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The use of phenology to define range readiness in riparian and upland habitats of the Cariboo-Chilcotin… Hemphill, Susan M. 1998

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THE USE OF PHENOLOGY TO DEFINE RANGE READINESS IN RIPARIAN AND UPLAND HABITATS OF THE CARIBOO CHILCOTIN REGION, BRITISH COLUMBIA by Susan M. Hemphill B Sc., The University of British Columbia 1994 B. Sc., The University of Michigan 1969 M. A., California State University 1971 A THESIS SUBMITTED IN PARTIAL FULFILMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE in THE FACULTY OF GRADUATE STUDIES (Department of Plant Science) We accept this thesis as corrforming to the required standard THE UNIVERSITY OF BRITISH COLUMBIA August, 1998 ©Susan M. Hemphill, 1998 In presenting this thesis in partial fulfilment of the requirements for an advanced degree at the University of British Columbia, I agree that the Library shall make it freely available for reference and study. I further agree that permission for extensive copying of this thesis for scholarly purposes may be granted by the head of my department or by his or her representatives. It is understood that copying or publication of this thesis for financial gain shall not be allowed without my writ ten permission. Department of \ \ <XW The University of British Columbia Vancouver, Canada Date A u q \ 1 \C{ ^ % DE-6 (2/88) Abstract The main objectives of this study were (1) to determine how yearly changes in climate affects phenology at sites in the Bunchgrass (BG) and the Sub-Boreal Spruce (SBS) Biogeoclimatic Zones of the Cariboo-Chilcotin region of British Columbia, (2) to clarify how phenology varies among plant associations which are related to soil moisture gradients in grasslands, (3) to clarify how phenology varies among plant associations which are related to soil moisture and canopy cover gradients in forested range, and (4) to describe and compare the phenological patterns found in riparian areas and surrounding uplands at sites in the BG and SBS. A set of permanent plots was established in each of the riparian and upland plant associations at the two BG sites and one SBS site. I observed and recorded the phenology of each species growing in each plot from April to September in 1995 and 1996. Soil moisture and degree day totals at each site were also measured. Hierarchial cluster analysis and ordering species by flowering dates were used to group species with similar phenologies. Spring degree day totals were close to 20 year averages in 1995, but in 1996, they were only 69% of average at the BG sites and 73% of average at the SBS site. Changes in phenology between 1995 and 1996 reflected these differences. Forty percent of the species at the three sites flowered two or more weeks later in 1996 than in 1995. Twenty-four percent of the species dispersed fruit later in 1996. The higher levels of winter precipitation in 1996 increased the time in spring that soils were saturated, and changes in phenology were greatest in the species growing in the transition areas between riparian and upland habitats. More species growing in these areas flowered two weeks later in 1996 than species growing in ii other areas. I categorized species into five general phenological groups based on flowering dates (Early Spring, Spring, Early Summer, Mid Summer, and Late Summer). The Early Spring and Late Summer groups each had distinct, uniform phenologies. The three mid season groups (Spring, Early Summer, and Mid Summer) overlapped in both flowering and fruit dispersal times and included a variety of phenologies. Individual upland and riparian plant associations differed in at least one aspect of their phenological patterns related to dates of flowering and fruit dispersal and/or proportion of species included in each phenological group. In addition, the phenology of some species growing in several associations varied among associations. These differences suggested that species growing in each plant association had phenological adaptations related to the habitat and microclimate where each association grew. The use of indicator species clarified the differences in upland and riparian phenologies. At the BG sites, only one riparian species flowered in early spring compared to 17% of the upland species. By early summer, only 50% of the riparian species had flowered compared to 63% of the upland species. Fruit dispersal was three weeks later in respective riparian phenological groups than in upland groups. At the SBS site, flowering times were similar in the riparian areas and uplands, but species in respective phenological groups dispersed fruit two to four weeks later in the riparian area than species growing in the upland areas. Many BG species common across the uplands flowered up to two weeks later growing in the associations with higher soil moisture (transition and deep swale plant associations). At the SBS site, species generally flowered earlier in the clear cut (low levels of canopy cover) than in the other upland forest associations (higher levels of canopy cover). iii Table of Contents Abstract 11 Table of contents iv List of Tables vi List of Figures ix Acknowledgements xi 1. Introduction 1 1.1 Literature Review 3 1.1.1 Use of Phenology to Understand Plant Communities 3 1.1.2 Range Management in Uplands and Their Associated Riparian Areas 5 1.2 Objectives 10 2. Study Area 11 2.1 Bunchgrass Study Area 13 2.2 Sub-Boreal Study Area 14 3. Methods 16 3.1 Site Selection and Establishment of Plots 16 3.2 Vegetation Composition and Coverage 17 3.3 Abiotic Measurements 22 3.4 Phenological Data 24 3.5 Statistical Analysis 2 5 4. Results 33 4.1 Growing Degree Days in 1995 and 1996 33 4.2 Soil Moisture and Precipitation 33 4.2.1 Plant Associations Related to Soil Moisture and Tree Canopy Gradients 33 4.2.2 Changes in 1995 and 1996 Levels of Precipitation and Soil Moisture 40 4.3 Vegetation Coverage in Each Plant Association 44 4.3.1 Grassland sites 44 4.3.2 Sub-Boreal Site 46 4.4 Phenology 47 4.4.1 Identifying Phenological Groups and Indicators 47 4.4.2 Phenological Groups 49 4.4.3 Bunchgrass Phenologies 5 5 4.4.4 Sub-Boreal Spruce Phenologies 67 5. Discussion 80 iv 5.1 Phenological Changes Related to Growing Degree Days 80 5.2 Effects on Phenology of Soil Moisture and Precipitation Changes Between 1995 and 1996 82 5.3 Variations in Species' Phenology Between Plant Associations 84 5.4 Phenological Groups Identified 86 5.5 Bunchgrass Riparian and Upland Phenologies 88 5.6 Sub-Boreal Spruce Riparian and Upland Phenologies 91 6. Summary 94 7. Management Recommendations 97 7.1 Phenological Summary of a Site as part of a Range Management Plan 97 7.2 Using Phenology to Design Range Management Plans 99 8. Literature Cited 105 9. Appendix 1: Ordered data matrixes for each plant association, BGxw and BGxw/h sites, Junction Sheep Range Park and SBS site, Horsefly, British Columbia, 1996 113 10. Appendix 2: Hierarchial cluster tree diagrams for riparian and upland summaries, BGxw site, Junction Sheep Range Park and SBS site, Horsefly, British Columbia, 1996 132 11. Appendix 3: Plant species found in each plant association BGxw, Bgxw/h sites, Junction Sheep Range Park, and SBS site, Horsefly, British Columbia, 1996 139 v List of Tables 1. Hours herb and shrub layers not shaded, SBS site, Horsefly, British Columbia, 1996 22 2. Phenology codes used at BG and SBS sites, Junction and Horsefly study areas, British Columbia, 1995/1996 25 3. Combined phenological code used for statistical analysis, Bunchgrass and Sub-Boreal Spruce study areas, British Columbia, 1995/1996 29 4. 