 18.0 14.0 0.0+2.5 10.9 193 2 CR 1186 10 23.6 19.0 0.0+0.0 14 .7 170 5 AL 1187 25 23.9 9.5 0.0+2.0 15.7 181 5 AL 1188 40 24.1 19.0 0.0+0.0 15.6 185 5 AL 1189 30 23.1 25.5 0.0+0.0* 15.6 170 5 AL 1190 50 24.1 16.0 0.0+0.5 15.5 225 5 AL 1191 90 24.6 7.0 0.5+3.5 • 15.3 170 5 AL 1192 120 25.1 23.0 0.0+0.0* 15.1 170 5 AL 1193 120 25.1 18.5 0.0+0.0* 15.1 170 5 AL 1194 100 24.9 19.0 0.0+0.0* 15.2 193 5 AL 1195 50 24.1 18.5 0.0+0.0 15.5 170 5 AL 5/ Cowichan Lake Forestry (AES) 6/ Campbell River (AES) 215 APPENDIX IV (continued) Table A7. Environmental data f o r i n d i v i d u a l sample p l o t s (continued). AVERAGE WATER AVERAGE POT.ANNUAL CLIMATE STATIONS PLOT ELEV. MAY-SEP AWSC STRESS MAY-SEP SOLAR RADN. USED IN EXTRAPOL. NO. (m) PRECIP. (cm) INDEX TEMP(°C) 10 3cal/cm 2/yr P p t . d i s t . / s o l a r 1196 205 26.2 12.5 0.0+1.0 14.7 166 5 AL 1197 470 29.5 12.0 0.0+0.5 14.3 125 7 AL 1198 470 29.5 20.5 0.0+0.0 14.3 204 7 AL 1199 500 29.7 4.0 3.0+2.0 14.1 170 7 AL 1200 585 17.0 12.0 0.5+2.5 11.8 193 2 CR 1201 500 14.9 15.5 0.5+2.5 12.2 201 2 CR 1202 475 14.2 16.0 0.5+2.5 12.4 201 2 CR 1203 770 21.6 9.0 0.0+1.5* 10.8 128 2 CR 1204 700 19.6 16.0 0.0+0.0 11.2 125 2 CR APPENDIX V: S i t e c h a r a c t e r i s t i c s f o r i n d i v i d u a l sample p l o t s by habitat type. 216 Table A8. Explanation of terms and abbreviations used i n s i t e c h a r a c t e r i s t i c t a b l e s . SLOPE POS'N TOPOGRAPHY Macro slope position/slope configuration, shape — i r r = i r r e g u l a r ; sm = smooth; c = coastal; f l = f l o o r PARENT MATERIAL Coll u v i a l / m o r a i n a l = c o l l u v i a l over morainal deposits, g l = g l a c i a l SOIL TEXTURE C = clay; L = loam; S = sand; S i = s i l t ; g = g r a v e l l y STONINESS : EST. = f i e l d estimates i n four classes f o r a l l rock materials. Scale: 1 = l e s s than 20% 2 = 20 - 50% 3 = 50 - 90% 4 = greater than 90% %WGT = the range i n percent coarse fragment content (over 2mm) from laboratory a n a l y s i s of a l l mineral horizons; rock fragments above gravel size (7.5 cm) excluded. STAND AGE : Taken at breast height using an increment borer, or from stumps i n adjacent c l e a r c u t s . CANOPY DENSITY Measured with a mirror densiometer. HT. OF DOMS Height of the dominant tree canopy layer measured with a Suunto clinometer and tape. BASAL AREA Given i n square meters per hectare, based on p l o t mensuration data. 217 APPENDIX V Ccqntinued),. ft. X* Ul Id <. m «1 g Cu o • • 2-a. M O E/1 —• s — O Q 3 s 3 o wo O CJ O m > n w o u rH a> 4-1 rH O rH 01 OJ 0> CD §• -a o • H • H e 3 W «» H 2 U M 3 f< c « 1/1 rH fH in i SO ^ °-O U CM — a a • CN CN ro m m rH 3 , 5 a w 0- < in br. a 'o a; 4-) CL, O -H O a1 u rc • fl • I • • rT3 • Ui U V U w 14 M -1 V« e u U •H - H £ -H -H -H > rH rH 4J •U *W 4J fl <a u •Tl c >J (8 C o c: >• -H >, -H rH •a 0 0) *H 13 tn a (0 T) TJ o TJ •H S d •H c* rJ •H EH S 1/1 in fl E r4 in > u > a •H m o m m o in rH 3 o •3 E O o o o in o m <£) in \r fl fl fl rH •H -H -H fl fl > > > 3 3 > > rH 3 3 rH rH rH 0 0 0 rH rH U o U fl ta rH 4-J •p tl) u U CJ t4 fl fl rH 4J -H N rH XI tfi a 'JI T5 0) cu U-l CJ l« TJ E o u e e rH fi E tn u e fl cn fl Ul > > m m in o O r- r*> CO m O in m CO m 1 CN (N o O O o in ro >x> cn OJ O rH 1-4 Ta'ole A12. Abiz6 gKandii/Polyi.tichum minitum (ABGR/POMU) habitat type. - iitz ckaiacteAiitia. HUMUS SOIL TEXTURE STONINESS SOIL DRAINAGE CANOPY HT. OF BASAL ELEV. % SLOPE POS'N PARENT ROOTING DE: PTH UPPER LOWER % (VFnTTPar.l STAND DENSITY DOMS. AREA PLOT NO. ASPECT SLOPE TOPOGRAPHY MATERIAL DEPTH (cm) LP a 25cm +25cm EST ' WGT. AGE (m) m3 /'na 1315 275 180 12 lower a l l u v i a l 59 6 5 LS-L SiL 1/1 7-37 mod.well 340 89 45 125 irr.concave 1119 300 55 70 lower c o l l u v i a l 84 4 4 L L 3/4 53-68 w e l l 300 87 55 112 i r r . s t r t . 1176 85 - 0 va l l e y f l . a l l u v i a l 52 3 0 SiL-LS LS 1/1 0-1 mod.well 220 90 50 201 sm. f l a t 1186 10 - 0 val l e y f l . a l l u v i a l 80 4 0 CL-L LS 1/3 0-68 w e l l 200 97 50 155 i r r . f l a t J 187 25 120 30 lower a l l u v i a l 75 5 0 SiL LS. 1/3 61-74 mod.well 500 8S 50 208 irr.concave 1183 40 120 75 middle g l . - f l u v i a l 74 4 0 L-SL L 1/2 35-68 mod.well '50C 96 50 102 1139 30 - 0 val l e y f l . a l l u v i a l 63 4 0 SiL-CL CL-SCL C/2 9-55 imperfect 100 95 55 193 i r r . f l a t (+vets) 1190 50 190 65 middle morainal 100 7 0 L SL 2/3 52-88 w e l l 100 92 40 66 i r r . s t r t . 1194 100 130 17 middle morainal 69 . 4 0 SL SCL 2/2 23-39 mod.well 120 96 35 186 irr.concave .1195 50 - 0 va l l e y f l . a l l u v i a l 90 5 3 SCL SL-S 1/2 40-81 mod.we11 90 97 35 59 i r r . f l a t Tabic A l 3. Thuja, plicata/LtjiidUXwn amitAicanum [THPL/LVM\) habitat type. - iitt chaAacteAiitici. 1178 170 60 5 lower gl.marine 14 1 4 SiCL SCL-CL 1/2 11-39 imperfect 190 94 26 59 i r r . f l a t 1192 120 - 0 c. p l a i n gl.marine 28 2 24 SCL CL 2/3 10-43 poor 110 95 30 99 i r r . f l a t 1193 120 - 0 c. p l a i n i r r . f l a t gl.marine 45 4 11 . SCL CL 2/3 50-58 imperfect 150 81 NR 90 Table A14. T&w&ga hztwiphylixilGauJUhvuja. ihallon (TSHE/GASH) habitat typz - ilXz chaAact&U&ticA. ELEV. SLOPE POS'N PARENT PLOT NO. (n) ASPECT SLOPE TOPOGRAPHY MATERIAL 1113 640 335 40 middle colluvial irr.concave 1114 580 302 50 up.middle colluvial sm.convex 1123 550 360 10 apex colluvial/ i r r . f l a t morainal 1147 660 360 35 middle gl.fluvial irr.concave ii-:a 580 20 45 middle morainal irr.concave 11S4 660 200 65 middle colluvial sm.strt. 1155 7 40 215 50 middle colluvial sm.strt 1160 580 210 70 lower morainal sm.strt. 1161 440 - 0 valley f l . gl.fluvial i r r . f l a t ROOTING DEPTH (c 80 80 34 80 50 110 100 120 60 1180 1164 1135 1200 1201 1202 5?0 725 635 585 500 475 270 83 142 145 160 160 50 10 35 27 45 35 lower irr . s t r t . middle irr.concave middle irr.concave middle irr.concave middle irr.concave middle irr.concave colluvial/ morainal morainal morainal morainal morainal morainal 56 53 77 79 3 1 5 0 2 5 3 1 10 10 2 -5 -6 3 3 2 3 2 1 0 SOIL TEXTURE UPPER LOWER 25cm +25cm STONINESS E S T - WGT. SOIL DRAINAGE (VERTICAL) STAND AGE CANOPY DENSITY (%) HT. OF DO.MS. !m) BASAL AREA m'/ha LS-SL SL 2/3 38 well 285 88 25 64 L-SL L-SL 4/4 42-47 well 300 79 28 59 SiL SiL 2/3 20-48 well 500 80 NR 66 SL-L LS 2/3 5-45 mod.well 315 85 30 76 LS gL LS gSL 3/3 3/3 37-76 62-80 mod.well-imperfect well 360 250 87 87 45 25 71 54 gL gL 3/3 70-75 well 270 89 22 47 gL-SL LS 3/3 43-67 well 285 89 35 84 LS gS 1/2 9-59 well 300 76 43 116 iphylla (TSHE/GASH.-BENE/ACTR) habitat type. - iite. chaAact.e.'ii&tici. SL SL 2/2 ND well 240 92 NR 96 L L-CL 2/3 41-54 mod.we11 260 95 28 122 L L 2/2 44-58 well 265 90 37 54 L SCL 2/2 49-70 mod.we11 250 93 40 70 LS-L L SiL-SiCL SCL 2/2 2/2 48-63 44-57 mod.we11 mod.we11 260 265 92 84 45 35 54 134 iS D td 2 D H X o o 3 rt H-3 C fD M M KO Table A16. Tiuga heXeAophylta/Volyiticham mutitum (TSHE/POMU) habitat type - iiXe chamcXeAlitia. HUMUS SOIL TEXTURE STONINESS SOIL DRAINAGE CANOPY HT. OF BASAL ELEV. SLOPE POS'N PARENT ROOTING DEPTH UPPER LOWER • STAND DENSITY DOKS. AREA PLOT NO. (m) ASPECT SLOPE TOPOGRAPHY MATERIAL DEPTH (cm) LF H 25cm +25cm EST ' WGT. AGE <%> fa) m* /ha 1122 500 2.40 25 lower c o l l u v i a l / 74 3 2 L L 2/1 31-35 imperfect: 300 97 34 76 irr.convex morainal (vets) 1123 272 60 40 lower g l . f l u v i a l 100 4 1 SL SL- 3/4 47-54 mod.well 300 95 60 32 irr.concave SiL 1135 240 295 65 lower morainal 90 4 6 SI. SL 3/3 60-64 wel l 120 . 93 27 129 irr.convex j.137 680 45 65 middle c o l l u v i a l / 40 1 2 SiL CL 3/4 52-73 rood.well 300 88 40 100 sm.strt. morainal 1138 610 30 60 middle c o l l u v i a l / 90 1 4 SiL SiL 3/2 57-73 mod.well/ 250 95 45 126 i r r . s t r t . morainal imperfect 1.143 500 360 50 lower g l . f l u v i a l 120 10 - SL LS-SiL 2/4 49-63 imperfect 360 88 34 150 i r r . s t r t . Table A17. ChairaecypaA.L& nootkateniii/GaattheAAa ihation [CHNO/GASH] habitat type. - hite. chanacteAibtici 1118 650 135 10 apex c o l l u v i a l / 63 5 - L L 2/4 48-58 mod.well 245 79 20 59 i r r . s t r t morainal 1156 900 215 45 middle c o l l u v i a l 40 2 2 L L 3/3 71-78 poor 275 68 25 79 i r r . s t r t . morainal 1157 820 215 45 middle c o l l u v i a l 35 3 5 gL gL 4/4 73-79 w e l l 270 93 28 99 i r r . s t r t . 1158 106C 220 30 upper morainal 70 7 3 SiL CL 3/3 55-59 poor 280 67 20 76 irr.convex 1159 930 235 35 upper c o l l u v i a l 60 4 1 L-SCL L-SCL 3/3 39-60 mod.we11 350 90 NR 113 i r r . s t r t . Table A18. Abiiu amabiUi/Vac.<Unitim aUikainit-Vaccunium pativi&olium (ABAM/MALjI/APA) habitat type - iite chatiacteAt&tia. PLOT KO. 1130 1140 1149 1150 1181 1197 i l'J9 1203 1204 Irr.convex Table Ai9. Abiu amabUUI Ac'nlyi VUphijUa-IiaxoXla tAtioliata (ABAM/ACTR-TITR) habitat type - iitz chaMcXeAi6tici. HUMUS SOIL TEXTURE STONINESS SOIL DRAINAGE CANOPY HT. OF BASAL ELEV. t SLOPE POS'N PARENT ROOTING DEPTH UPPER LOWER EST. % (VERTICAL) STAND DENSITY DOMS. AREA <m) ASPECT SLOPE TOPOGRAPHY MATERIAL DEPTH (cm) LP H 25cm +25cn> WGT. AGE (%) On) m3 /na 760 270 25 upper colluvial 24 2 2 CL _ .3 59 well 270 82 25 61 irr.convex 860 60 50 upper colluvial/ 80 4 2 SiL SiL 3/4 41-61 well 290 84 40 95 irr.convex morainal 740 30 40 middle morainal 48 2 3 SL L NR 40-57 mod.wall 300 96 45 123 i r r . s t r t . 820 30 40 upper morainal 100 2 5 . ' SL L 2/2 16-47 imperfect/ 280 95 40 120 830 i r r . s t r t mod.we11 82 5 middle morainal 46 1 - SL-LS LS 1/3 26-60 well 250 93 24 113 irr.concave 470 10 25 middle morainal 54 2 8 SL SCL 2/2 42-59 mod.we11 275 96 40 37 irr.concave 500 7S 10 middle morainal 39 2 4 SL SCL 3/4 12-51 mod.well 130 94 40 87 i r r . s t r t . 770 340 25 middle morainal 38 4 9 sc sc 2/3 43-54 mod.well/ • 225 94 40 105 360 irr.concave imperfect 695 32 middle morainal 46 11 7 L SL 1/3 32-54 mod.wall NR 93 30 100 1117 560 45 50 middle irr.convex colluvial/ morainal 100 5 - SiL SiL 4/3 43-61 mod.well 340 94 37 B5 1132 10S5 235 80 upper irr.concave colluvial 110 3 2 L SL 3/4 69-74 well 300 85 60 111 1133 940 260 80 middle i r r . s t r t . colluvial 120 3 - L L 4 75 well 265 83 45 141 1139 780 50 ao upper colluvial 90 5 3 SCL SL 3/3 72-74 well 250 79 50 90 sm. strt. 1179 620 262 55 middle irr.concave colluvial/ morainal 70 6 - SL L-SL 4/4 65-73 well 280 94 47 111 W o H X o • rt K-3 C CD Table A20. Abiej, ambilU/Vaccuiium aixukaenit/StAeptopuA bpp. (ABAM/l/AAL/STREPTOPUS) habitat, type - bite. chaAacteAibticj,. HUMUS SOIL TEXTURE STONINESS SOIL DRAINAGE CANOPY HT. OF BASAL ELE1/. % SLOPE POS'N PARENT ROOTING DEPTH UPPER LOWER EST. % (VERTICAL) STAND DENSITY DOMS. AREA PLOT NO. in) ASPECT SLOPE TOPOGRAPHY .MATERIAL DEPTH (cm) LF H 25cm +25cm WGT. AGE (*) to) ra" Aia 1136 510 45 10 v a l l e y f l . g l . f l u v i a l 90 2 6 L L 3/4 26-72 mod.we11 300 89 32 91 i r r . f l a t 1170 1040 70 35 middle morainal 70 5 2 Si l . SCL 2/3 39-74 imperfect 260 95 45 119 i r r . s t r t . 1173 940 60 80 middle c o l l u v i a l 50 6 12 gL gSiL 4/4 69-80 w e l l 300 94 38 113 i r r . s t r t . 1174 790 45 45 middle morainal 60 5 15 CL CL 3 47 mod.well 300 89 45 91 i r r . s t r t . 1183 310 45 10 middle morainal 52 3 17 SL SL 1/3 33-60 mod.well 240 93 36 76 i r r . s t r t . Table A21. Abie-i arrabiZib/Vacc-inijum aiabkaeMerVaccAnUun ovaliioiiwn/Rubub pedatab 1 ABAM/l/AAL (Of) /RUPE) habitat type - bite chanacteAibticb. 1141 965 45 35 upper morainal 70 1 4 SiCL SiCL- 4/4 69-75 imperfect/ 280 93 37 124 i r r . s t r t . CL poor 1151 900 20 35 middle c o l l u v i a l / 90 2 10 L SL-L 1/3 18-38 mod.well 265 84 40 88 i r r . s t r t . morainal 1152 930 20 60 upper c o l l u v i a l 60 2 4 SiL SiL 3/3 41-61 poor 240 87 25 73 i r r . s t r t . 1169 1000 315 10 lower morainal 20 2 6 L-SCL - 2 20-51 imperfect 250 88 42 123 i r r . f l a t . 1132 915 75 10 upper morainal 38 4 - L-SCL L 2/2 11-59 mod.well 290 92 32 93 i r r . s t r t . Table A22. Kbieb amabitcb/Oplopanax hoVu.du.rn !ABAM/OPHO) habitat type - bite chaAacteAibtixu>. 1171 830 70 40 lower morainal 80 2 8 SCL SCL 3/3 60-65 w e l l 195 ?7 52 114 i r r . s t r t . 1172 800 - 0 vall e y f l . a l l u v i a l 65 4 2 SiCL S 2/4 51-93 ra p i d 165 95 51 117 IS H a o H o o 3 r+ r (D sm.flat 223 APPENDIX VI: Assoc i a t i o n t a b l e s f o r the habitat types. Table A23. Explanation of terms and abbreviations used i n the a s s o c i a t i o n t a b l e s . ST. NO. = Strata number (see canopy layers given i n Table A2). SYNTHETIC VALUES: P = Presence—percentage occurrence of a species i n the type. MS = Mean species s i g n i f i c a n c e — t h e average cover, computed using the midpoints of the cover classes below, and given as a cover scale value i n tenths, (e.g. 5.5. i s midway between the means of cover classes 5 and 6) RS = Range i n s i g n i f i c a n c e — u p p e r and lower l i m i t s of coverage of a species i n the type. COVER AND VIGOR: Cover Classes Vigor r a t i n g + - trace 0 = dead 1 = 0.1 - 1.0 % + = poor 2 = 1.1 - 2.2 1 - f a i r 3 = 2.3 - 5.0 2 = good 4 = 5.1 - 10.0 3 = e x c e l l e n t 5 = 10.1 - 25.0 6 = 25.1 - 33.0 7 = 33.1 - 50.0 8 = 50.1 - 75.0 9 - over 75.0 -A species with a value of 7.1 i s i n cover c l a s s 7, with a vigor r a t i n g of 1. -Note: no vigor ratings are given f o r tree species. VI (continued).. Table A 2 4 . Association table for the P S M E T A R K E / G A S H habitat type. P L O T 1 S Y N T H E T I C 1 1 1 1 1 I N U M B E R 1 V A L U E S J)1 24 |11 1G |11 10 | i 1 0 9 (1« 7 5 J S T . N O . S P E C I E S j P MS R S i C O V E R AND V I G O R A3 B 1 82 C 1 P S E U D O T S U G A M E N Z I E S I I 1 0 O . 0 7 7 5 - 8 | 8 8 7 7 s I P S E U D O T S U G A M E N Z I E S I I 4 0 0 2 4 0 - 4 4 4 2 A R B U T U S M E N Z I E S I I 2 0 0 2 4 0 - 4 4 3 P I N U S C O N T O R T * 2 0 0 2 4 0 - 4 4 P S E U D O T S U G A M E N Z I E S I I 1 0 0 0 4 1 •»-5 3 3 5 + 3 A R B U T U S M E N Z I E S I I 6 0 0 7 0 0 - S 7 8 a P I N U S C O N T O R T A 2 0 0 1 5 0 - 3 3 4 H O L O D I S C U S D I S C O L O R 2 0 0 + 2 0 - 1 1 2 5 L O N I C E R A H I S P I O U L A 2 0 0 4 2 0 - 1 1 2 6 R O S A G Y M N O C A R P A 2 0 0 2 0 - 1 1 1 7 G A U L T H E R I A S H A L L O N 1 0 0 0 7 1 1 - 8 7 1 8 1 8 2 4 2 1 2 8 B E R B E R I S N E R V O S A 1 0 0 0 3 1 1 - 3 1 1 2 1 3 1 3 2 3 2 R O S A G Y M N O C A R P A 8 0 0 4 1 0 - 5 2 2 5 2 4 1 2 1 H O L O D I S C U S D I S C O L O R 8 0 0 2 6 0 - 3 2 2 3 2 + 4^  3 2 9 S Y M P H O R I C A R P O S A L B U S 6 0 0 1 9 0 - 3 1 2 1 1 3 1 1 0 L O N I C E R A C I L I O S A 4 0 0 1 7 0 - 3 1 2 3 2 1 1 S Y M P H O R I C A R P O S M O L L I S 4 0 0 1 S 0 - 3 3 1 4- 1 12 A R C T O S T A P H Y L O S C O L U M B I A N A L O N I C E R A H I S P I O U L A 4 0 2 0 0 0 1 1 2 5 0 - 2 0 - 3 1 1 2 4-3 2 13 R U B U S U R S I N U S 2 0 0 1 0 0 - 2 2 1 14 A M E L A N C H I E R A L N I F O L I A 2 0 0 + 2 0 - 1 1 1 15 B E R B E R I S A Q U I F O L I U M 2 0 - 0 2 0 - 1 1 2 1S M E N Z I E S I A F E R R U G I N E A 2 0 0 2 0 - 1 1 1 17 C Y T I S U S S C O P A R I U S 2 0 0 + 0 0 - * •4 + 18 F E S T U C A O C C I O F . N T A L I S 1 0 0 0 3 3 4--4 3 2 4 2 3 1 1 2 + 2 19 G O O D Y E R A O B L O N G I F O L I A 1 0 0 0 1 1 4-- 1 1 1 + + 1 2 4- 1 + + 2 0 B R O M U S V U L G A R I S 8 0 0 5 6 0 - 9 1 2 1 2 4 2 9 2 R U B U S U R S I N U S 8 0 0 2 0 0 - 3 1 + 3 4 4- 4 1 1 21 T R I E N T A L I S L A T I F O L I A 8 0 0 2 0 0 - 3 1 1 1 1 3 1 4- 1 2 2 A R E N A R I A M A C R O P H Y L L A 6 0 0 2 7 0 - 4 2 2 4 2 1 2 2 3 M A D I A S A T I V A 6 0 0 2 1 0 - 3 2 2 3 2 1 2 2 4 C O L L O M I A H E T E R O P H Y L L A GO 0 1 4 0 - 2 2 2 2 2 4 1 2 5 H I E R A C I U M A L B I F L O R U M 6 0 0 1 4 0 - 2 2 2 1 1 1 2 2 6 C A M P A N U L A S C O U L E R I GO 0 1 0 0 - 1 1 2 4 1 1 1 2 7 P O L Y S T I C H U M M U N I T U M GO 0 1 0 0 - 1 4 + 1 2 1 2 2 8 C H I M A P H I L A U M B E L L A T A 4 0 0 2 6 0 - 4 4 2 2 2 2 9 D A N T H O N I A S P I C A T A 4 0 0 2 5 0 - 4 4 2 1 2 3 0 L A T H Y R U S N E V A D E N S I S 4 0 0 2 3 0 - 3 3 2 3 2 31 A D E N O C A U L O N B I C O I . O R 4 0 0 1 6 0 - 3 3 1 4- 1 3 2 M O N T I A P E R F O L I A T A 4 0 0 1 0 0 - 2 4-+ 2 + 3 3 F R A G A R I A V E S C A 4 0 0 + 8 O - 1 1 1 1 2 3 4 P O L Y S T I C H U M L O N C H I T I S 4 0 0 + 4 0 - 1 4- 1 1 2 3 5 E L Y M U S G L A U C U S 4 0 0 4 0 0 - + 1 + 2 3 6 L A C T U C A M U R A L I S 4 0 0 + 0 0 - + + 1 4 1 3 7 T H A L I C T R U M O C C I D E N T A L S 4 0 0 * 0 0 - + + 4- + 3 8 E L Y M U S S P P 2 0 0 3 4 0 - 5 5 2 