1996 ordered data matrix, dry grassland association, BGxw site, Junction Sheep Range Park, British Columbia 32 5. Plant associations, BGxw site, Junction Sheep Range Park, British Columbia, 1995/1996 37 6. Plant associations, BGxw/h site, Junction Sheep Range Park, British Columbia, 1995/1996 38 7. Plant associations, SBS site, Horsefly, British Columbia, 1995/1996 39 8. Riparian area summary ordered data matrix, BGxw site, Junction Sheep Range Park, British Columbia, 1996 57 9. Riparian area summary ordered data matrix, BGxw site, Junction Sheep Range Park, British Columbia, 1995 58 10. Number of species reaching flowering and fruit dispersal later in 1996 than in 1995, BGxw site, Junction Sheep Range Park, British Columbia 1995/1996 59 11. Number of life forms found in each phenological group, BGxw site, Junction Sheep Range Park, British Columbia, 1996 60 12. Upland area summary ordered data matrix, BGxw site, Junction Sheep Range Park, British Columbia, 1996 63 13. Upland area summary ordered data matrix, BGxw site, Junction Sheep Range Park, British Columbia, 1995 64 vi 14. Riparian area summary ordered data matrix, SBS site, Horsefly, British Columbia, 1996 70 15. Riparian area summary ordered data matrix, SBS SITE, Horsefly, British Columbia, 1995 71 16. Number of species reaching flowering and fruit dispersal later in 1996 than in 1995, SBS site, Horsefly, British Columbia, 1995/1996 72 17. Number of life forms found in each phenological group, SBS site, Horsefly, British Columbia, 1996 .' 73 18. Upland area summary ordered data matrix, SBS site, Horsefly, British Columbia, 1996 74-75 19. Upland area summary ordered data matrix, SBS site, Horsefly, British Columbia, 1995 76-77 20. Summary of the phenological progress, BGxw site, Junction Sheep Range Park, British Columbia, 1996 103 21. Summary of the phenological progress, SBS site, Horsefly, British Columbia, 1996 104 Appendix Tables: A. 1.1. 1996 ordered data matrix, open water and wet riparian association, BGxw site, Junction Sheep Range Park, British Columbia 115 A. 1.2 1996 ordered data matrix, dry riparian association, BGxw site, Junction Sheep Range Park, British Columbia 115 A. 1.3 1996 ordered data matrix, transition association, BGxw site, Junction Sheep Range Park, British Columbia 116 A. 1.4 1996 ordered data matrix, dry grassland association, BGxw site, Junction Sheep Range Park, British Columbia 117 A. 1.5 1996 ordered data matrix, shallow swale association, BGxw site, Junction Sheep Range Park, British Columbia 118 A. 1.6 1996 ordered data matrix, deep swale association, vii BGxw site, Junction Sheep Range Park, British Columbia A. 1.7 1996 ordered data matrix, open water and wet riparian association, BGxw/h site, Junction Sheep Range Park, British Columbia A. 1.8 1996 ordered data matrix, dry riparian association, BGxw/h site, Junction Sheep Range Park, British Columbia A. 1.9 1996 ordered data matrix, transition association, BGxw/h site, Junction Sheep Range Park, British Columbia A. 1.10 1996 ordered data matrix, dry grassland association, BGxw/h site, Junction Sheep Range Park, British Columbia A. 1.11 1996 ordered data matrix, shallow swale association, BGxw/h site, Junction Sheep Range Park, British Columbia A. 1.12 1996 ordered data matrix, deep swale association, BGxw/h site, Junction Sheep Range Park, British Columbia A. 1.13 1996 ordered data matrix, open water association, SBS site, Horsefly, British Columbia A. 1.14 1996 ordered data matrix, riparian association, SBS site, Horsefly, British Columbia A. 1.15 1996 ordered data matrix, open edge association, SBS site, Horsefly, British Columbia A. 1.16 1996 ordered data matrix, forest association, SBS site, Horsefly, British Columbia A. 1.17 1996 ordered data matrix, path association, SBS site, Horsefly, British Columbia A. 1.18 1996 ordered data matrix, clearcut association, SBS site, Horsefly, British Columbia viii List of Figures 1. Location of study areas in British Columbia, 1995/1996 12 2. Side and top view, BGxw site, Junction Sheep Range Park, British Columbia, 1995/1996 18 3. Side and top view, BGxw/h site, Junction Sheep Range Park, British Columbia, 1995/1996 19 4. Side view, SBS site, Horsefly, British Columbia, 1995/1996 20 5. Top view, SBS site, Horsefly, British Columbia, 1995/1996 21 6. Tree diagram: dry grassland association, BGxw site, Junction Sheep Range Park, British Columbia, 1996 30-31 7. Weekly degree day totals and monthly averages, BGxw and BGxw/h sites, Junction Sheep Range Park, British Columbia, 1995/1996 34 8. Weekly degree day totals and monthly averages, SBS site, Horsefly, British Columbia, 1995/1996 35 9. 1995, 1996, and 20 year average precipitation, BG sites, Junction study area, BC 42 10. 1995, 1996, and 20 year average precipitation, SBS site, Horsefly area, BC 42 11. Percent soil moisture of each plant association, BGxw site, Junction Sheep Range Park, British Columbia, 1995/1996 43 12. Average dates for initiation of flowering for each phenological group, BG and SBS sites, Junction Sheep Range Park, and Horsefly, British Columbia, 1995/1996 49 13. Phenologies of selected species in the Early Spring and Late Spring Phenological groups, Bunchgrass and Sub-Boreal Spruce sites, British Columbia, 1996 52 14. Phenologies of selected species in the Early Summer and Mid Summer Phenological groups, Bunchgrass and Sub-Boreal Spruce sites, ix British Columbia, 1996 53 15. Phenologies of selected species in the Mid Summer and Late Summer Phenological groups, Bunchgrass and Sub-Boreal Spruce sites, British Columbia, 1996 54 16. Graphs of upland and riparian area phenologies with group flowering and fruit dispersal initiation dates, BGxw site, Junction Sheep Range Park, British Columbia, 1995/1996. 