3 9 B R O M U S S P P B E R B E R I S A Q U I F O L I U M 2 0 2 0 0 0 2 + 4 2 0 - 4 0 - 1 1 2 4 3 4 0 D E S C H A M P S I A E L O N G A T A 2 0 0 4 2 0 - 1 1 2 4 1 G A L I U M T R I F L O R U M 2 0 0 + 2 0 - 1 1 2 4 2 G E R A N I U M S P P 2 0 0 2 0 - 1 1 2 4 3 H Y P O C H A E R I S R A D I C A T A 2 0 0 + 2 0 - 1 1 2 4 4 U U N C U S S P P 2 0 0 + 2 0 - 1 1 2 4 5 L I N N A E A B O R E A L I S 5 0 0 4- 2 0 - 1 f 2 4 6 V I C I A A M E R I C A N A 2 0 0 4 2 0 - 1 1 2 4 7 V I C I A S A T I V A 2 0 0 4- 2 0 - 1 1 1 4 8 A C H I L L E A M I L L E F O L I U M 2 0 0 + 0 0 - * 4 + 4 9 A G R O S T I S T E N U I S A M E L A N C H I E R A L N I F O L I A 2 0 2 0 0 0 4-4-0 0 0 - * 0 - 4 + 1 4 5 0 A N E M O N E L Y A L I I 2 0 0 4 0 0 - * 4 4 5 1 H Y P O P I T Y S M O N O T R O P A 2 0 0 4 0 0-4 + 2 5 2 L U Z U L A S P P 2 0 0 4-0 0 - + + 1 5 3 M E L I C A S U B U L A T A 2 0 0 4 0 0 - + + 2 5 4 P O L Y P O D I U M G L Y C Y R R H I Z A 2 0 0 4 0 0 - 4 4 4 5 5 P T E P O S P O R A A N D R O M E D I A 2 0 0 4 0 0 - * 4-5 6 R H A M N U S P U R S H I A N A 2 0 0 + 0 0-4 4- 1 APPENDIX VI (continued) T a b l e A2 5. A s s o c i a t i o n t a b l e f o r the PSME/GASH-BENE h a b i t a t t ype. PLOT 1 SYNTHETIC 1 1 I 1 1 1 1 NUMBER | VALUES |1l 12 |(12 1 |I125|119 1 |i1 1 1 |f 136 | ST.NO. SPECIES | P MS RS | COVER AND VIGOR A 1 A2 A3 C 1 PSEUDOTSUGA MENZIESII 100 .0 6 . 3 5-7 5 7 . 7 . 5. 5 . 6 2 THUJA PLICATA 66 . 7 5 . 2 0-7 4 . 5. 7 . 5 3 TSUGA HETEROPHYLLA 50 .0 3 . 3 0-4 3 . 4 . 4 4 ABIES GRAND IS 16 . 7 2 . 2 0-4 4 . 5 PINUS CONTORTA 1G . 7 f- . 7 0-2 2 TSUGA HETEROPHYLLA 66 . 7 5 . 2 0-7 3. 1 7 . 5 . 5 THUJA PLICATA 66 . 7 4 .5 0-6 6 . 3 . 4 . 4 PSEUDOTSUGA MENZIESII 50 .0 3 .6 0-5 5 3 . 3 . ABIES GRAND IS 16 . 7 3 . 2 0-5 5 . e CORNUS NUTTALL1I 16 . 7 3 .2 0 -5 5. 7 ALNUS RUBRA 16 7 1 . 4 0-3 3 . TSUGA HETEROPHYLLA 83 3 5 .0 0-G 3 . 3 . 4 . 5 . 6 PSEUDOTSUGA MENZIESII 66 7 3 . 5 0-5 5 3 . 2. 1 . ABIES GRAND IS 50 0 4 . 5 0-7 1 7 . 3. THUJA PL I CATA 50 0 3 . 1 0-4 + . 4 . 4 8 ACER MACROPHYLLUM CORNUS NUTTALL11 33 16 3 7 1 3 . 7 .2 0-3 0-5 3. 5 . 2 . 9 TAXUS BREVIFOLIA ALNUS RUBRA 16 16 7 7 3 + 2 . 7 0 -5 0-2 5. 2. 10 VACCINIUM PARVIFOLIUM 50 0 2 5 0-4 4.2 1 . 3 1 . 2 2| 1 1 HOLODISCUS DISCOLOR 16 7 4-0 O- 1 1 . 2 • I 12 GAULTHERI A SHALLON 100 0 7 4 5-9 5 1 8.2 9 . 3 5 . 3 5 . 1 7 2 13 BERBER IS NERVOSA 100 0 5 3 2-7 3 + 7.2 5 . 3 4 . 3 2 . 1 5 2 VACCINIUM PARVIFOLIUM 10O 0 3 3 1-4 1 + 1 . 2 1 . 2 4 . 3 2 . 1 4 2 14 RUBUS URSINUS 50 0 1 1 0-2 2 . 1 1 . 1 * . 1 15 ROSA GYMNOCARPA HOLODISCUS DISCOLOR 33 33 3 3 3 1 2 0 0-5 0-2 1 + 5.2 2 . 2 1 . 2 16 LONICERA SPP 16 7 + 0 0 - + + + 17 SYMPHORICARPOS ALBUS 16 7 + 0 0 - + * + 18 TRIENTALIS LATIFOLIA 100 0 4 0 + -6 1 1 6 . 2 1 . 1 * . 1 2 . 2 1 2 19 POLYSTICHUM MUNITUM 83 3 3 5 0-5 1 . 2 3 . 2 + .2 5.2 t 1 20 PTERIDIUM AOUILINUM 83 3 3 1 0-4 2 1 2 . 2 4 . 2 2 . 1 3 2 2 1 ACHLYS TRIPHYLLA 66 7 4 7 0-7 7 . 2 4 . 3 2 . 2 1 2 22 LINNAEA BOREALIS 50 0 1 9 0-3 2 1 1 . 2 3 2 23 FESTUCA OCCIDENTALIS 33 3 + 6 0-1 1 1 1 . 2 24 PACHYSTI MA MYRSINITES 33 3 •4 2 0 - 1 1 + 4- . + VACCINIUM PARVIFOLIUM 33 '3 4- 2 0-1 1 . 3 + . 2 25 ARCTOSTAPHYLOS UVA-URSI 16 7 1 4 0-3 3 1 26 GALIUM SPP 16 7. 1 4 0-3 3 . 2 27 ADIANTUM PEDATUM 16 7 + 0 0 - 1 1 . 2 28 AGROPYRON SPP 16 7 + 0 0 - 1 1 . 1 29 ANEMONE LYALI I '.6 7 + 0 0-1 1 . 2 30 DE SCHAMPSI A ELONGATA 16 7 + 0 0 - 1 1 . 1 3 1 TIARELLA TRIFOLIATA 16 7 + 0 0 - 1 1 . 2 32 BOSCHNIAKI A HOOKERI 16 7 + 0 0 - * 4- . 2 33 COPTIS SPP 16 . 7 + 0 0--"- + . 1 34 HYPOCHAERIS RAOICATA 16 . 7 4-0 0 - + t- . 1 35 LACTLICA MURALIS 1G. 7 4-0 0 - * + . 1 36 LIST ERA CORDATA 16 . 7 + 0 0 - + + . 2 37 . POLYPODIUM GLYCYRRHIZA RUBUS URSINUS 16 . 16 . 7 7 4 + 0 0 0 - + 0 - + 4- . 1 38 TRILLIUM OVATUM 16 . 7 + 0 0 - + + . 2 39 VIOLA ORBICULATA 16 . 7 4-0 0 - * + . + 40 VIOLA SPP 16 . 7 + 0 0 - * 4- . 1 APPENDIX VI (continuedL T a b l e A26. A s s o c i a t i o n t a b l e f o r the PSME/HODI/POMU h a b i t a t t y p e . PLOT NUMBER 1 SYNTHETIC | VALUES ||127|l12o|t12e|l145|l177| ST NO. SPECIES | P MS RS | COVER AND VIGOR A1 A2 C 1 PSEUDOTSUGA MENZIESII 100. 0 8 . 7 7-9 9 . 7 . 9. 8 . 8. 2 THUJA PLICATA 20. 0 2 . 4 0-4 4 . 3 TSUGA HETEROPHYLLA 20. 0 2 . 4 0-4 4 . PSEUDOTSUGA MENZIESII 80. 0 4 . 4 0-5 3 . 4 . 4 . 5. TSUGA HETEROPHYLLA 40. 0 4 . 0 0-5 5 . 4 . 4 ACER MACROPHYLLUM 20. 0 4 . 0 0- + . PSEUDOTSUGA MENZIESII 80. 0 3 . 4 0-4 3 . 3. 3 . 4 . TSUGA HETEROPHYLLA GO. 0 4 . 3 0-5 5 4 . 4 . ACER MACROPHYLLUM 60. 0 3 . 0 0-4 4 . 4 3 . THUJA PL I CATA 60. 0 1 . 8 0-3 1 4 . 3 . 5 ABIES GRANDI5 20. 0 1 5 0-3 3 . 6 HOLODISCUS DISCOLOR I 80 0 4 0 0-4| 4 2| 4 3| 3 . 2|4 2 |7 .3 | 7 ROSA GYMNOCARPA 80 0 4 2 0-5 3 2 5 2 3 2 4 . 1 8 VACCINIUM PARVIFOLIUM 80 0 3 8 0-5 3 2 1 . 1 2 2 5 2 9 BERBERIS NERVOSA 80 0 3 3 0-4 4 2 3 . 1 2 2 3. 1 10 GAULTHERI A SHALLON 40 0 5 5 0-9 1 4 9 2 1 1 LONICERA UTAHENSIS 40 0 1 6 0-3 + 1 3 1 12 RUBUS SPECTABILIS HOLODISCUS DISCOLOR 40 20 0 0 4 8 6 0-1 0-7 1 . 1 1 1 13 RUBUS URSINUS 20 0 1 5 0-3 3 1 14 SYMPHORICARPOS ALBUS 20 0 1 5 0-3 3 3 15 RIBES SPP 20 0 4 2 0- 1 1 1 1G AMELANCHIER ALNIFOLIA 20 0 4 0 0- + + + 17 POLYSTICHUM MUNITUM 100 0 8 2 3-9 7 1 9 3 8 2 9 3 3 1 18 ACHLYS TR I PHYL.LA 100 0 3 7 3-4 3 2 4 2 3 2 3 3 3 2 19 TRIENTALIS LATIFOLIA 80 0 3 3 0-4 3 2 2 2 3 2 4 2-20 GALIUM TRIFLORUM 80 0 2 3 0-3 3 2 2 2 1 1 2 1 21 POLYPODIUM GLYCYRRHIZA 80 0 1 0 0-1 + 1 + 4 1 1 1 2 22 LACTUCA MURAL IS 60 0 3 1 0-4 4 1 2 2 3 2 23 LINNAEA BOREALIS 60 0 2 5 0-3 1 3 2 3 3 24 TIARELLA l.ACINATA 60 0 2 1 0-3 1 2 2 2 3 3 25 TIARELLA TRIFOLIATA 60 0 2 1 0-3 1 2 2 2 3 3 2G ARENARI A MACROPHYLLA 60 0 1 6 0-2 1 1 2 2 2 1 27 BROMUS INERMIS 40 0 2 3 0-3 3 2 3 3 28 ADENOCAULON BICOLOR 40 0 1 7 0-3 3 2 1 2 29 LATHYRUS NEVADENSIS 40 0 1 4 0-2 2 2 2 1 30 CHIMAPHILA UMBELLATA 40 0 + 8 0- 1 1 1 1 2 31 CAMPANULA SCOULERI 40 0 + 4 0- l' + 1 1 1 32 MONTIA SIBIRICA 40 0 + 4 0- 1 4 2 1 2 33 VIOLA GLABELLA 40 0 + 4 0- 1 4 1 1 3 34 VIOLA ORBICULATA 40 0 + 4 0- 1 + . 1 1 2 35 LISTERA CORDATA 40 0 4 0 o-<- 4 . 4 4 1 3G BROMUS SPP 20 0 1 .5 0-3 3 3 37 FRAGARIA VESCA 20 .0 1 .5 0-3 3 2 38 MEL ICA SUBULATA 20 .0 1 .5 0-3 3 2 39 PTERIDIUM AOUILINUM 20 .0 1 . 5 0-3 3 . 3 40 TRAUTVETTERIA CAROL INIENS IS 20 .0 1 . 5 0-3 3 . 2 4 1 ATHYRIUM FILIX-FEMINA 20 .0 1 .0 0-2 2 . 2 42 CAREX SPP 20 .0 t .0 0-2 2 . 2 43 MONTIA PARVIFOLIA 20 .0 1 . 0 C - 2 . 1 44 STACHYS COOLEYAE 20 .0 1 .0 0-2 2 . 2 45 ACTAEA RUBRA 20 .0 4 . 2 0- 1 1 . 2 4G ADIANTUM PEDATUM 20 .0 + . 2 0- 1 1 . 3 47 CHIMAPHILA MENZIESII 20 .0 + . 2 0- 1 1 . 1 48 LILIUM COLUMBIANUM 20 .0 + . 2 0-1 1 .2 49 POLYSTICHUM LOMCHITIS 20 .0 + . 2 0- 1 1 . 1 50 RANUNCULUS UNCINATUS 20 .0 + . 2 0- 1 1 . 2 51 TRILLIUM OVATUM 20 .0 + . 2 0- 1 1 . 2 52 GOODYERA OBLONGIFOLI A RUBUS URSINUS 20 20 .0 .0 + + .0 .0 0- + o-+ 4 4 . 1 . 1 APPENDIX VI (continued) T a b l e A 2 7. A s s o c i a t i o n c a b l e f o r t h e ABGR/POMU h a b i t a t t y p e . P L O T 1 S Y N T H E T I C 1 j I 1 I 1 j 1 | 1 I N U M B E R 1 V A L U E S | M 9 0 | l l 1 9 | | 1 7 6 |1195|11 15 | l ! 9 4 | <188 | l 187 J1! BO (1186 j S T . N O . S P E C I E S | P MS R S | COV£R A NO V I G O R A2 A 3 C 1 P S E U D O T S U G A M E N Z I E S I I 8 0 . 0 5 . 9 0 - 7 7 6 7 6 7 5 . 7 4 2 T H U J A P L I C A T A 5 0 . 0 4 . 5 0 - 5 5 5 5 5 3 A B I E S G R A N D I S 4 0 . 0 4 . 4 0 - 7 4 3 7 5 4 A C E R M A C R O P H Y L L U M 3 0 . 0 4 . 8 0 - 7 7 . 4 7 5 T S U G A H E T E R O P H Y L L A 3 0 . 0 4 . 4 0 - 8 8 1 4 6 P O P U L U S T R I C H O C A R P A 1 0 . 0 1 . 0 0 - 3 3 T S U G A H E T E R O P H Y L L A 6 0 . 0 4 . 6 0 - 5 5 4 5 . 3 5 A C E R M A C R O P H Y L L U M 5 0 . 0 4 . 4 0 - 6 5 3 6 . 3 5 T H U J A P L I C A T A 4 0 . 0 3 . 0 0 - 4 3 4 4 3 A B I E S G R A N D I S 4 0 . 0 2 . 4 0 - 4 3 3 4 2 P S E U D O T S U G A M E N Z I E S I I 3 0 . 0 3 . 5 0 - 5 5 5 3 7 A L N U S R U B R A 20 . . 0 1 .5 0 - 3 3 3 T S U G A H E T E R O P H Y L L A 1 0 0 . 0 5 . 6 1-7 2 5 4 1 5 6 5 . 7 5 7 A B I E S G R A N D I S 7 0 . 0 3 . 2 0 - 4 2 2 3 3 3 4 . 4 T H U J A P L I C A T A 6 0 . 0 2 . 4 0 - 3 3 3 1 1 3 3 A C E R M A C R O P H Y L L U M 4 0 . 0 3 . 6 0 - 5 5 2 5 3 P S E U D O T S U G A M E N Z I E S I I 3 0 . 0 1 . 7 0-3 2 3 3 8 C O R N U S N U T T A L L I I A L N U S R U B R A 1 0 . 