65-66 17. Graphs of upland and riparian area phenologies with group flowering and fruit dispersal initiation dates, SBS site, Horsefly, British Columbia, 1995/1996 78-79 Appendix Figures: A.2.1 Tree diagram from clustering of riparian indicators, BGxw site, Junction Sheep Range Park, British Columbia, 1996 133 A.2.2 Tree diagram from clustering of upland indicators, BGxw site, Junction Sheep Range Park, British Columbia, 1996 134-135 A.2.3 Tree diagram from clustering of riparian indicators, SBS site, Horsefly, British Columbia, 1996 136 A. 2.4 Tree diagram from clustering of upland indicators, SBS site, Horsefly, British Columbia, 1996 137-138 x Acknowledgements I would like to thank my research supervisor, Dr. Michael Pitt, for his patience and encouragement through this entire project. Dr. Pitt has the rare ability to guide but not direct his students so that they learn how to make decisions for themselves and to correct their own errors. Thanks also to my committee members, Peter Jolliffe and Peter Marshall for their time and interest throughout the process. I would also like to thank Val LeMay for her guidance in the initial stages of my data analysis. A note of thanks to Matt Fairbarns for the initial idea for this study and for his early guidance in the field. My sincere thanks to Fred Knezevitch and Anna Roberts for their time. Fred shared his knowledge of grazing practices and his years of observations on the results of grazing in the Cariboo Chilcotin. Anna spent long hours helping me identify the sedges and rushes of the riparian areas we both appreciate. Thank-you also to Francis Njenga for his constant help with my computer problems and to Njenga and Tracey Hooper for acting as sounding boards for my ideas. Finally, I want to thank my family and and especially my husband Jurgen Hornburg. They encouraged me throughout even though my preoccupation with this project left me less time to spend with them. This research was funded in part by the Grazing Enhancement Program. xi 1. Introduction The Range Management Guidebook (B.C. Ministry of Forests 1995a) describes the objectives that must be met in any range use plan including desired plant community objectives, riparian area objectives, and landscape level biodiversity objectives. A strong emphasis is placed on ensuring the conservation of biological diversity when managing resources including range in British Columbia. An ecosystem management approach has been taken that is based on providing suitable habitat conditions for all native species. In this way, habitat diversity is used as a surrogate to maintain biodiversity (B.C. Ministry of Forests 1995c). An important part of any range management plan is denning range readiness for an area (the time when grazing can begin). The criteria used to determine readiness must prevent damage to the vegetation and soils of all habitats within the range site to maintain habitat diversity. To do this successfully requires knowledge about the ecosystem of the range area. The establishment of a sustained livestock industry dependent upon natural vegetation must include ecological, phenological, and physiological studies of individual species, and comprehensive knowledge of plant community dynamics in response to grazing ( Klemmedson et al. 1978, Kinsinger 1979). In the Cariboo-Chilcotin, both grassland and forest ecosystems are commonly used for grazing cattle. Riparian areas are common in both ecosystems. Riparian areas are one of the most productive habitats in British Columbia (Van Ryswyk et al. 1992). Although they may cover only a small portion of the total landscape, there is growing awareness of how important they are to the health and diversity of ecosystems. Riparian areas frequently contain 1 the highest number of plant and animals species found in forests, and provide critical habitats, home ranges, and travel corridors for wildlife. There are no other landscape features within the natural forest that provide the natural linkages of riparian areas (B.C. Ministry of Forests 1995b). These same areas are used extensively by livestock supplying water, shade, and forage (Bunnel et all992). Until recently, range management plans in the Cariboo-Chilcotin were designed to maintain key species in upland communities (Knezevich, Cariboo Forest Region Range Reference Area Agrologist pers. comm.). Little consideration was given to riparian communities, despite the essential role they play for native wildlife and habitat diversity and their often heavy use by livestock. The Riparian Management and Range Management guidebooks now ask that range use plans be designed to meet riparian and upland objectives for plant community maintenance. The upland communities in both forest and grassland ecosystems present further habitat complexities. In the grasslands, habitat variations are associated with various levels of soil moisture found in the swales common to this area. In the forest range, the habitat variation is associated with levels of tree canopy cover. To maintain the diversity of upland habitats and riparian areas being used as range requires information on how plant growth and range readiness in the various habitats are related. The phenologies of species in a community have been used by range managers as one way to indicate range readiness (Bertiller et al. 1990). Phenology is the study of (1) the seasonal timing of life cycle events, (2) the causes of their timing related to biotic and abiotic forces, and (3) the interrelation among phases of the same or different species (Lieth 1974). 2 The Range Management Guidebook (B.C. Ministry of Forests 1995a) suggests managers choose range readiness criteria based on plant phenology and soil conditions for each area. Despite the Guidebook's suggestion to base readiness criteria on the phenology of plants in the area, almost no information is available to the manager (except local knowledge) on how phenology may differ in the various habitats that make up a range site. This study was designed to track phenology along soil moisture and tree canopy coverage gradients to clarify and describe these differences. This information can then be used to choose readiness criteria that protect vegetation and the diversity of the varied habitats that make up grassland and forest range in the Cariboo-Chilcotin. 1.1 Literature Review 1.1.1 Uses of Phenology to Understand Plant Communities An understanding of plant phenology is valuable in planning the protection of vegetation under forage use (Bertiller et al. 1990). There are several studies that describe phenological patterns of grassland communities (Blaisdell 1958, Dickinson and Dodd 1976, Sauer and Uresk 1976). Other studies focus on a few or single species common to grasslands (Rice 1950, Quinton et al. 1982, Stout et al. 1981, Bertiller et al. 1990). The life cycles of grassland plants have been used by several authors to group plants into phenological groups to better assess the quantity and quality of forage plants (Tisdale 1947, Pitt and Wikeem 1990, Dickenson and Dodd 1976). These three studies along with Sauer and Uresk (1976) also discuss the effects of varying levels of precipitation and soil moisture on the phenology of grassland plants. Budd and Campbell (1959), Reid (1979), and B.C. Ministry of Forests (1995a) use the flowering of local flora to indicate range readiness. 3 The timing of range readiness varies from year to year and its annual determination is important to the range manager (Reid 1979). A number of models have been explored to quantify the seasonal progression of plants as a function of environmental factors. Most of them indicate that there are a number of important abiotic factors, but air temperature is one of the strongest regulators (Blaisdell 1958, French and Sauer 1974, Sauer 1978, Sparks and Carey 1995, Diekmann 1996). Growth is usually related to air temperature in terms of growing degree days (average daily temperature above a base temperature) or a similar temperature summation unit. Frank and Hofmann (1989) state that plant growth stages can be predicted from accumulated growing degree days and growing degree days alone could be used for predicting grazing readiness. Other studies point out that although temperature is important, the phenology of a plant results from interactions between the genetics of the plant and a variety of environmental factors including air and soil temperature, soil moisture, day length, and insolation (French and Sauer 1974, Rathcke and Lacey 1985, Salisbury and Ross 1992, White 1995, Sparks and Carey 1995). Individual species in a community vary in their response to a variety of climatic factors depending on the phenological stage they are in, and this variation is difficult to predict unless all climatic factors are known (Blaisdell 1958, Dickinson and Dodd 1976, Sparks and Carey 1995, Diekmann 1996,). Climatic factors are usually known only for large geographical areas and there can be wide variation in the microclimates of the individual habitats that make up each area. There has been some work done to describe the effects of microclimate on the phenology of plants. Ratcliff and Turkington (1989) found that the phenologies of some species of alpine plants were affected by aspect and others were not. Jackson (1966) looked at herbaceous plants, shrubs, and trees growing in a wide variety of microclimates found in an open field and deciduous forest in Indiana. She found species of spring flowers whose first day of flowering varied up to eleven days from one site to another depending on microclimate. Bertiller et al. (1990) studied the effects of topography on the phenology of Festuca pallescens. They report that F. pallescens shows delayed growth in areas with cooler temperatures and more humid conditions. French and Sauer (1974) describe this delay in flowering related to temperature and humidity of microclimates also. Because of the number of biotic and abiotic factors involved, Wang (1960) recommends that a system for prediction of flowering or other phenological stages should include several environmental factors (temperature, soil moisture, day length, insolation, and wind) taken directly at the plants (microclimates), and that this system must allow for the non-linearity of plant-environmental relationships. Such a system would require sophisticated time-consuming measurements of several environmental factors (Diekman 1996). Several studies have shown that despite variation in individual plant phenologies from year to year, that species were relatively constant in their flowering patterns with respect to the order in which species flowered (Anderson and Schelfhout 1979; Pitt and Wikeem 1990). White (1979) states " the date that a selected index species flowered was the simplest and probably the most practical method of estimating flowering dates if a one day loss of accuracy was acceptable." 1.1.2 Range Management in Uplands and Associated Riparian Areas Range readiness is defined by the Society for Range Management as the stage of plant growth at which grazing may begin without permanent damage to soil or vegetation (Reid 1979). Definitions of range readiness in B.C. are based on a number of factors but usually emphasize the level of soil moisture and the phenological stage and/or height of key indicator species in upland areas found in both grassland and forested range (B.C. Ministry of Forests 1995a). In the areas of the grasslands in B.C. that are grazed for the entire season, turnout occurs when the soil is firm enough to hold the weight of animals without damage and key species are at certain stages of development such as: bluebunch wheatgrass (Agropyron spicatum) has 15-18 cm (6-7 in.) of new growth, rough fescue (Festuca altaica) has 20 cm, needle-and-thread (Stipa comata) has about 12 cm (5 in.) of new growth, Sandberg's bluegrass (Poa secunda) and Junegrass (Koeleria macrantha) are flowering, and balsumroot {Balsammorhiza sagittata) is in bloom (B.C. Ministry of Forests 1995a). If cattle will not remain in the area all season the bunchgrass areas are grazed in early spring and then not until fall to allow the bluebunch seed to mature. In these areas, range readiness is determined by the soil's ability to withstand the trampling of cattle moisture (Fred Knezevich pers. comm.). Nordstrom (1984) stated that forested range is made up of two types: (1) herb and shrub dominated forest range in early successional forest ecosystems (e.g. clearcuts and burns.) and (2) tree dominated forest range in late successional and climax forest ecosystems According to guide books used by range managers in B.C., forest ranges are ready for grazing when pinegrass (Calamagrostis rubescens) has 10 cm (4 in.) of new growth or Kentucky bluegrass (Poa pratensis) has 8 cm (3 in.) of new growth (B.C. Ministry of Forests 1995a and Reid 1979). The guidelines for range readiness for forested range do not distinguish between types of forest and it is not clear to which type the height and phenological stages listed in the Range Management Guidebook (B.C. Ministry of Forests 1995a) relate. The valuable role that healthy riparian ecosystems play in regional diversity of plant and wildlife communities is just beginning to be recognized (Schultz and Leininger 1990). Skovlin (1984) summarized this role: It provides vegetation that furnishes food, cover, and nesting habitat for a multitude of terrestrial wildlife. The vegetation provides an ecotone of diverse structure and edge mosaic for a productive fauna adjacent to their necessary water. For fish and other aquatic species, it can furnish shade and cover for buffering microclimate and water temperature and is a major source of nutrients to the aquatic system. The riparian zone reduces the amount of sediment and other pollutants that enter streams, and the vegetation stabilizes channel banks. Riparian areas support the greatest number of wildlife species within the grasslands of the Cariboo-Chilcotin, and Blue- and Yellow-listed species use riparian areas more than any other type of habitat (Hooper and Pitt 1995). Riparian areas were the most frequently listed habitat of concern by respondents to interviews conducted as part of the Problem Analysis for the Chilcotin-Cariboo Grassland Biodiversity (Hooper and Pitt 1995) for a number of