0 1 0 . 0 4 .2 0 - 2 + . 0 0 - 1 2 9 T A X U S B R E V I F O L I A 1 0 . 0 + . 0 0-* 4 1 0 V A C C I N I U M P A R V I F O L I U M I 3 0 . 0 1 . 2 0 - 3 1 | 1 2 | 1 3 i 3 3 1 I . 1 1 I 1 1 R U B U S S P E C T A B I L I S • 1 0 . 0 + . 0 ^ - 1 ! | l 3 I • ! 1 12 B E R B E R I S N E R V O S A 9 0 . 0 5 . 0 0 - 7 5 3 7 3 4 2 3 2 4 1 5 3 3 .2 3 3 3 2 V A C C I N I U M P A R V I F O L I U M 9 0 . 0 3 . 0 0 - 5 1 2 1 2 2 3 5 3 1 4 1 2 1 . 2 4 1 4 2 13 R U B U S U R S I N U S R U B U S S P E C T A B I L I S 6 0 . 0 6 0 . 0 1 . 4 0-3 4 .5 0 - 1 1 4 2 2 1 4 4 1 3 1 2 2 4 2 4 . 2 1 2 4 2 1 1 2 3 14 G A U L T H E R I A S H A L L O N 4 0 . 0 1 . 3 0 - 3 4 2 1 4 3 2 1 1 15 R O S A G Y M N O C A R P A 2 0 . 0 4.1 0 - 1 1 4 1 4 16 O P L O P A N A X H O R R I D U M 1 0 . 0 1.0 0 - 3 3 2 17 S Y M P H O R I C A R P O S A L B U S 1 0 . 0 + . 2 0 - 2 2 3 18 H O L O D I S C U S D I S C O L O R 1 0 . 0 + . 0 0-4 4 1 19 R I B E S L A C U S T R E 1 0 . 0 * . 0 0-4 4 2 2 0 P O L Y S T I C H U M M U N I T U M 1 0 0 . 0 8 . 2 5 - 9 9 3 7 3 9 3 8 3 5 2 8 3 9 . 3 7 3 7 3 7 3 21 T I A R E L L A T R I F O L I A T A 9 0 . 0 3 . 7 0-5 1 1 4 2 1 2 3 1 3 2. 1 . 2 4 3 1 2 5 3 2 2 A C H L Y S T R I P H Y L L A 9 0 . 0 3 . 0 0 - 4 4 2 3 2 1 2 1 2 3 2 3 . 2 4 2 2 2 2 3 2 3 T R J E N T A L I S L A T I F O L I A 8 0 . 0 2 .1 0-3 4 2 4 1 3 2 4 1 4 4 3 . 2 1 2 3 3 2 4 G A L I U M T R I F L O R U M 8 0 . 0 2 . 1 0 - 4 4 2 1 3 4 2 1 2 1 . 2 4 3 1 2 1 3 2 5 T R I L L I U M O V A T U M 8 0 . 0 1 . 1 0 - 2 4 I 4 2 4 1 1 1 4 2 2 3 4 2 1 2 2 6 T I A R E L L A L A C I N A T A 7 0 . 0 1 . 3 0 - 3 4 1 4 2 3 1 4 2 1 3 4 2 1 3 2 7 A D I A N T U M P E D A T U M 7 0 . 0 1 . 1 0 - 2 4 2 4 2 1 2 1 2 2 . 2 4 2 1 3 2 8 A D E N O C A U L O N B I C O L O R 6 0 . 0 1 . 2 0 - 2 1 2 1 2 1 2 2 3 4 1 1 3 2 9 L A C T U C A M U R A L I S 6 O . 0 1 . 2 0 - 2 1 3 1 2 2 1 1 . 3 4 2 1 3 3 0 D R Y O P T E R I S A U S T R I A C A 6 0 . 0 1 . 0 0 - 2 4 1 2 4 4 1 4 . 2 1 2 1 2 31 A T H Y R I U M F I L I X - F E M I N A 5 0 . 0 + . 9 0 - 1 1 1 1 2 1 . 2 1 2 4 2 3 2 P T E R I D I U M A O U I L I N U M 5 0 . 0 +'. 4 0 - 1 4 1 1 1 1 2 4 1 4 4 3 3 B R O M U S P A C I F I C U S 4 0 . 0 2 . 4 0 - 4 4 . 1 4 3 1 2 4 3 3 4 S T R E P T O P U S A M P L E X I F O L I U S 4 0 . 0 * . 8 0 - 2 1 2 4 1 4 2 2 3 3 5 C A R E X S P P 4 0 . 0 • 1 0 - 1 4 2 1 .1 4 1 4 1 3 6 P O L Y S T I C H U M L O N C H I T I S R U B U S U R S I N U S 4 0 . 0 3 0 . 0 4.1 0 - 1 • 4 0 - 2 * 4 2 1 2 . 2 4 4 . 1 4 4 1 1 3 7 P O L Y P O O I U M G L Y C Y R R H I Z A 3 0 . 0 4 .0 0 - 1 4 2 4 . 2 1 . 3 3 8 V E R A T R U M V I R I D E 3 0 . 0 4 .0 o- + 4 2 4 2 4 2 3 9 G A L I U M S P P 2 0 . 0 1 . 1 0 - 3 4 . 4 3 . 1 d O T R A U T V E T T E R I A C A R O L I N I E N S I S 2 0 . 0 1 . 1 0 - 3 4 . 1 3 2 4 1 B R O M U S V U L G A R I S 2 0 . 0 4.1 0 - 1 1 2 1 2 4 2 F E S T U C A O C C I D E N T A L I S 2 0 . 0 • . 1 0 - 1 1 2 1 2 4 3 G Y M N O C A R P I U M D R Y O P T E R I S 2 0 . 0 4.1 0 - 1 1 . 2 1 . 1 4 4 E O U I S E T U M S P P 2 0 . 0 » .0 0 - 1 1 . 2 4 . 2 4 5 M O N T I A S I 3 I R I C A 2 0 . 0 + . 0 0 - 1 4 . 2 1 . 3 4 6 S T A C H Y S C O O L E Y A E 2 0 * . 0 0-1 4 2 1 . 3 4 7 L I N N A E A B O R E A L I S V A C C I N I U M P A R V I F O L I U M 2 0 . 0 2 0 . 0 * . 0 0 - * 4 O 0 - * 4 1 4 . 2 4 . 3 4 2 4 8 L A T H Y R U S N E V A D E N S I S 1 0 . 0 • . 2 0 - 2 2 2 4 9 MA I ANT HE MUM D l L A T A T U M 1 0 . 0 4 .2 0 - 2 2 . 3 5 0 B L E C H N U M S P I C A N T 1 0 . 0 + . 0 0 - 1 1 . 2 5 1 B R O M U S S P P 1 0 . 0 4 .0 0 - 1 1 . 2 5 2 C A L Y P S O 3 U L B 0 S A 1 0 . 0 * . 0 0 - 1 1 . 2 53 E L Y M U S G L A U C U S 1 0 . 0 » .0 0 - 1 1 . 3 5 4 H E U C H E R A M I C R A N T H A 1 0 . 0 4 .0 0 - 1 1 . 3 5 5 L Y S I C H I T U M A M E R I C A N U M 1 0 . 0 * . 0 0 - 1 1 . 2 5 6 O S M O R H I Z A C H I L E N S I S R O S A G Y M N O C A R P A 1 0 . 0 1 0 . 0 * . 0 0 - 1 * . 0 0-1 1 . 2 1 . 3 5 7 T H A L I C T R U M O C C I D E N T A L S 1 0 . 0 + . O 0 - 1 1 . 3 5 8 V I O L A G L A B E L L A - 1 0 . 0 4 .0 0 - 1 1 . 3 5 9 C L A P O T H A M N U S P Y R O L A E F L O R U S 1 0 . 0 4 .0 0-4 4 , 46 0 L U Z U L A S P P 1 0 . 0 + . 0 0 - * 4 2 6 1 V I O L A 0 P B 1 C U L A T A 1 0 . 0 4 . 0 0-4 4 4 j APPENDIX VX. ^ continued.!. Table A2 8. A s s o c i a t i o n table for the THPL/LYAM ha b i t a t type. PLOT SYNTHETIC NUMBER VALUES 1192 (193 1178 ST.NO. SPECIES | P MS RS |COVER AND VIGOR A1 C 1 TSUGA HETEROPHYLtA 100 0 5 7 3-6 3. 6 . 6 . 2 THUJA PLICATA 66 7 5 3 0 -6 5. 6 . 3 POPULUS TRI CHOCARPA 66 7 5 1 0 -5 5 . 5 . 4 PSEUDOTSUGA MENZIESI I 66 7 5 1 0-5 5. 5 . 5 PICEA SITCHENSIS 66 7 4 0 0-4 4 . 4 . 6 ALNUS RUBRA 33 3 4 1 0 -5 5. ALNUS RUBRA 100 0 4 9 3-5 3 . 4 . 5 . TSUGA HETEROPHYLLA 100 0 3 7 2-4 2 . 3 . 4 . THUJA PLICATA 66 7 4 0 0-4 4 . 4 . 7 ACER MACROPHYLLUM 33 3 2 1 0 -3 3 . PSEUDOTSUGA MENZIESI I 33 3 2 1 0 -3 3 . PICEA SITCHENSIS 33 3 1 2 0-2 2. TSUGA HETEROPHYLLA 133 3 5 2 1-5 1 . 3 5 . THUJA PLICATA 66 7 5 3 0 -7 7 . 3 . ALNUS RUBRA 66 7 2 1 0 -3 + 3 . PICEA SITCHENSIS 66 7 2 0 0-2 2 . 2 PSEUDOTSUGA MENZIESI I 33 3 2 1 0 -3 3 . ACER MACROPHYLLUM 33 3 + 6 0 - 1 1 8 RUBUS SPECTABIL IS 33 3 + 6 0-1 • 1 2| 9 VACCINIUM PARVIFOLIUM 33 3 + 6 0-1 • 1 3 VACCINIUM PARVIFOLIUM 100 0 3 0 + -3 + . 1 3 3 3 3 RUBUS SPECTABIL IS 66 7 3 5 0-4 4 3 3 2 10 RHAMNUS PURSHIANA 66 7 1 0 0 - 1 + 2 1 1 1 1 GAULTHERIA SHALLON 33 3 4 1 0 -5 5 1 12 OPLOPANAX HORRIDUM 33 3 + 6 0-1 1 1 13 RIBES SPP 33 3 + 6 0-1 1 1 14 RUBUS URSINUS 33 3 4- 6 0-1 1 3 15 SAMBUCUS RACEMOSA 33 3 + 0 o-+ + 2 16 POLYSTICHUM MUNITUM 100 o 4 5 2-5 2.3 5 3 3 2 17 ATHYRIUM F IL IX -FEMINA 100 0 4 3 1 -5 1 . 3 5 3 2 1 18 LYSICHITUM AMERICANUM 100 0 3 3 + -4 2 . 3 + 2 4 2 19 DRYOPTERIS AUSTRIACA 100 0 1 7 + -2 + . 1 1 2 2 2 20 T IARELLA LACINATA 100 0 1 0 + - 1 + . 3 + 2 1 + 21 BLECHNUM SPICANT 66 7 3 1 0-4 + . 2 4 2 22 EOUISETUM SPP 66 7 1 .3 0-2 + . 1 2 2 23 VERATRUM VIRIDE 66 7 1 . 2 o - i 1 3 1 2 24 LACTUCA MURALIS 66 7 1 .0 0 - 1 + 2 1 1 25 POLYSTICHUM LONCHITIS . 66 . 7 1 .o 0 - 1 + 1 1 1 26 STREPTOPUS AMPLEXIFOLIUS 66 . 7 1 .0 0 - 1 + . 1 1 1 27 TIARELLA TRIFOLI ATA 66 7 1 .0 0-1 + 2 1 1 28 PT ERIDIUM AOUILINUM 33 . 3 2 . 1 0 -3 3 3 29 ACHLYS TRIPHYLLA 33 .3 + .6 0-1 1 3 30 GALIUM TRIFLORUM 33 . 3 +• .6 0 - 1 1 2 RUBUS URSINUS 33 . 3 4 .6 O- 1 1 1 3.1 STACHYS COOLEYAE .33 .3 +• .6 o-1 1 2 32 TRILLIUM OVATUM 33 . 3 + .0 o-+ +• .2 APPENDIX VI (continued). Table A2 9. Association table 'for the TSHE/GASH habitat type. PLOT 1 SYNTHETIC 1 1 1 I 1 1 1 ! 1 1 NUMBER 1 VALUES J11 13|I1 14 | i 1 5 5 | M 5 4 |<14 7 |(14 8 |l160|l16 1 |1123| ST.NO. SPECIES | P MS RS | COVER AND VIGOR A1 A2 A3 C 1 PSEUDOTSUGA MENZIESII 100 0 7 9 6-8 7 7 8 8 8 . 7 8 . S . 7 2 TSUGA HETEROPHYLLA 55 G 4 5 0-5 4 5 . 5 5. 4 3 THUJA PLICATA 44 4 3 4 0-5 3 5 4 . 2 4 CHAMAECYPARIS NOOTKATENSIS 1 1 1 1 1 0-3 3. 5 PINUS CONTORTA 1 1 1 + 3 0-2 2 PSEUDOTSUGA MENZIESII 100 0 5 2 4 - 7 7 5 3 5 3 . 4 6. 4 . 2 TSUGA HETEROPHYLLA 65 7 3 7 0-5 2 5 4 . 4 2 . 3 THUJA PLICATA 33 3 3 1 0-5 3 5. 2 . 6 ABIES AMABILIS CHAMAECYPARIS NOOTKATENSIS 1 1 1 1 1 1 1 + 1 0 0-3 0- 1 1 . 3. TSUGA HETEROPHYLLA 88 9 5 0 0-7 2 7 4 3 . 5 3. 5. 5 THUJA PLICATA 77 8 4 0 0-5 2 5 5. A 2. 3 . 4 PSEUDOTSUGA MENZIESII 55 6 2 2 0-3 3 3 3 1 4- . ABIES AMABILIS 1 1 1 1 7 0-4 4 . CHAMAECYPARIS NOOTKATENSIS 1 1 1 1 1 0-3 3. 7 PINUS MONTICOLA PINUS CONTORTA 1 1 1 1 1 1 4 + 3 0 0-2 0- + 2 4 8 VACCINIUM PARVIFOLIUM " 1 1 1 0-3 • • • |3 a l 9 GAULTHERIA SHALLON 100 0 9 3 8-9 9 2 9 2 9 2 9 2 9.2 9 3 9.2 8.3 9 2 10 BERBERIS NERVOSA 44 4 + 9 0-2 + 2 2. 1 1 . 2 4 4-VACCINIUM PAR.."r0LIUM 33 3 3 0 0-5 + + 3 2 5 2 1 1 ROSA GYMNOCARPA 33 3 4 0 0 -4 + 1 + + 4 1 12 VACCINIUM ALASKAENSE 22 2 1 8 0-4 1 2 4.2 13 ARCTOSTAPHYLOS MEDIA 1 1 1 4 0 0- 1 1 4 14 JUNIPERUS COMMUNIS 1 1 1 4 0 0- 1 1 4 15 AMELANCHIER ALNIFOLIA 1 1 1 + 0 0- + 4 4 1G CHIMAPHILA UMBELLATA 88 9 3 2 0-4 4 2 3 2 1 1 3 . 2 1 1 4 . 2 4 . 2 3 2 17 LINNAEA BOREAL IS 77 8 4 9 0-7 2 2 1 1 * . 1 1 2 1 . 1 7 . 2 7 3 18 LI ST ERA CORDATA 7 7 8 2 0 0-3 3 2 4 4- 4 4 2 . 1 1 1 4 . 4 3 2 19 GOODYERA OBLONGIFOLIA 77 8 + 5 0- 1 4 2 * 2 4 2 4 1 4 . 1 4- 4 1 1 20 CHIMAPHILA MENZIESII 55 6 1 4 0-3 1 1 1 1 * . 1 1 1 3 . 1 2 1 POLYPODIUM GLYCYRRHIZA 44 4 4 2 0- 1 4 • 4 1 4 4 1 . 