reasons including: • they are rare, often small, and not very resilient • they provide essential habitat • they are affected by a number of factors, including livestock grazing and watering. Studies of cattle behaviour show general over use of riparian areas at various seasons depending on the quantity and quality of the forage of the associated uplands when compared to use of the associated forested or grassland areas. Roath and Krueger (1982) in a study done in forested mountainous range in the southern Blue Mountains of northern Oregon found the riparian meadows covered 2% of the summer range, produced 21% of the total herbaceous biomass, yet provided 81% of the forage consumed over a two-year period. Marlow and Pogachik (1986) compared cattle use of riparian areas (aspen, aspen/willow, bog, meadow, and stream bank vegetation) and uplands (grassland, sagebrush, and aspen park vegetation) in Montana at 1400-2000 m and found that cattle spent a disproportionate amount of their feeding time in the riparian zone during late summer and early fall. Bryant (1985) found that livestock in the Blue Mountains of northern Oregon made between 12 and 20 times the use of riparian zones than would be expected by chance during the first third of the summer. Elmore (1989) found that if riparian areas associated with grassland in central Oregon are used for grazing during seed ripe stage of the grassland, the riparian area is utilized at 80-90% while uplands are at 60% utilization. Riparian areas including stream edge, sedge/grass meadows, fens, and shrub carrs are common in both grasslands and forests and make up about 10% of the land area of the central interior of B.C. (Nordstrom 1984). About half of the livestock feed supply of the Cariboo-Chilcotin is produced on riparian areas (Van Ryswyk et al. 1992). Riparian use in the Chilcotin grasslands is dictated by water levels. Stream banks with solid soils may be grazed season long. A seasonally inundated area will not be used until water subsides. In forested range in the Cariboo-Chilcotin, the wetlands are not usually grazed until summer or fall and some not until freeze-up. The level of use will range from 0-90% in both ecosystems depending on both quality and quantity of vegetation in the riparian area and surrounding uplands (Knezevich pers. comm.). Cattle directly impact riparian vegetation in three ways: (1) compaction of soil, which increases runs off and decreases water availability to plants, (2) herbage removal, which allows soil temperatures to rise and (3) physical damage to vegetation by rubbing, trampling and browsing (Severson and Boldt 1978). Direct effects of these impacts on the riparian 8 vegetation are not consistent, which may result from differences in plant species, environmental conditions, and livestock concentrations found in various types of riparian areas (Platts and Raleigh 1984, Clary 1995). Some general if not uniform effects, however, have been reported. First, riparian woody species are impacted negatively showing altered size, shape, and volume (Carothers 1977, Knopf and Cannon 1982, Kovalchik and Elmore 1991, and Green and Kaufmann 1995). Secondly, grazed riparian areas show an increase in the number and cover of forbs and weedy species including Kentucky blue grass (Kaufmann et al. 1983, Schultz and Leininger 1990, Popolizio et al. 1994, Clary 1995, Green and Kaufmann 1995). Large sedges (Carex rostrata and Carex nebrascensis) can show a decrease in phytomass in areas where the growing season is short (Pond 1961, McClean 1979, Kaufmann et al. 1983) or are not affected by grazing in others (Platts and Nelson 1989, Schultz and Leininger 1990, Allen and Marlow 1994). Litter increases when grazing is discontinued in riparian areas (Schultz and Leininger 1990, Popolizio et al. 1994). Finally, the phenology of riparian species can be up to two weeks later in ungrazed areas (Kaufmann et al. 1983). Despite the recognition of the importance of maintaining the diversity of riparian ecosystems included in areas used for range, management continues to focus on range readiness in terms of the main range resource of upland ecosystems (Platts and Raleigh 1984, Skovlin 1989, Knezevich pers. comm.). The Range Management Guidebook (B.C. Ministry of Forests 1995a) does list the readiness level of a few common riparian species but these species are not considered when assessing range readiness each spring (Knezevich pers. comm.). 9 Information about the phenology of plants is recognized as being essential to understanding an ecosystem (French and Sauer 1974, Dickenson and Dodd 1976, Bazzaz and Parish 1979) and is used by range managers to determine range readiness. I could find little in the literature, however, that relates either the phenology or range readiness of upland species (grasslands or forested range) with the phenology or readiness of species found in the associated riparian areas. There is also little information on the relationship of phenological patterns of species found growing under the various levels of canopy cover (insolation) found in the forested range of B.C. Without this information, it is difficult to achieve prescribed management goals for range use that simultaneously allow for the maintenance of diversity of the various habitats within a range area. This study was designed as a case study and intended to be descriptive in order to provide more information on these relationships. Because of the importance of and pressures on riparian habitats imposed by grazing, I chose to focus on the phenological patterns found across a wet/dry gradient in both grassland and forested range. In addition, phenological patterns related to canopy closure were examined in the forested range, to allow for the two types of forested areas (herb and shrub dominated and tree dominated) used as range. Objectives The objectives of this research were: a. To determine how yearly changes in climate affect phenology at sites in the Bunchgrass and the Sub-Boreal Spruce Biogeoclimatic Zones of the Cariboo-Chilcotin region of B.C.; 10 b. To clarify how phenology varies among plant associations which are related to soil moisture gradients in grasslands; c. To clarify how phenology varies among plant associations which are related to soil moisture and canopy cover gradients in forested range; d. To describe and compare the phenological patterns found in riparian areas and surrounding uplands at sites in the BG and SBS; e. To identify species that are consistently good indicators of the phenological progression found in the uplands and associated riparian areas and indicators of range readiness; f. To integrate the phenological patterns of individual plant associations into a description of the phenological progression of a site using these indicators; and g. To make range management suggestions based on the observed phenological patterns to help maintain the diversity of the plant associations found in the various habitats within the grassland and forest ecosystems 2. Study Area I chose two study areas located in two biogeoclimatic zones in the Cariboo-Chilcotin region of British Columbia (BC). One area is located in the grasslands of the Junction Sheep Range Park (Junction) (51° 51' N , 122° 21' W) west of Williams lake and the Fraser River (Figure 1). These grasslands are at the northern limit of the Bunchgrass (BG) biogeoclimatic zone (Roberts 1992). The second area is in a section of forest south of Horsefly (52° 13'N, 121° 29'W). This area is located on the southeastern boundary of the Sub-Boreal Spruce (SBS) biogeoclimatic zone (Figure 1). 