1 22 POLYSTICHUM MUNITUM VACCINIUM PARVIFOLIUM 33 33 3 3 1 1 1 0 0-3 0-2 4 2 4 4 4 4 2. 1 3 . 1 2. 1 23 TRIENTALIS LATIFOLIA 33 3 1 0 0-2 1 1 2 . 1 1 2 24 FESTUCA OCCIDENTALIS 33 3 + 6 0- 1 1 2 1 2 1 . 1 25 LACTUCA MURAL IS 22 2 1 3 0-3 3 2 2 . 2 26 BOSCHNIAKI A HOOKER I 22 2 + 0 0-1 4 2 1 2 27 CORALLORHIZA MACULATA 22 2 + 0 0-1 1 . 2 4 2 28 HIERACIUM ALBIFLORUM 22 2 4 0 0-1 4 2 1 2 29 VIOLA ORB ICULATA 22 2 + 0 0- 1 4- 1 1 2 30 CORAL LORHIZA MERTENSIANA 22 2 + 0 0- + 4 . 1 4 2 3 1 POLYSTICHUM LONCHITIS 22 2 4 0 0 -4 4 2 4 1 32 ARCTOSTAPHYLOS UVA-URSI 1 1 1 1 1 0-3 3 2 33 CAMPANULA SCOULERI GAULTHERIA SHALLON 11 \ 1 1 1 1 0-3 0-3 3 2 3 1 34 VIOLA GLABELLA 1 1 1 1 1 0-3 3.2 35 CORNUS CANADENSIS 1 1 1 + 3 0-2 2.2 3S ALLOTROPA VIRGATA 1 1 1 4 0 0-1 1 3 37 HEUCHERA MICRANTHA 1 1 1 + 0 0- 1 1 2 38 MONT 1A PARVIFOLIA 1 1 1 4- 0 0- 1 1 2 39 TIARELLA LACINATA 1 1 1 4- 0 0- 1 1 . 2 40 TIARELLA TRIFOLIATA 1 1 1 4- 0 0- 1 1 . 1 4 1 ACHLYS TRIPHYLLA 1 1 1 4 0 0-4 4 4 42 CLADOTHAMNUS PYROLAEFLORUS 1 1 1 4 0 0-4 4 2 43 LISTERA CAURINA 1 1 1 4 0 0 -4 * . 1 44 PYROLA PICTA 1 1 1 4 0 0-4 4 2 APPENDIX VI (continued) T a b l e A 3 0 . A s s o c i a t i o n t a b l e f o r t h e T S H E / G A S H - B E N E / A C T R h a b i t a t t y p e . PLOT 1 SYNTHETIC | 1 1 1 1 1 1 NUMBER | VALUES jl200|l1B.1 |ltn.5|t18O(l201 f202| ST.NO. SPECIES | P MS RS | COVER AND VIGOR B2 1 PSEUDOTSUGA MENZIESII 100. 0 7 . 9 7-8 7 . 7 . 8 . 7 . 7 . 8 . 2 TSUGA HETEROPHYLLA 66 . 7 5. 1 0-7 4 . 5. 3. 7 . 3 THUJA PLICATA 60 . 7 4 . 5 0-5 3 . 5 . 5. 4 . 4 PINUS MONT I COLA 16 . 7 1 . 4 .0-3 3 . TSUGA HETEROPHYLLA 100. 0 5. 2 4-5 5 . 4 . 5. 4 . 5. 5. PSEUDOTSUGA MENZIESII 66 . 7 4 . 0 0-5 3 . 4 . 3 . 5 . THUJA PLICATA 50. 0 2 . 9 0-4 2 . 4 . 3 . TSUGA HETEROPHYLLA 100. 0 6. 8 4-7 7 . 7 . 5. 7 . 7 . 4 . THUJA PLICATA 83 . 3 4 . 4 0-5 5 . 3. t . 3 . 5 . PSEUDOTSUGA MENZIESII 66 . 7 1 . 6 0-2 2 . 1 2 . 1 . 5 ABIES AMABILIS 16 . 7 1 . 4 0-3 3 . 6 TAXUS BREVIFOLIA 16 7 1 . 4 0-3 3 . 7 GAULTHERIA SHALLON 100. 0 7 2 3-9 5 . 1 3 . 2 9 2 5 . 2 9. 3 4 . 1 8 ROSA GYMNOCARPA 100 0 4 6 2-6 2 . 2 3 . 1 4 3 2. 2 4 . 2 6. 2 9 BERBERIS NERVOSA 83 3 3 1 0-4 1 1 4 . 2 3 2. 3. 2 1 . 2 10 VACCINIUM PARVIFOLIUM 83 3 2 4 0-3 1 1 1 . 2 3 2 3 2 1 . 1 1 1 VACCINIUM ALASKAENSE 66 7 3 3 0-5 + 1 5 . 1 2. 1 4 1 12 RUBUS URSINUS 66 7 1 7 0-3 4 2 3 2 1 2 1 2 13 SYMPHORICARPOS ALDUS 50 0 1 9 0-3 3 2 1 2 2 2 14 SYMPHORICARPOS MOLLIS 50 0 1 0 0- 1 1 2 1 2 1 2 15 VACCINIUM OVALIFOLIUM 50 0 4 8 0- 1 4 1 1 2 1 1 16 AMELANCHIER ALNIFOLIA. 50 0 + 4 0- 1 4- 4 1 1 4 1 17 PACHYSTIMA MYRSINITES 16 7 4 0 0- 1 1 1 18 VACCINIUM MFMBRANACEUM 16 7 4 0 0-1 1 1 19 RIBES LACUS1RE 16 7 +• 0 0-4 4 2 20 RUBUS PARVIFLORUS 16 7 4 0 0-4 4 4 2 1 ACHLYS TRIPHYLLA 100 0 5 0 3-5 4 2 5 2 4 2 4 3 3 2 5 2 22 LATHYRUS NEVADENSIS 100 0 4 1 4-5 4 1 4 2 4 2 4 4 4 1 5 2 23 LINNAEA BOREALIS 100 0 2 7 1-3 1 2 3 2 2 2 1 2 3 2 2 2 24 CHIMAPHILA UMBELLATA 100 0 1 8 4-2 2 2 1 2 4 1 2 2 1 2 1 2 25 ADENOCAULON BICOLOR too 0 1 6 -4-3 4 4 4 4 4 2 4 1 4 1 3 2 26 GOODYERA OBLONGIFOLIA 100 0 1 2 4- 1 4 2 1 2 4- 1 1 2 4 ' 1 2 27 VIOLA ORBICULATA 83 3 3 7 0-4 4 2 4 1 4 2 3 2 4 2 28 CORNUS CANADENSIS 83 3 3 2 0-4 1 2 4 2 2 2 4 2 4 2 29 PEDICULARIS RACEMOSA 83 3 3 0 0-4 4 2 1 2 4 1 3 2 2 2 30 PTERIDIUM AOUILINUM 6G 7 2 2 0-4 4 1 4 2 4 1 4 1 31 FESTUCA OCCIDENTALS VACCINIUM PARVIFOLIUM S6 66 7 7 1 1 7 0-3 1 0-2 1 2 2 i 1 4 2 1 4 i 1 2 3 4 2 1 32 OSMORHIZA CHILENSIS 66 7 + 5 0-1 4 1 4 1 4 2 1 2 33 CHIMAPHILA MENZIESII GS 7 + 1 0-* 4 1 4 4 4 4 4 2 34 LACTUCA MURAL IS 50 0 + 0 0-* 4 1 4 4 4 2 35 POLYSTICHUM MUNITUM 50 0 4 0 0-4 4 4- 4 4 4 4 36 TIARELLA TRIFOLIATA 33 3 3 2 0-5 5 2 1 2 37 CAMPANULA SCOULERI 33 3 2 .3 0-4 1 2 4 2 38 TRIENTALIS LATIFOLIA VACCINIUM ALASKAENSE 33 33 . 3 . 3 1 4 .4 0-3 .6 0-1 4 . 1 1 1 1 . 1 3 . 2 39 PYROLA ASARIFOLIA 33 . 3 + .2 0-1 4 . 1 1 2 40 LILIUM COLUMBIANUM 33 . 3 +-.0 0-4 4 . 4 4 . 1 4 1 LISTERA CORDATA 33 . 3 + .0 0-4 4 + 4 . 4 42 POLYSTICHUM LONCHITIS 33 . 3 4 .0 0-4 4 . 2 4 . 4 43 STREPTOPUS AMPLEXIFOLIUS 33 . 3 4-.0 0-4 4 . 2 4 . 1 44 TIARELLA LACINATA 33 . 3 + .0 0-4 4 2 4 . 2 45 TRILLIUM OVATUM ROSA GYMNOCARPA 33 1G . 3 . 7 4 1 .0 O-* .4 0-3 4 . 1 4 3 . 4 . 2 46 ARENARI A MACROPHYLLA 16 . 7 4 .0 0-1 1 . 2 47 FRAGARI A VESCA RUBUS URSINUS 16 1G .7 ..7 4 4-.00-1 .00-1 1 . 1 1 . 1 48 VIOLA GLABELLA AMELANCHIER ALNIFOLIA 16 16 .7 . 7 4 4 .0 0-1 .0 0-4 1 4 .2 . 4 49 ANEMONE LYALII 1G . 7 4 .0 0-* 4 . 1 50 ATHYRIUM FILIX-FEMINA 16 . 7 4 .0 0-4 4 . 4 5 1 GAULTHERIA OVATIFOLIA 16 . 7 4 .0 0-4 4-. 2 52 HIERACIUM ALB IF LORUM 16 . 7 4 .0 C-4 4 . 2 53 LISTERA CAURINA 16 . 7 4 .0 0-4 4 . 2 54 POLYPODIUM GLYCYRRHIZA RIBES LACUSTRE 16 16 . 7 . 7 4. .0 O-* .0 0-4 4 4-. 2 55 STENANIHIUM OCCIDENTALE 16 . 7 4 .0 0-4 + . 2 56 TIARELLA UNIFOLIATA 16 . 7 .4 .0 0-4 4 • 4 231 APPENDIX VI (continued).. Table A 31. A s s o c i a t i o n table f o r the TSHE/POMU habitat type. PLOT 1 SYNTHETIC 1 1 1 1 1 1 NUMBER 1 VALUES 114 3 |1122 |1135 |11 28 |f1 37 |1138| ST.NO. SPECIES | P MS RS COVER AND VIGOR A1 C 1 PSEUDOTSUGA MENZIESII 100 0 6 7 3-8 7 . 3. 5 . 7 5 8 2 TSUGA HETEROPHYLLA 66 7 5 7. 0 -8 7 . 4 8. 5 3 THUJA PLICATA 33 3 5 3 0 -9 3 . 9 . TSUGA HETEROPHYLLA 6G 7 5 8 0 -8 8 . 8 4 + THUJA PLICATA 50 0 3 0 0-4 4 . 3. 3 4 ACER MACROPHYLLUM 16 7 1 4 0 -3 3. 5 ABIES GRANDIS 16 7 + 0 0 - 1 1 . 6 PINUS MONTICOLA PSEUDOTSUGA MENZIESII 16 16 7 7 + + 0 0 0 - 1 o-+ 1 . + . TSUGA HETEROPHYLLA 100 0 5 8 3-7 5. 7 . 5. 7 3 5 THUJA PLICATA 66 7 2 2 0-4 + . 4 . + + PSEUDOTSUGA MENZIESII 33 .3 2 1 0 -3 3. 3 7 ABIES AMABILIS ABIES GRANDIS 16 16 7 7 + + 0 0 o-+ o-+ + . + 8 VACCINIUM PARVIFOLIUM 16 7 + 0 0 - • • • 1 I 9 BERBER I 3 NERVOSA 100 0 4 7 1-7 1 . 2 2 . 1 4 . 2 1 2 7 2 2 1 VACCINIUM PARVIFOLIUM 100 0 2 4 + -3 3 . 1 1 . + 1 . 2 1 1 3 1 + 1 10 GAULTHERIA SHALLON 50 0 1 2 0 -2 2 . 2 1 . + 1 1 1 1 GAULTHERIA OVATIFOLIA 16 7 + 0 o-+ +. 1 12 POLYSTICHUM MUNITUM 100 0 7 4 3-9 3 . 2 4.2 8.2 9 3 5 2 8 2 13 ACHLYS TRIPHYLLA 100 0 4 0 + -5 2 . 1 + . 1 3 . 2 2 1 5 2 4 2 14 T IARELLA TRIFOLIATA 100 0 2 5 + -3 3 . 2 + . + 2 : 2 1 2 1 2 3 2 15 TRILLIUM OVATUM 100 o +• 4 + - + + . 2 + . 1 * . 2 + 1 + 1 +• 2 16 TIARELLA LACINATA 66 7 2 0 0 -3 2 . 2 1 2 1 2 3 2 17 CHIMAPHILA UMBELLATA 66 7 1 8 0-3 2 . 2 + . + +• 4- 3 2 18 VIOLA ORBICUL-ATA 66 7 1 1 0 - 1 1 . 2 + . 1 1 2 1 2 19 GOODYERA OBLONGIFOLIA • 66 7 -f 5 0-1 + .2 1 . 1 + 2 + 2 20 TR IENTALIS LATIFOLIA 50 0 1 5 0 -3 + . 1 3 . 2 + 1 21 CHIMAPHILA MENZIESII 50 0 1 1 0 -2 2 . 2 1 2 + 1 22 LACTUCA MURAL IS 50 0 1 1 0 -2 1 . 1 + 4- 2 2 23 POLYSTICHUM LONCHITIS 50 0 1 0 0 -2 + . 1 + 2 2 1 24 GALIUM TRIFLORUM 33 3 1 0 0-2 2 . 1 1 1 25 LI ST ERA CORDATA 33 3 + 0 0 - + + . + + 1 26 RUBUS URSINUS VACCINIUM PARVIFOLIUM 33 16 3 7 + 3 0 2 0-+ 0 -5 4- . + + + 5 1 27 BLECHNUM SPICANT 16 7 2 2 0-4 4 2 28 LINNAEA BOREALIS 16 7 1 4 0 -3 3 2 29 CORNUS CANADENSIS 16 7 + 7 0 -2 2 2 30 L ISTERA CAURINA 16 •7 + 7 0 -2 2 . 2 31 LUZULA PARVIFLORA 16 7 + 7 0 -2 2 2 32 LYCOPODIUM CLAVATUM 16 7 + 7 0 -2 2 2 33 ATHYRIUM F IL IX-FEMINA 16 7 + 0 0 - 1 1 1 34 C0RALL0RH1ZA MACULATA 16 7 + O O- 1 1 2 35 ADENOCAULON BICOLOR 16 7 + o 0 - + + . 1 36 ADIANTUM PEDATUM 16 7 + 0 o-+ + .2 37 GALIUM SPP 16 7 + 0 0 - + + 1 38 MADIA SATIVA 16 7 0 0 - + + . 1 39 MONOTROPA UNIFORA. 16 7 + 0 0 - + 2 40 PTERIDIUM AOUILINUM 16 7 + 0 0 - + 41 RUBUS PEDATUS 16 7 4- 0 0 - + + 1 42 VIOLA GLABELLA 16 7 0 0 - + 232 APPENDIX VI (continued). Table A 32. A s s o c i a t i o n table f o r the u n c l a s s i f i e d samples i n the TSHE zone, PLOT NUMBER | SYNTHETIC 1 VALUES jl14s|l131 |*134 j,129| ST.NO. SPECIES | P MS RS |COVER AND VIGOR A 1 A2 A3 B1 B2 1 PSEUDOTSUGA MENZIESI I 2 THUJA PLICATA PSEUDOTSUGA MENZIESI I 3 TSUGA HETEROPHYLLA THUJA PLICATA PSEUDOTSUGA MENZIESI I TSUGA HETEROPHYLLA THUJA PLICATA 4 HOLODISCUS DISCOLOR 5 BERBER IS NERVOSA 6 ROSA GYMNOCARPA HOLODISCUS DISCOLOR 7 SYMPHORICARPOS ALBUS ' 8 ACER GLABRUM 9 VACCINIUM PARVIFOLIUM 10 PRUNUS EMARGINATA .11 RUBUS SPECTABIL IS 12 LINNAEA BOREAL IS 13 CHIMAPHILA UMBELLATA 14 TR IENTALIS LATIFOLIA 15 GOODYERA OBLONGIFOLIA 16 CAMPANULA SCOULERI 17 CHIMAPHILA MENZIESI I 18 FESTUCA OCCIDENTALIS 19 HIERACIUM ALBIFLORUM 20 LACTUCA MURALIS 21 ACHLYS TRIPHYLLA 22 ARENARIA MACROPHYLLA 23 MONT IA PARVIFOLIA 24 VIOLA ORBICULATA 25 POLYSTICHUM LONCHITIS 26 CALYPSO BULBOSA 27. FRAGARIA VESCA 28 BROMUS INERMIS 29 POLYSTICHUM MUNITUM VACCINIUM PARVIFOLIUM 30 POLYPODIUM GLYCYRRHIZA 31 PYROLA PICTA BERBER IS NERVOSA 32 ADENOCAULON BICOLOR 33 BROMUS VULGARIS 34 GALIUM TRIF LORUM 35 MELICA SUBULATA 36 MADIA SATIVA. 