11 FIGURE 1. Location of study areas in British Columbia, 1995/1996. 12 2.1 Bunchgrass Study Area The climate of the Bunchgrass zone is characterized by warm to hot dry summers and moderately cold winters (Nicholson et al. 1991) with a 120 day frost-free period. Annual precipitation averages 300 mm and is usually bimodal with a peak in December and January in the form of snow and another in June (Nicholson et al. 1991). The 20 year averages for precipitation at the Junction do not show this peak in June; precipitation averages 38 mm each month, May through August (Environment Canada 1973-1993). Springs can be dry, and the summer precipitation does not always recharge the soil moisture. Depletion of soil moisture can stress plants as summer progresses (Williams 1983). This precipitation pattern also affects the associated riparian areas where water levels vary considerably from year to year (Roberts 1992). The soils of the Junction's grasslands are Brown, Dark Brown, Black, and Dark Gray Chernozems (Nicholson et al. 1991). The soils have a thick (>15 cm) Ah horizon with generally good drainage except in shallow depressions or at the base of cliffs where wetland communities may occur (Nicholson et al. 1991). There are two grassland subzones- the Very Dry Hot (BGxh) found from valley bottom to about 700 m and the Very Dry Warm (BGxw) between 700 and 1000 m. The vegetation of the BG zone consists mainly of widely spaced bunchgrasses and lichen species that encrust the soil surface, but the vegetation also reflects changes in topography, aspect, and drainage (Nicholson et al. 1991). Moist depressions (swales) are common and are characterized by a greater variety of forbs and grasses and/or groves of trembling aspen (Populous tremuhides) associated with western snowberry 13 (Symphoricarpos occidentalis) (Nicholson et al. 1991). The Junction was grazed by domestic cattle until 1978 when it became a protected area. The first site located in the Junction was within the Bunchgrass Very Dry Warm subzone (BGxw site) at 850 m. Bluebunch wheatgrass, pasture sage (Artemisia frigida), and Junegrass dominated the dryer parts of the uplands of this subzone. Deep swales with stands of Douglas-fir (Pseudotsuga menziesii) and trembling aspen trees were common. The associated riparian area was an awned sedge (Carex atheroides) fen marsh association (Steen and Roberts 1988). A baltic rush (Juncus balticus)-fie\d sedge (Carex praegracilis) meadow surrounded it. The second site in the Junction was located at 700 m in a transition between the BGxw subzone and the Bunchgrass Very Hot Dry subzone (BGxh). I will refer to this as the BGxw/h site. The same species dominated the uplands here as at the BGxw site, but cover was less dense and there was more bare ground. Deep swales were rare with only one or two large Douglas-fir trees growing in them. The same type of riparian area was associated with these uplands. 2.2 Sub-Boreal Spruce Study Area The Sub-Boreal Spruce biogeoclimatic zone is located from valley bottom to 1100-1300 m elevation over a latitudinal range of 51° 30' to 59° N and has a continental climate with seasonal extremes of temperature, severe, snowy winters, and relatively warm, moist and short summers (Meidinger et al. 1991). Annual precipitation can range from 440 to 900 mm, and the spring and summer portion is distributed evenly during the growing season (Meidinger et al. 1991). The frost-free period averages 60 days (Lord 1984). This biogeoclimatic zone is 14 dominated by upland coniferous forests made up of hybrid white spruce (Picea englemannii x glauca), subalpine fir (Abies lasiocarpa), lodgepole pine (Pinus contorta var. latifolia), trembling aspen, paper birch (Betulapapyri/era), and Douglas-fir (Meidinger et al. 1991). The study site in this area was located in a section of forest at 1050 m in the Horsefly Forest District. This area is within the dry warm subzone of the Sub-Boreal Spruce biogeoclimatic zone (SBSdw). The most common soils of this subzone are Podzolic Gray Luvisols (Lord 1984). Up to 8 cm of forest litter commonly occurs on these soils, but there is little or no incorporation of this organic matter into the mineral soil such that no Ah horizon exists (Valentine et al. 1978). Because clay accumulates in the B horizon, water penetration is restricted and these soils are very wet in spring (Valentine et al. 1978) The uplands of this area were dominated by lodgepole pine, hybrid white spruce, and trembling aspen. There was a well developed but less dominant shrub and herb layer. Riparian areas make up 4% of the total land in the SBS zone and consist of bogs, fens, and marshes. The riparian areas associated with the forested uplands of this site were fens, (wetlands that have accumulations of non Sphagnum peat, are less acid, and are more mineral rich than bogs). There were three types of fens within this Sub-Boreal site (SBS site) with a variety of plant associations. One was a shallow mesic fen located at the edge of a large (2 ha) shallow open water that was part of a linked basin system (Runka and Lewis 1981). This fen had two associations: a slender sedge-moss fen association (Carex lasiocarpa-Drepanocladus aduncus); and, a beaked sedge-water sedge fen association (Carex utriculta-Carex aquatilis) (Steen and Roberts 1988). A floating mesic fen (Runka and Lewis 1981) with a buckbean-slender sedge fen association (Menyanthes trifoliata-Carex 15 lasiocarpa) (Steen and Roberts 1988) also occurred in this site. There was also a shallow mesic fen (Runka and Lewis 1981) surrounding open water in a closed basin. This site has been grazed by domestic cattle periodically over the last 100 years. It was not grazed the two years of the study. 