37 OSMORHIZA CHILENSIS 38 BROMUS SPP 39 CORALLORHIZA'MACULATA 40 GAULTHERIA SHALLON 4 1 GERANIUM SPP 42 LATHYRUS NEVADENSIS 43 LISTERA CORDATA 44 NOTHOCHELONE NEMOROSA 45 PTEROSPORA ANDROMEDIA 100.0 8.2 7-8 8. 25 .0 2.7 0-4 8. 4 . 7. 100.0 5.2 +-5 3 . i - . 5. ' 5. 5 0 . 0 1.8 0 -3 3 . 4 . 2 5 . 0 4.4 0 -6 5 . 100.0 4 .0 +-5 3 . 4 . 4 . 5. 5 0 . 0 4 .0 0-5 5 . 3 . 2 5 . 0 4.4 0 -6 G . | 25.O 1.8 0 - 3 | I | 3 2 | I 5 0 . 0 5.0 0 -7 2 . 1 7 . 2 5 0 . 0 4 . 0 0 -5 5.2 3. 2 25 .0 3.7 0 -5 5 . 2 25 .0 1.8 0 -3 3. 1 2 5 . 0 + . 3 0-1 1 . 1 1 2 5 . 0 +.3 0-1 1 . 25 .0 +.0 0-+ + . + 25 .0 + 0 0-+ + . 1 1 0 0 . 0 4 .0 1-5 1 . 1 1 . 2 2.2 5 2 100.0 3.0 +-4 + . 1 + . 1 2 . 2 4 2 100.0 1.6 +-2 + . 2 1 . 1 2. 1 1 2 100.0 +.5 +-+ 4. 2 + . 2 + . 1 + 2 7 5 . 0 2.9 0-4 1 . + + . 2 4 2 7 5 . 0 2.3 0 -3 1 . 2 2 . 2 3- 2 7 5 . 0 2.3 0 -3 1 . 1 2.2 3 2 75 .0 2.0 0 -3 + 1 3.2 1 2 7 5 . 0 1 9 0 -3 + 4 + . 1 3 . 2 7 5 . 0 1.4 0-2 1 + + + 2 . 1 75 .0 1.4 0 -2 + 1 2 .2 1 . 1 7 5 . 0 1.4 0-2 1 2 2.2 +• .2 7 5 . 0 1 1 0-1 1 2 1 2 4- .2 75 .0 +.8 0-1 + 1 + 2 1, 1 75 .0 +.2 0-+ + 1 4- 1 + . 1 5 0 . 0 2.0 0-3 3.2 1 . 2 5 0 . 0 1.8 0 -3 4- 1 3.2 5 0 . 0 1.1 0-2 +• + 2 . 1 5 0 . 0 1.0 0-1 1 . + 1 . + 5 0 . 0 +.6 0-1 + . 1 1 . 1 5 0 . 0 +.6 0-1 + .2 1 . 2 5 0 . 0 + 0 0-+ + . + 4- . + 2 5 . 0 1.8 0 -3 3.2 25 .0 1.8 0 -3 3 . 2 25 .0 1.8 0 -3 3.2 2 5 . 0 1.8 0 -3 3.2 25.O 1,1 0-2 2.2 2 5 . 0 1.1 0-2 2 . 1 25 .0 +-.0 0 - + + . 1 2 5 . 0 +.0 0-+ + . 1 2 5 . 0 +.0 0-+ . + 25 .0 +.0 0-+ + . 1 2 5 . 0 +.0 O-+ + .2 2 5 . 0 +.0 0-+ + . 2 25 .0 +.0 0-+ + . 2 25 .0 + .0 0 - + + .3 APPENDIX VI (continued) Table A33. Association table for the CHNO/GASH habitat type. PLOT NUMBER 1 SYNTHETIC 1 VALUES |l1 18 | i i 58 |l1 S9 i«15G jl157 | ST.NO. SPECIES | P MS RS | COVER AMD VIGOR 233 A 1 A2 A3 B2 1 PSEUDOTSUGA MENZIESII 2 TSUGA HETEROPHYLLA 3 THUJA PLICATA 4 CHAMAECYPARIS NOOTKATENSIS 5 TSUGA MERTENSIANA 6 ABIES AMABILIS TSUGA HETEROPHYLLA THUJA PLICATA CHAMAECYPARIS NOOTKATENSIS PSEUDOTSUGA MENZIESII 7 PINUS MONTICOLA TSUGA HETEROPHYLLA CHAMAECYPARIS NOOTKATENSIS THUJA PLICATA PINUS MONTICOLA TSUGA MERTENSIANA ABIES AMABILIS PSEUDOTSUGA MENZIESII 8 GAULTHERIA SHALLON 9 VACCINIUM ALASKAENSE 10 VACCINIUM MEMBRANACEUM 11 VACCINIUM PARVIFOLIUM 12 CHIMAPHILA UMBELLATA 13 LINNAEA BOREALIS 14 GAULTHERIA OVATIFOLIA 15 PYROLA SECUNDA VACCINIUM PARVIFOLIUM VACCINIUM MEMBRANACEUM 16 CHIMAPHILA MENZIESII 17 LISTERA CAURINA 18 HYPOPITYS MONOTROPA 19 CAMPANULA SCOULERI 20 FESTUCA OCCIDENTALIS 21 LACTUCA MURAL IS 22 LISTERA CORDATA 23 MADIA SATIVA 24 MONTIA PARVIFOLIA 25 POLYSTICHUM LONCHITIS VACCINIUM ALASKAENSE 26 PTEROSPORA ANDROMEDIA 27 GOODYERA OBLONGIFOLIA 28 BERBERIS NERVOSA 29 CORALLORHIZA MERTENSIANA 30 TRIENTALIS LATIFOLIA 31 VIOLA ORBICULATA 100. 0 3 4 1-4 3 . 1 1 2 4 . 2 3. 2 2 . 2 80. 0 1 8 0-3 3 . 1 + 1 + . 1 1 . 1 60 0 1 8 0-3 1 1 3 1 1 60 0 + 0 o-+ + 2 + 1 +. + 40 0 4 0 0-5 4 1 5 2 40 0 1 7 0-3 3 1 1 1 40 0 1 2 0-2 2 1 1 1 40 0 1 2 0-2 1 2 2 1 40 0 + 0 o-+ + 2 + 2 20 0 1 5 0-3 3 1 20 0 1 5 0-3 3 2 20 0 1 5 0-3 3 1 20 0 1 5 0-3 3 1 20 0 1 5 0-3 3 2 20 0 '1 .5 0-3 3 1 20 0 1 . 5 0-3 3 1 20 .0 1 .5 0-3 3 . 1 20 .0 1 .0 0-2 2 2 20 .0 + . 2 0- 1 1 . 1 20 .O + .0 o- + + . 1 20 .0 + .0 o-+ + .2 . 1 20 .0 + .0 o-+ + 20 .0 + .0 o-+ + . + 100. 0 5 . 9 3-8 3 . 5 . 5 . 5. 8 . 80. 0 7 . 7 0-8 8 . 7 . 8 . 8. 40. 0 3 0 0-4 3. 4 . 40. 0 2 3 0-3 3 . 3 . 40 0 2 0 0-3 3. 2 . 20 O + O o-+ + . 100 0 4 8 + -5 4- 4 . 5 . 5 . 3. 60 O 2 9 0-3 3 . 3. 3. 60 0 4 9 0-3 3 . 1 . 1 . 40 0 2 5 0-4 1 . 4 . 20 0 1 5 0-3 3. 80 0 5 2 0-7 7 . 3 . 5. 3. 80 0 4' 2 0-5 3 . 4 . 2 . 5 . 40 0 3 7 0-5 3 . 5 . 40 0 2 0 0-3 3 2 . 20 .0 + . 2 0-1 1 20 .0 + .0 o-+ + . 20 .0 + .0 o-+ 4- . 100 .0 8 . 1 5-9 9. 1 8 + 5 . 1 8. 1 7 . 1 60 .0 1 .0 0-1 + . 1 1 1 1 . 1 20 .0 4 . 6 0-7 7 3 20 .0 1 . 5 0-3 3 . 1 APPENDIX VI (continued) Table A34. Association table for the ABAM/VAAL-VAPA habitat type. PLOT- I S Y N T H E T I C 1 1 1 ! 1 1 1 1 1 1 NUMBER | V A L U E S |1l40|l'49|l15oit130|l203 |i20<1 |lt97 |l199|1181 | S T . N O . S P E C I E S | P MS RS |• COVER AND VIGOR A 1 A2 A3 B2 C 1 TSUGA HETEROPHYLLA 100 0 7 2 3-8 7 . 7 . 8 . 7 5 7 . 7 . 7 3 2 PSEUDOTSUGA MENZIESII 88 9 6 4 0-8 4 . 6. 4 . 6 7 7 . 7 . 8 3 THUJA PLICATA 66 7 4 6 0-5 5. 5. 4 . 5 . 4 . 4 4 ABIES AMABILIS 22 2 3 5 0-6 3 . 6 5 CHAMAECYPARIS NOOTKATENSIS 1 1 1 1 7 0-4 4 e TSUGA MERTENSIANA 1 1 1 1 7 0-4 4 7 PINUS MONTICOLA 1 1 1 + 3 0-2 2 TSUGA HETEROPHYLLA 100 0 4 6 4.-5 4 . 4 . 4 4 4 5. 4 . 5 3 PSEUDOTSUGA MENZIESII 66 7 3 4 0-5 3 . 3 2 5. 2 . 3 THUJA PLICATA 44 4 3 5 0-5 4 . 5 2 . 4 ABiES AMABILIS 22 2 2 8 0-5 1 5 CHAMAECYPARIS NOOTKATENSIS 22 2 1 8 0-4 4 1 TSUGA HETEROPHYLLA 100 0 5 5 1-8 4 . 4 . 1. 8 7 4 . A . 4 5 ABIES AMABILIS 77 8 3 8 0-5 3 . 4 . 4 . 3 5. 4 1 THUJA PLICATA 77 8 3 7 0-4 3. 1 . 3. 4 4 . 4 . 4 CHAMAECYPARIS NOOTKATENSIS 55 6 3 9 0-5 3 . 3. 5 2 5 PSEUDOTSUGA MENZIESII 22 2 1 6 0-3 3 . 3 8 VACCINIUM ALASKAENSE 77 8 4 3 0-5 1 . 1 3 . 1 5 2 3. 1 4 . 1 5 2 5 2 9 BERBERIS NERVOSA 66 7 1 4 0-3 4 . 1 4 2 1 2 1 . 1 3 2 4 1 10 VACCINIUM PARVIFOLIUM ' 55 6 3 1 0-4 3 . 1 4 2 3 . 1 1 . 1 4 2 1 1 VACCINIUM MEMBRANACEUM 55 6 2 6 0-4 3 2 4 1 1 . 1 1 . 1 3 2 12 VACCINIUM OVALIFOLIUM 44 4 3 2 0-5 5 2 2 . 1 1 2 4. 2 13 GAULTHERIA SHALLON 33 3 4 5 0-2 2. 1 4 . | 4 1 14 ROSA GYMNOCARPA 22 2 + 0 0-4 4 2 4.2 IS RHODODENDRON ALBIFLORUM 1 1 1 •4 3 0-2 2 1 16 SORBUS SITCHENSIS .' 1 1 1 4 0 0- + 4 1 17 LINNAEA BOREALIS 100 0 3 4 + -5 1 . 2 4 . 2 1 . 2 5 2 3 2 1 . 2 4 . 2 1 2 4 2 VACCINIUM PARVIFOLIUM 77 8 3 3 0-4 3 . 2 4 2 1 1 4 2 3 . 1 2 . 1 3 2 18 CHIMAPHILA UMBELLATA 77 8 3 1 0-4 1 . 2 3 . 3 3 2 4 2 1 .2 4 2 4 2 19 CHIMAPHILA MENZIESII 77 8 1 1 0-2 4 . 1 1 . 2 4 . 1 4 2 1 . 2 2 . 1 4 2 20 ACHLYS TRIPHYLLA 66 7 1 9 0-3 1 1 1 2 3.2 4 . 4 4 2 3 2 2 1 VIOLA ORBTCULATA 66 7 1 4 0-2 4 . 1 1 2 2 2 1 . 2 1 . 1 2 2 22 RUBUS PEDATUS 66 7 4 9 0-1 4 . 1 4 . 4 1 2 1 2 1 .2 4 1 23 GOOOYERA OBLONGIFOLIA 66 7 4 7 0- 1 4 . 1 4 2 4 2 4 . 1 1 . 2 1 2 24 TIARELLA TRIFOLIATA 55 6 2 1 0-4 4 . 1 1 2 4 2 1 . 1 2 2 25 CORNUS CANADENSIS 55 6 1 5 0-3 1 .2 3 2 1 .2 1 2 1 2 26 LISTERA CORDATA 55 6 + 8 0-1 1 .2 1 3 4 4 4 . 4 1 4 27 POLYSTICHUM MUNITUM 44 4 4 0 0- + 4 1 4 . 4 4 . 4 4 1 28 TRILLIUM OVATUM 44 4 4 0 0- + 4- 1 4 . V 4 2 4 1 29 PEOICULARIS RACEMOSA 33 3 1 1 0-3 4 1 4 2 3 2 30 CLINTONIA UNIFLORA 33 3 4 6 0- 1 1 2 1 2 1 2 3 1 LISTERA CAURINA 33 3 4" 3 0- 1 1 2 4 . 1 1 2 32 PYROLA SECUNOA 33 3 + 1 0-1 1 2 4 2 4.2 33 TIARELLA LACINATA 33 3 4 1 0-1 1 2 4 2 4 . 1 34 HYPOPITYS MONOTROPA VACCINIUM ALASKAENSE 33 22 3 2 4 1 0 2 0-4 0-3 4 . 2 1 . 1 4 1 4 2 3 2 35 GAULTHERIA OVATIFOLIA 22 2 4 7 0-2 1 2 2 2 36 LATHYRUS NEVADENSIS 22 'j 4 2 0- 1 1 2 1 2 37 LYCOPUDIUM CLAVATUM 22 2 4 2 0- 1 1 .2 1 2 38 STENANTHIUM OCCIOENTALE 22 2 -4 2 0- 1 1 2 1 2 39 ATHYRIUM FI LIX-F EMINA BERBERIS NERVOSA 22 22 2 2 4 4 0 0 0-4 0-4 4 . 4 4 1 4 . 4 4 . 4 40 POLYSTICHUM LONCHITIS 22 2 4- 0 0-4 4 . 1 4 . 2 4 1 PYROLA ASARIFOLIA 22 2 + 0 0-4 4 1 4 2 42 STREPTOPUS ROSEUS 22 2 + 0 0-4 4 1 4 1 43 STREPTOPUS STREPTOPOIDES VACCINIUM OVALIFOLIUM 22 22 2 2 4 4 0 0 0-4 0-4 4 . 1 4 . 1 4 1 4 1 44 ARNICA LATIFOLIA 1 1 1 4 3 0-2 2 2 45 BLECHNUM SPICANT 1 1 1 4 3 0-2 2 2 46 COUALLORHIZA MACULATA 1 1 1 + 0 0- 1 1 . 3 47 LYSICHITUM AMERICANUM VACCINIUM MEMBRANACEUM 1 1 4 4 0 0 0- 1 0- 1 1 . 4 1 2 48 AMELANCHIER ALNIFOLIA 11 1 4- 0 0-4 4 4 49 CORALLORHIZA SPP 11 1 • 0 0-4 4 . 2 50 FESTUCA OCCIOENTALIS GAULTHERIA SHALLON JJ 1 1 4 4 0 0 0-4 0-4 4 2 4 4 51 LACTUCA MURAL IS 11 1 4 0 0-* 4 . 1 52 LILIUM C0LUM8IANUM 11 1 + 0 0-4 4 1 53 OSMORHIZA CHILENSIS 11 1 4 0 O- > 4 1 APPENDIX VI (continued) Table A35. A s s o c i a t i o n t a b l e f o r the ABAM/ACTR-TITR h a b i t a t t y p e . PLOT SYNTHETIC NUMBER I VALUES |I1 79 |11 33 |11 39 |11 1 7 |1132 | ST.NO. SPECIES I P MS RS | COVER A NO VIGOR A 1 A2 A3 B2 C 1 PSEUDOTSUGA MENZIESII 80. 0 7 . 4 0-8 8 . 8 . 5 . 8. 2 TSUGA HETEROPHYLLA GO. 0 5 . 3 0-8 3 . 8 . 4 . 3 THUJA PLICATA 40. 0 4 . 0 0-5 4 . 5 . 4 ABIES AMABILIS 40. 0 3 . 7 0-5 5 . 3 . TSUGA HETEROPHYLLA 40. 0 5 . 2 0-8 8 . 4 . PSEUDOTSUGA MENZIESII 40. 0 3. 2 0-4 4 . 4 . ABIES AMABILIS 40. 0 3 . 0 0-4 4 . 3 . THUJA PLICATA 40. 0 2 G 0-4 4 . 2 . TSUGA HETEROPHYLLA 80. 0 5 . 6 0-8 8 . 3 . 5 . 5. THUJA PLICATA GO. 0 3 . 4 0-5 5 . 4 4 . 5 TSUGA MERTENSI ANA ABIES AMABILIS 40. 20. 0 0 1 3 6 0-3 4 0-5 4 . 3 . 5 . 6 TAXUS BREVIFOLIA PSEUDOTSUGA MENZIESII 20. 20. 0 0 + + 2 0-1 0 0-4 1 . 4 . 7 VACCINIUM PARVIFOLIUM 60 0 1 9 0-3 1 . 1 3 . 2 1 1 8 BERBERIS NERVOSA GO 0 1 0 0-1 1 . 1 1 . 1 4 . 2 9 VACCINIUM ALASKAENSE 40 0 1 2 0-2 1 2 2 . 1 10 ROSA GYMNOCARPA 40 0 + 40-1 1 1 4 . 4 1 1 PACHYSTIMA MYRSINITES 20 0 4 2 0- 1 1 2 12 VACCINIUM MEMBRANACEUM 20 0 4 2 0- 1 1 1 13 SYMPHORICARPOS ALBUS 20 0 4 0 0-4 4 1 14 ACHLYS TRIPHYLLA 100 0 4 9 1-7 1 2 2 2 1 2 3 2 7 2 15 TIARELLA TRIFOLIATA 100 0 3 8 4-5 3 2 4 2 1 2 2 2 5 3 1G TIARELLA LAC INATA 100 0 2 8 4-4 4 2 4 2 1 2 2 2 4 3 17 CHIMAPHILA MENZIESII 1O0 0 2 3 1-3 1 2 1 2 1 2 2 2 3 2 18 VIOLA ORBICULATA 80 0 1 4 0-2 2 2 1 2 4 1 1 1 2 19 POLYSTICHUM MUNITUM 80 0 1 3 0-2 4 1 2 4 1 1 4 20 LACTUCA MURAL IS 80 0 4 7 0-1 4 4 4 1 4 1 1 1 2 1 TRILLIUM OVATUM 80 0 4 7 0-1 1 1 4 1 4 1 4 2 22 LINNAEA BOREALIS SO 0 1 2 0-2 2 2 1 2 4 1 23 CAMPANULA SCOULERI 60 0 1 0 0- 1 1 2 4 1 1 2 24 CHIMAPHILA UMBELLATA 60 0 1 0 0-1 1 2 4 4 1 2 25 GOODYERA OBLONGIFOLI A 60 0 1 0 0-1 1 2 1 2 4 2 2G POLYSTICHUM LONCHITIS 60 0 4 5 0- 1 4 1 4 2 1 1 27 PYROLA SECUNDA 60 0 4 5' 0-1 1 2 4 2 4 2 28 POLYPCDIUM GLYCYRRHIZA 60 0 4 0 0-4 4 4 4 4 4 4 29 BLECHNUM SPICANT VACCINIUM PARVIFOLIUM 40 40 0 0 1 1 6 0-3 2 0-2 2 2 1 1 4 2 3 2 30 BROMUS SPP 40 0 1 0 0-2 2 2 4 2 3 1 CORNUS CANADENSIS 40 0 4 8 0- 1 1 1 1 1 32 NOTHOCHELONE NEMOROSA 40 0 4 4 0- 1 1 2 4 2 33 PYROLA PICTA 40 0 4 .40-1 1 2 4 1 34 RUBUS PEDATUS 40 0 4 .40-1 1 2 4 1 35 TRIENTALIS LATI FOLIA 40 .0 4 .4 0-1 4 1 1 1 36 ARENARIA MACROPHYLLA 40 0 4 0 0-4 4 2 4 4 37 CLINTONIA UNI FLORA 40 0 4 .0 0-4 4 4 4 1 38 LILIUM COLUMBIANUM 40 0 4 .0 0-4 4 4 4 1 39 RIBES LACUSTRE 40 0 4 .0.0-4 4 1 4 . 4 40 LISTERA CORDATA 20 .0 1 .5 0-3 3 2 4 1 TIARELLA UNIFOLIATA VACCINIUM MEMBRANACEUM 20 20 .0 .0 4 4 .20-1 .2 0-1 1 1 . 2 . 1 42 VIOLA GLABELLA 20 .0 4 .20-1 1 . 2 43 VIOLA SPP 20 .0 4 .20-1 1 . 1 44 ADENOCAULON BICOLOR 20 .0 4 .00-4 4 . 4 45 ARNICA LATI FOLIA 20 .0 4 .0 0-4 4 . 2 46 GAULTHERIA OVATIFOLIA 20 .0 4 .0 0-4 4 . 2 47 HIERACIUM ALBIFLORUM 20 .0 4 .0 0-4 4 . 2 48 HYPOPITYS MOMOTROPA 20 .0 4 .0 0-4 4 .2 49 LUZULA SPP 20 .0 4 .0 0-4 4 . 2 50 MADIA SATIVA 20 .0 4 .0 0-4 4 . 1 5 1 MONT I A PARVIFOLIA 20 .0 4 .0 0-4 4 . 4 52 MONT I A SIBIRICA 20 .0 4 .OO-i 4 . 2 53 PTEROSPORA ANOROMEOI A 20 .0 4 .0 0-4 4 .2 54 RUBUS NIVALIS 20 .0 4 .00-4 4 . 2 236 APPENDIX VI (continued) Table A36. Association table for the ABAM/VAAL/STREPTOPUS habitat type. PLOT 1 SYNTHETIC I 1 1 1 1 1 NUMBER | VALUES |ll83|l13G|1t74|(173|«70| ST.NO. SPECIES | P MS RS ICOVER AND VIGOR A 1 82 C t TSUGA HETEROPHYLLA 100.0 7 . 8 6-8 7 . 6. 8 . 7 . 8. 2 PSEUDOTSUGA MENZIESII 80.0 5. 1 0-5 5. 5. 5. 4 . 3 ABIES AMABILIS 60.0 4 . 7 0-5 4 . 5 . 5. TSUGA HETEROPHYLLA I 80.0 4 4 o-s l 5 . I 4 . I I 4 . I 3. I PSEUDOTSUGA MENZIESII I 20.0 2 4 0-4 | -1 . I I I I I ABIES AMABILIS 100.0 4 1 4-5 4 . 5 3 . 1 . 4 . TSUGA HETEROPHYLLA 60.0 5 0 0-5 5 . 5 5 . 4 THUJA PLICATA 40.0 1 0 0-2 2 . 4 . 5 VACCINIUM ALASKAENSE 80.0 4 1 0-5 Cj 2 3 2 1 . 2 4 . 1 6 VACCINIUM 0VAL1F0LIUM 80.0 2 6 0-3 2 2 3 2 3. 2 1 . 2 7 VACCINIUM PARVIFOLIUM 60.0 4 0 0-5 5 3 4 2 1 1 8 BERBERIS NERVOSA' 40.0 1 7 0-3 3 2 1 2 9 VACCINIUM MEMBRANACEUM 20.0 1 5 0-3 3 3 10 ARUNCUS SYLVESTER 20.0 + 2 0- 1 1 . 2 1 1 ACER GLABRUM 20.0 + 0 0-4 4 4 12 MENZIESIA FERRUGINEA 20.0 * 0 0-4 4 1 13 RHODODENDRON ALBIFLORUM 20.0 4 0 0-4 4 4 14 RIBES LACUSTRE 20.0 4 0 0-4 + 1 15 SAMBUCUS SPP 20.0 4 0 0-4 4 4 16 ACHLYS TRIPHYLLA 100.0 3 6 2-4 2 2 4 3 3 2 3 2 3 2 17 TIARELLA TRIFOLIATA 100.0 3 4 1-4 2 2 3 2 1 2 3 2 4 1 18 PYROLA SECUNDA 100.0 2 2 4-3 4 2 1 3 2 2 1 2 3 2 19 STREPTOPUS AMPLEXIFOLIUS 100.0 1 3 4-2 4 2 1 ^ 4 2 4 2 2 1 20 TIARELLA LACINATA 100.0 1 3 4-2 * 2 4 2 4 2 1 2 2 1 2 1 RUBUS PEDATUS 80.0 3 0 0-4 1 2 4 2 4 2 3 1 VACCINIUM PARVIFOLIUM 80.0 2 7 0-4 4 1 4 3 2 2 4 1 22 STREPTOPUS ROSEUS 80.0 1 3 0-2 1 2 4 2 4 1 2 1 23 CHIMAPHILA MENZIESII 80.0 1 0 0- 1 4 1 1 2 4 2 1 2 24 TRILLIUM OVATUM 80.0 + 7 0- 1 4 2 4 2 4 2 1 2 1 25 STREPTOPUS STREPTOPOIDES 60.0 3 6 0-5 2 2 5 3 2 26 LINNAEA BOREALIS 60.0 2 3 0-3 3 2 3 3 + 2 27 LYCOPODIUM CLAVATUM 60.0 2 3 0-3 3 3 3 2 4 4 28 CORNUS CANADENSIS 60.0 2 0 0-3 3 2 2 3 4 2 29 POLYSTICHUM MUNITUM GO.O 1 8 0-3 * 1 1 2 3 2 30 STENANTHIUM OCCIDENTALS 60.0 1 2 0-2 1 2 4 2 2 2 31 CHIMAPHILA UMBELLATA 60.0 1 1 0- 1 1 2 1 2 1 2 32 BLECHNUM SPICANT 60.0 1 0 0- 1 1 2 4 2 1 1 33 LISTERA CAURINA 60.0 + 5 0-1 1 2 4 2 4 2 34 LUZULA SPP 60.0 + 0 0-4 4 2 4 2 4 2 35 POLYSTICHUM LONCHITIS SO.O 4 0 0--:- 4 - + 4 2 4 3 36 CLINTONIA UMIFLORA 40.0 2 .0 0-3 3 2 2 2 37 VIOLA ORB ICULATA 40.0 4 . 8 0- 1 1 2 1 .2 38 LISTERA CORDATA 40.0 4 . 4 0- 1 4 2 1 . 3 39 OSMORHIZA CHILENSIS 40.0 + . 4 0-1 1 2 4 2 40 PYROLA PICTA VACCINIUM MEMBRANACEUM 40.0 40.0 4 4 .0 .0 0-4 0-4 4 1 4 . 2 4 4 2 t 4 1 ADIANTUM PEDATUM 20.0 + . 2 0- 1 1 2 42 GALIUM SPP 20.0 4 . 2 0- 1 1 .2 43 GYMNOCARPIUM DRYOPTERIS 20.0 + . 2 0- 1 1 . 2 44 POA SPP VACCINIUM ALASKAENSE 20.0 20.0 4 + . 2 . 2 0-1 0-1 1 .2 1 . 2 45 VIOLA GLABELLA 20.0 4 . 2 0- 1 1 2 46 ADENOCAULON BICOLOR 20.0 4 .0 0-* 4 . 4 47 ARENARI A MACROPHYLLA 20.0 4 .0 0-4 4 1 48 A7HYRIUM FILIX-FEMINA 20.0 4 .0 0-4 4 . 2 49 CALYPSO BULBOSA 20 0 4 .0 0-4 4 . 2 50 DRYOPTERIS AUSTRIACA 20.0 + .0 0-4 4 4 5 1 EPILOBIUM ANGUSTIFOLIUM 20.0 4 .O 0-4 4 4 52 GALIUM TRIFLORUM 20.0 4 .0 0-4 4 . 2 53 GAULTHERIA OVATIFOLIA 20.0 4 .0 0-4 4 . 2 54 HEUCHERA MICRANTHA 20.0 4 .0 0-4 4 . 2 55 HYPOPITYS MONOTROPA 20.0 4 .0 0-4 4 . 2 56 LACTUCA MURAL IS 20.0 4 .0 0-* 4 . 4 57 MEL I CA SL'SULATA 20.0 * .0 r\~ 4 4 . 2 58 MONTIA PARVIFOLIA 20.0 4 .0 4 .2 59 MONTIA S1BIRICA 20.0 4 .0 0-» 4 . 2 60 NOTHOCHSLONE NEMOROSA 20.0 * .0 0--> 4 .2 61 PEDICULARIS RACEMOSA 20.0 4 .0 0-4 4 2 62 POLYPODIUM GLYCYRRHIZA 20.0 4 .0 o--» 4 . 2 63 SMILACtNA RACEMOSA . 20.0 4 .0 0-* 4 . 2 64 VERATRUM VIRIDE 20.0 4 .0 0- + 4 . 1 237 APPENDIX VI (continued)". Table A37. As s o c i a t i o n table f o r the ABAM/VAAL(OV)/RUPE habitat type. PLOT 1 SYNTHETIC 1 1 I I I ! NUMBER 1 VALUES |,14 1 |115 1 |1169|1152|1182| ST.NO. SPECIES | P MS RS | COVER AND VIGOR A 1 A2 A3 B2 C 1 TSUGA HETEROPHYLLA 100. 0 8 . 5 7-9 9 . 8 . 8. 7 . 8 . 2 TSUGA MERTENSIANA 80. 0 5 . 2 0-7 4 . 3 . 7 . 5. 3 ABIES AMABILIS 60. O 4 . 1 0-5 3 . 4 . 5. 4 PSEUDOTSUGA MENZIESII 40. 0 3 . 2 0-4 4 . 4 . 5 CHAMAECYPARIS NOOTKATENSIS 40. 0 2 . 3 0-3 3 . 3 . 6 THUJA PLICATA 20. 0 2 . 4 0-4 4 . TSUGA HETEROPHYLLA 100. 0 5 . 1 + -5 5 . 5 . + 4 . 5 . ABIES AMABILIS 60. 0 3 . 2 0-4 4 . 3 . 3 . TSUGA MERTENSIANA 60. 0 2 5 0-3 3 . 3. 2 . CHAMAECYPARIS NOOTKATENSIS 40. 0 1 7 0-3 3 . 1 . ABIES AMABILIS 100 0 5 1 3-5 5 . 3 . 5 . 5. 3. THUJA PLICATA 80 0 1 8 0-3 r 1 . + . + . CHAMAECYPARIS NOOTKATENSIS 80 0 1 3 0-2 1. + . + . 2 . TSUGA HETEROPHYLLA 60 0 3 9 0-5 3 3 . 5. TSUGA MERTENSIANA 20 0 + 0 0- + + 7 VACCINIUM ALASKAENSE 100 0 6 0 1-8 8 2 3 1 7 . 1 1 . 1 5 2 8 VACCINIUM OVALIFOLIUM 80 0 3 1 0-4 + 2 2 1 3 . 1 4 2 9 VACCINIUM MEMBRANACEUM 60 0 1 9 0-3 1 2 3 . 1 1 1 2 10 RHODODENDRON ALBIFLORUM 60 0 + 0 0- + + 2 + + 2 . 1 + . 1 1 MENZIESIA FERRUGINEA 20 0 1 0 0-2 12 SORBUS SITCHENSIS 20 0 1 0 0-2 2. 1 13 VACCINIUM PARVIFOLIUM 20 0 + 0 o-+ + 2 14 RUBUS PEDATUS 100 O 4 4 + -6 3 2 + + 6.2 1 1 3 2 15 PYROLA SECUNDA 80 0 1 0 0- 1 1 2 + 2 + 1 1 2 1S TIARELLA TRIFOLI ATA 60 0 + 5 0-1 + 1 + 1 1 2 17 STREPTOPUS STREPTOPOIDES 40 0 2 3 0-3 3.2 3 2 18 LYCOPODIUM CLAVATUM 40 0 1 6 0-3 3 2 + 2 19 CHIMAPHILA UMBELLATA 40 0 + 4 0- 1 1 2 + 1 1 VACCINIUM PARVIFOLIUM 40 0 + 4 0-1 1 1 + 20 VERATRUM VIPIDE 40 0 + 4 0-1 + 1 1 2 2 1 HYPOPITYS MONOTROPA 40 0 + .0 0- + + 2 4- 2 22 LISTERA CAURINA 40 .o + .0 o-+ 4- . 1 4- 2 23 TIARELLA LACINATA 40 o +• .o 0- + + 1 + T 24 8LECHNUM SPICANT 20 0 1 .5 0-3 3 2 25 CORNUS CANADENSIS 20 .0 1 .0 0-2 2 . 2 26 GYMNOCARPIUM DRYOPTERIS 20 .O + o O- 1 1 . 2 27 LINNAEA BOREALIS VACCINIUM ALASKAENSE 20 20 .0 .0 + + .2 . 