3 Methods 3.1 Site Selection and Establishment of Plots I chose sites in both areas to represent the varied plant associations and to express the vegetation changes across a wet-to-dry gradient associated with a riparian area. The two sites in the BG zone reflected the most common variation in vegetation related to changes in topography. In the SBS zone, variation in vegetation was related more to the level of canopy cover (insolation), and this site included three levels of insolation common to this zone. To allow for these variations, the sites had to be large (1-3 ha) and irregularly shaped. At each site in the BG zone, I differentiated and mapped seven different plant associations related to soil moisture levels from the centre of the riparian area (wetlands) to the uplands (Figures 2 and 3). Except for the swales, the plant associations encircled each riparian area. The swales ran generally perpendicular to each wetland basin. I staked ten plots each two metres on a side within each plant association. To average the effects of aspect, the plots were spaced evenly along the centre line of each association on all sides of the wetland basin. At the site in the SBS zone the upland forested area had three types of fens associated with it as described earlier, and each had a different plant association surrounding it. To allow for this variation, I established plots each two metres on a side in each of the 16 associations surrounding the three fens. Two of the riparian associations were only large enough for five plots. To be consistent in the riparian associations, I placed five plots in each of the riparian associations. For the forested uplands, I included a path that ran through the forest and a nearby one-hectare clearcut along with the general forest cover to compare the effects of the different levels of tree canopy closure in the uplands. Ten plots were staked in the clear cut and 10 in the forest. I sampled the path along its length to include the diverse but in some cases uncommon species present (Figures 4 and 5). This choice of site and inclusion of path and small clearcut allowed for sampling the varied forest habitats related to canopy closure and soil moisture used by cattle in the SBS zone (Nordstrom 1984). 3.2 Vegetation Composition and Coverage Herb and shrub vegetation (<10 m) was sampled along 30-m transects placed in the middle of each plant association. A plot frame 20 x 50 cm was placed at three-metre intervals along each transect starting at 0 (10 samples per transect). Because the wetlands differed in size, it took from two to four transects per plant association to sample on all sides of each wetland. I recorded the cover of each species and the amount of bare ground using classes based on Daubenmire (1959): present -1% (1); >l-5% (2); >5-25% (3); >25-50% (4); >50-75% (5); and >75% (6). Each species' average cover per transect was combined with the other transects in that association to give a percent cover class for each species in each association. I visually estimated the cover of cryptogams for each association. I sampled the BG zone in mid June 1996 and the SBS zone in mid July 1996. Plant taxonomic and common names were taken from Douglas et al. (1989, 1990, 1991) and Meidinger (1987). 17 AWW5tVGL A^OCIATION orotic field rush, eedaz/ • wetter m m \v\\rwr&\ edit clay Legend 1 Open water association 2-4 Wet riparian, dry riparian, and transition associations 5 Dry grassland association 6 Shallow swale association 7 Deep swale association FIGURE 2. Side and top view, BGxw site, Junction Sheep Range Park, British Columbia, 1995/1996. 18 62 arctic f j 'eid <?idqej ruzh; giant rye/ •Y.Tir.r.r. a water organic (feM.) clay (c 9<F Aspen Legend 1 Open water association 2-4 Wet riparian, dry riparian, and transition associations 5 Dry grassland association 6 Shallow swale association 7 Deep swale association FIGURE 3. Side and top view, BGxw/h site, Junction Sheep Range Park, British Columbia, 1995/1996. 19 WATEFi SEP2E> > 1.0 AA r r~| mater ^ organic (peat) | M clay FIGURE 4. Side view, SBS site, Horsefly, British Columbia, 1995/1996. 20 for&sk 4 OEAKEP SEDGE--WAT5& SVPGZ.-FZv\ AeZCClAjlON f lowing m A^OdATION Legend 1 Open water association 2 Riparian association 3 Open edge association 4 Forest association 5 Path association 6 Clearcut association FIGURE 5. Top view, SBS site, Horsefly, British Columbia, 1995/1996. 21 Tree canopy cover (>10 m) in the SBS zone was the combined coverage of the vertically projected crowns of the trees in the canopy layers on the forest floor (Luttmerding et al. 1990). In mid-July, I recorded the number of hours per day each association was shaded by the tree canopy (Table 1). Table 1. Hours herb and shrub layers not shaded, SBS site, Horsefly, BC, 1996 Area Tree canopy coverage (%) Hours not shaded Centre of wetland 0 12 (constant) Clear cut 0 12 (constant) Wetland edge 10-20 6-8 (constant) Path 10-20 3-6 (intermittent) Forest 40-50 1-2 (intermittent) 3.3 Abiotic Measurements I measured soil moisture gravimetrically at each site at approximately two-week i intervals from April to August in 1995 and 1996 . Each sample was an aggregate made up of ten individual samples collected along a transect placed at the centre of each association. I collected soil samples from soil at 0-10 cm and 10-20 cm depths at regular intervals along the transect starting at a different point each time. The aggregate samples were sealed in a bag, weighed, unsealed, and air-dried at 25° C until the weight was constant. I calculated the moisture content in terms of percentage of air-dried soil. I recorded the height of the standing water at the centre of each riparian association using a metre stick and did not measure the soil moisture until the standing water had receded. 22 I measured precipitation at each study area using a standard rain gauge from April to August. Precipitation for the rest of the year was based on the nearest Environment Canada Weather Station (Wineglass, 20 km from the B G sites and Olchitree, 23 km from the SBS site). Both Diekman (1996) and White (1979) compared different methods of relating meteorological measurements to the phenology of plants and concluded that the sum of growing degree days was the most practical to use with very little loss of accuracy. Degree days is the cumulative sum of average daily temperatures above a threshold base temperature from a chosen starting day (January 1 in this study). Environment Canada uses 5° C as a threshold temperature usually in reference to agricultural crops. Diekman (1996), Caprio (1974), and Frank and Hofmann (1989), however, used 0° C as the base temperature for their studies. I used a 0° C base temperature, because it allows for the growth that can take place in the warmed soils of late spring even if there are low air temperatures. In 1995,1 used the temperature data from the nearest weather stations to provide the daily minimum and maximum temperatures for each site. In order to have more exact temperature data for each site separately, I installed Hobo Temp Temperature Loggers (Onset Computer Corporation, Pocasset, Ma.) at each site 1.5 m above the ground in 1996. From April 15 to September 30, each logger recorded the temperature four times daily. I used these data for the daily minimum and maximum temperatures for 1996. I wanted to be able to compare community and species phenology between sites and between years and to relate this to the degree day totals. I, therefore, altered the 1995 weather station degree-day data by the same percent differences I found between the 1996 site and station data. The BGxw site's 23 weekly degree totals were 85% of Wineglass totals from April 15- July 21 and 83% from July 22- September 30. The BGxw/h site's totals were 88% and 86% of Wineglass totals for the same time periods respectively. The SBS site's totals were 120% of Ochiltree totals from April 15- May 31 and the same as Ochiltree totals from June 1-September 30. 