2 0-1 0- 1 1 1 . 2 . 2 28 CHIMAPHILA MENZIESII 20 .0 4-.0 o-+ . 2 29 DRYOPTERIS AUSTRIACA 20 .0 4. .0 o-+ + . 2 30 GAULTHERIA OVATIFOLIA 20 .o + .0 o-+ + . 2 . 1 31 LISTERA CORDATA 20 .0 + .0 o-+ 1 + 32 STREPTOPUS AMPLEXIFOLIUS 20 .0 + .0 o-+ + 33 VIOLA ORBICULATA 20 .0 + .0 0- + + . + APPENDIX VI (continued) 238 Table A 3 8 . A s s o c i a t i o n table for the ABAM/OPHO habi t a t type. PLOT 1 SYNTHETIC 1 1| 1| NUMBER | VALUES | l 7 l | l 7 2 | ST.NO. SPECIES | P MS RS | COVER AND VIGOR C 1 ABIES AMABILIS 1 100 .0 8 .0 7-8 Is |7, 2 TSUGA HETEROPHYLLA 1 100 .0 6 .5 5-7 U h • ABIES AMABILIS I 100 .0 5 .0 3-5 I3 I5 TSUGA HETEROPHYLLA I 50 .0 2 .6 0 -3 |3 I • ABIES AMABILIS j 100 .0 5 . 5 4-6 I4 Is TSUGA HETEROPHYLLA j 100 .0 2 . 7 + -3 |3 I* • 3 OPLOPANAX HORRIDUM 100 .0 4 . 7 + -5 + 2 5 . 3 4 RUBUS SPECTABIL IS 50 .0 1 .6 0 -2 2 .2 5 RIBES LACUSTRE 50 .0 + .0 0 - + + 1 6 VACCINIUM PARVIFOLIUM 50 .O + .0 0 - + +• + 7 TIARELLA TRIFOLIATA 100 0 5 .8 5-6 5 2 6 3 8 ACHLYS TRIPHYLLA 100 0 4 .9 2-5 2 2 5 3 9 POLYSTICHUM MUNITUM 100 0 3 6 1-4 4 2 1 1 10 ATHYRIUM F IL IX-FEMINA 100 0 2 9 1-3 1 2 3 3 1 1 STREPTOPUS ROSEUS 100 0 2 "7 + -3 + 2 3 2 12 STREPTOPUS AMPLEXIFOLIUS 100 0 2 0 1-2 1 2 2 3 13 TIARELLA LACINATA 100 0 1 5 1-1 1 2 1 3 1.4 TRILLIUM OVATUM 100 0 1 5 1-1 1 2 1 2 15 MONTIA S IBIRICA 100 0 1 2 + - 1 + 2 1 2 16 OSMORHIZA CHILENSIS 100 0 1 2 +'- 1 + 2 1 2 VACCINIUM PARVIFOLIUM 100 0 + 5 + - + + + + 1 17 GYMNOCARPIUM DRYOPTERIS 50 0 4 7 0 -5 5 3 18 TRAUTVE T TERIA CAROL INIENSIS 50 o 3 5 0-4 4 2 19 ADENOCAULON BICOLOR 50 0 1 0 0 - 1 1 2 20 DRYOPTERIS AUSTRIACA 50 0 1 0 0-1 1 . 2 2 1 RUBUS PEDATUS 50 o 1 0 0 - 1 1 3 22 VALERIANA SCOULERI 50 0 1 0 0 - 1 1 2 23 VIOLA GLABELLA 50. 0 1 0 0-1 1 . 2 24 ADIANTUM PEDATUM 50. 0 + 0 0 - + + . 2 25 LACTUCA MURALIS 50. 0 + 0 0 - + + . 1 26 LISTERA CAURINA 50. 0 + o 0 - + 2 27 POLYPODIUM GLYCYRRHIZA 50. 0 + 0 0 - + 2 28 POLYSTICHUM LONCHITIS 50. 0 0 0 - + 2 29 STREPTOPUS STREPTOPOIOES 50. 0 + . o 0 - + f . 2 30 TIARELLA UNIFOLI ATA 50. 0 + . 0 o-+ + . 2 31 VERATRUM VIRIDE 50. o 0 0 - + + . 2 APPENDIX VI (continued) 239 Table A39. A s s o c i a t i o n table for samples i n the ABAM-TSME vegetation zone. PLOT NUMBER 1 SYNTHETIC 1 VALUES jlt.12 |l144 |l153 |ll6?. ll1G3 !ne><l |ll65 |l16S |l1G7 JllGsl ST.NO. SPECIES | P MS RS | COVER AND VIGOR At A 2 A 3 B 2 1 T S U G A M E R T E N S I A N A 100.0 5 . 8 4 - 7 7 . 4 . 7 . 7 . 5 . 4 . 4 . 6 . 4 . 6 . 2 A B I E S A M A B I L I S 9 0 . 0 5 . 9 0 - 8 4 . 7 . 4 . 4 . 5 . 8 . 5 . 8 . 5 . 3 T S U G A H E T E R O P H Y L L A 5 0 . 0 4 . 5 0 - 5 5 . 5 . 5 . 4 . 5 . 4 C H A M A E C Y P A R I S N O O T K A T E N S I S 50.0 3 . 6 0-5 4 . 5 . 4 . 4 . 2 . 5 A B I E S L A S I O C A R P A 2 0 . 0 4 . 6 0 - 7 7 . 7 . 6 P S E U D O T S U G A M E N Z I E S I I 1 0 . 0 2 . 6 0 - 5 5 . T S U G A M E R T E N S I A N A 90.0 3 . 9 0 - 5 l 4 . 4- . 5 . 4 . 3 . 4 4 . 3 . 4 . A B I E S A M A B I L I S 7 0 . 0 5 . 0 0 - 6 5 . n 4 . 4 . 5 . 6 . 4 . C H A M A E C Y P A R I S N O O T K A T E N S I S 5 0 . 0 3 . 2 0 - 4 3 . 4 . 4 . 4 . 3 . T S U G A H E T E R O P H Y L L A 3 0 . 0 1 . 2 0 - 3 1 . 3 . 1 . A B I E S L A S I O C A R P A 2 0 . 0 3 . 4 0 - 5 5 . 5 . A B I E S A M A B I L I S 1 0 0 . 0 5 8 1 - 7 7 . 6 . 5 . 7 . 7 . 7 . 1 . 2 . 3 . 4 . T S U G A M E R T E N S I A N A 1 0 0 . 0 4 5 4 - 5 5 . 4 3 4 . 5 . 2 . 5 . 3 3 . 3 . C H A M A E C Y P A R I S N O O T K A T E N S I S 6 0 . 0 2 0 0 - 3 + . 3 3 . 3 4 . + • A B I E S L A S I O C A R P A 3 0 . 0 2 9 0 - 5 4 5 . 3 T S U G A H E T E R O P H Y L L A 3 0 . 0 2 9 0 - 5 3 . 5 * • 7 T H U J A P L I C A T A 1 0 . 0 * 0 0-4 8 V A C C I N I U M M E M B R A N A C E U M 1 0 0 . 0 4 2 + - 5 3 2 3 2 1 1 4 . 2 5 2 3 1 5 2 4 2 3 1 4 2 9 V A C C I N I U M A L A S K A E N S E 9 0 . 0 5 1 0 - 8 8 2 1 2 5 2 5. 2 1 2 4 1 3 2 4 1 5 2 1 0 V A C C I N I U M O V A L I F O L I U M 9 0 . 0 4 7 0 - 5 1 2 4 2 5 2 5 2 3 1 1 2 5 2 3 1 5 2 11 R H O D O D E N D R O N A L B I F L O R U M 8 0 . 0 4 7 0 - 6 5 2 6 3 3 2 1 2 5 2 5 2 3 1 1 2 1 2 S O R B U S S I T C H E N ; ' . S 4 0 . 0 * 3 0 - 1 1 2 4 4 4 1 1 2 1 3 M E N Z I E S I A F E R R U G I N E A 2 0 . 0 * 1 0-1 1 2 1 2 1 4 P H Y L L O D O C E E M P E T R I F O R M I S 2 0 . 0 * 1 0 - 1 1 2 1 2 1 5 R I B E S L A C U S T R E 2 0 . 0 * 0 0 - 1 4 2 1 2 16 C L A D O T H A M N U S P Y R O L A E F L O R U S 1 0 . 0 * 0 0 - 1 1 2 17 R U B U S P E D A T U S 1 O 0 . 0 4 9 + - 6 6 2 5 2 1 1 3 1 4 1 5 2 5 2 3 2 3 2 3 1 1 8 P Y R O L A S E C U N D A 7 0 . 0 . 1 5 0 - 3 1 2 4 1 3 2 1 2 1 2 1 2 1 2 1 9 T I A R E L L A T R I F O L I A T A 6 0 . 0 1 3 0 - 3 4 2 1 2 4 2 4 1 1 2 3 2 2 0 V E R A T R U M V I R I D E 6 0 . 0 4 9 0 - 2 4 2 •»- 2 4 1 4 2 1 2 2 1 2 1 S T R E P T O P U S S T R E P T O P O I D E S 4 0 . 0 2 5 0 - 4 4 3 4 1 2 2 2 1 2 2 2 V I O L A O R B I C U L A T A 4 0 . 0 T 3 0 - 3 1 2 1 2 3 2 1 2 3 L I S T E R A C A U R I N A 4 0 . 0 1 0 0 - 2 2 2 4 1 1 2 1 2 2 4 G O O D Y E R A O B L O N G I F O L I A 4 0 . 0 * 1 0 - 1 4 2 4 1 1 2 4 1 2 5 L U Z U L A S P P 4 0 . 0 + 0 0 - * 4 2 4 2 4 2 4 2 1 2 6 A R N I C A L A T I F O L I A 3 0 . 0 1 7 0 - 4 + 2 4 2 2 2 7 C L I N T O N I A U N I F L O R A 3 0 . 0 * 2 0 - 1 1 2 4 2 1 2 2 8 P O L Y S T I C H U M L O N C H I T I S 3 0 . 0 + 0 0 - + 4 2 * 2 + 1 2 9 V A L E R I A N A S C O U L E R I 3 0 . 0 + 0 0 - + 4 2 4 2 4 2 3 0 L U E T K E A P E C T I N A T A 2 0 . 0 1 1 0 - 3 3 2 + + 3 1 A C H L Y S T R I P H Y L L A 2 0 . 0 + 0 0 - + 2 3 2 B L E C H N U M S P I C A N T 2 0 . 0 * . 0 0-1 4 1 1 1 3 3 S T R E P T O P U S R O S E U S 2 0 . 0 + . 0 0 - 1 + 2 1 2 3 4 A N E M O N E L Y A L I I 2 0 . 0 * . 0 0 - + * 1 4 2 35 A T H Y R I U M F I L I X - F E M I N A 2 0 . 0 + . 0 0 - + 4 1 4 4 3 6 H E U C H E R A G L A B R A 2 0 . 0 + . 0 o- + 4 2 4 . 2 3 7 P O L Y S T I C H U M M U N I T U M 2 0 . 0 + .0 0-* 4 4 4 4 3 8 S T R E P T O P U S A M P L E X I F O L I U S 2 0 . 0 . * . 0 0 - + 4 . 2 4 . 2 3 9 C O R N U S C A N A D E N S I S 1 0 . 0 + . 0 0-1 1 1 4 0 G A U L T H E R I A O V A T I F O L I A 1 0 . 0 + . 0 0 - 1 1 . 2 4 1 L I N N A E A B O R E A L I S 1 0 . 0 * .00-1 1 2 . 2 4 2 L U Z U L A P A R V I F L O R A 1 0 . 0 * .00-1 1 4 3 O S M O R H I Z A C H I L E N S 1 S 1 0 . 0 * . 0 0 - 1 1 . 2 4 4 T I A R E L L A L A C I N A T A 1 0 . 0 + . 0 0-1 1 . 2 4 5 V A C C I N I U M P A R V I F O L I U M 1 0 . 0 * .0 0-1 1 . 1 4 G V I O L A G L A B E L L A 1 0 . 0 • . 0 0 - 1 1 . 3 4 7 C H I M A P H I L A U M B E L L A T A 1 0 . 0 4 .0 0-+ 4 . 1 4 8 F R I T I L L A R I A C A M S C H A I C E N S I S 1 0 . 0 + . 0 0 * 4 . 2 4 9 H Y P O P I T Y S M O N O T R O P A 1 0 . 0 * . 0 0 - + + . 2 5 0 L 1 L I U M C O L U M B I A N U M 1 0 . 0 + . 0 0 - * 4 . 2 5 1 L O N I C E R A S P P 1 0 . 0 + . 0 0 - < 4 . 2 5 2 L Y C O P O D I U M C L A V A T U M 1 0 . 0 * . 0 0 - * . 2 5 3 P E O I C . ^ A R I S R A C E M O S A 1 0 . 0 * . 0 0 - + * . 2 5 4 P O L Y P O D I U M G L Y C Y R R H I Z A 1 0 . 0 * . 0 0 - + 4 . 4 5 5 P Y R O L A A S A R I F O L I A 1 0 . 0 + .0 0-* 4 . 2 R I B E S L A C U S T R E 1 0 . 0 + . 0 0 - + 4 . 1 5 6 T R I L L I U M O V A T U M 1 0 . 0 * . 0 0-4 . 2 

Cite

Citation Scheme:

        

Citations by CSL (citeproc-js)

Usage Statistics

Share

Embed

Customize your widget with the following options, then copy and paste the code below into the HTML of your page to embed this item in your website.
                        
                            <div id="ubcOpenCollectionsWidgetDisplay">
                            <script id="ubcOpenCollectionsWidget"
                            src="{[{embed.src}]}"
                            data-item="{[{embed.item}]}"
                            data-collection="{[{embed.collection}]}"
                            data-metadata="{[{embed.showMetadata}]}"
                            data-width="{[{embed.width}]}"
                            async >
                            </script>
                            </div>
                        
                    
IIIF logo Our image viewer uses the IIIF 2.0 standard. To load this item in other compatible viewers, use this url:
https://iiif.library.ubc.ca/presentation/dsp.831.1-0095194/manifest

Comment

Related Items