3.4 Phenological Data I collected phenological data on a weekly basis from the last week in April until September 7 for a total of eighteen weeks in 1995 and 1996. By checking all plots within a plant association, I sampled ten or more plants of each species and determined a modal phenological stage for each species in an association. Some species were so rare, however, that they did not occur in enough plots, and I would check for these species throughout each association. It was often difficult to determine generative stage without handling plants or collecting and examining the plant using a microscope. To avoid damaging plot plants, I examined and collected samples at the modal stage of development from outside the plot. The 1996 data are generally more accurate and contain more species, because I was better able the second year to recognize early stages of more plants. I began by using the phenological codes for herbs and grasses from Luttmerding et al. (1990) which were adapted from Dierschke (1972). During 1995,1 altered these codes slightly (Table 2) to better describe the stages I was observing and to have a common phenological code for all vegetation forms (herbs, grasses, and shrubs). 24 Table 2. Phenology codes used at B G and SBS sites, Junction and Horsefly study areas, British Columbia, 1995/1996 Vegetative Generative 0 Without new shoots above ground 0 Without blossom buds or closed buds 1 Shoots without unfolded leaves 1 Inflorescence recognizable or buds with green tips 2 First leaf unfolded 2 Buds swollen and/or visible 3 2 or 3 leaves unfolded 3 Flower visible but not open 4 Most leaves unfolded not full size 4 Flower open 5 Almost all leaves unfolded range 5 Up to 25% pollenizing in size or in blossom 6 Growing and adding size and leaves 6 Up to 50% pollenized 7 Plant fully developed 7 Full Bloom 8 First leaves turning yellow 8 Fading inflorescence 9 Yellowing up to 50% 9 Fully faded inflorescence and early fruit 10 Yellowing over 50% 10 Bearing green fruit 11 Dead or dormant 11 Bearing green and ripe fruit 12 Regrowth 12 Fruit dispersal 13 Fruit dispersed 3.5 Statistical Analysis I used the SYSTAT (1990) hierarchial clustering program to identify groups of species in each association with similar phenological patterns. The first step in this process is to calculate the similarity or dissimilarity between all pairs of samples to produce a (dis)similarity matrix. The samples are then sorted into groups based on the information in the data matrix (Kent 1992). There are a variety of choices for the measure of (dis)similarity and for the sorting technique depending on the form of the data. I used Euclidean distance as the measure of (dis)similarity. Despite the concern expressed by Everett (1993) that Euclidean distance can be a poor measure of 25 similarity/dissimilarity when variable values are correlated with one another as they are in phenological data, the resulting clusters did group species with similar phenologies. Ward's minimum variance (Ward 1963) was used as the sorting procedure, because it is less likely to include outliers and forms tighter groups (Gauch 1991). This was important because each site included some species with unique phenologies which could have affected the groups formed by the clustering process. The hierarchial clustering was useful in both clarifying groups of species with similar phenology and indicating how these groups related to each other through the growing season. This clustering was done first using each species' generative phenology. The vegetative phenology varied very little and did not allow for meaningful clusters. Using the generative phenology did produce clusters, but the resulting clusters were determined by very detailed characteristics of flowering (e.g., 25% pollenizing, up to 50% pollenizing, fading, and fully faded) which are difficult to assess for a large number of species in the field. Equally important, the generative code did not allow for early season variation in phenology, which is usually indicated by vegetative phenology, to affect the clustering. I therefore amalgamated the vegetative and generative code into a combined code that allowed a more complete assessment of a species' phenology over the growing season in only one set of variables (Table 3). The resulting clusters grouped species into phenological groups that better reflected the complete phenology of each species. Hierarchial clustering grouped species with very similar phenologies well, but larger groupings were not as accurate especially in terms of which groups species with unusual phenologies (e.g., long flowering periods, evergreen plants) were included in. Using cluster 26 information and deciding which groupings represent a common pattern (of phenology in this case) is a subjective decision which relies on the knowledge and experience of the user (Kent 1992). I felt it was important to have species grouped together that were similar in growth stages that are easy to recognize in the field. I decided to focus on flowering stages as did Dickenson and Dodd (1976) who stated that "Even if time of flowering can be a poor indicator of vegetative activity or fruit dissemination, there are few other criteria that may be used easily and consistently to note phenological similarity." I used the 1996 phenological data to order the species of each plant association by the date of initiation of flowering (stage seven) and produced ordered data matrixes of all species found in each plant association at each site. In most cases, the groups thus formed were similar to those formed in the hierarchial clustering process for each association except for species with unusual phenologies. The species were placed in the same order in the 1995 data matrixes as they were in the 1996 matrixes to allow for comparison of flowering order between years. The ordered matrixes made it possible to identify phenological groups in each association and to compare groups between associations and between years. The cluster tree diagram (Figure 6) and ordered data matrix of the BGxw dry grassland association for 1996 (Table 4) are included here as an example. The species are listed in abbreviated form followed by the number of the association in which they are being recorded (species list and codes are in appendix three). The flowering stages (seven, eight, and nine) are shaded in the matrix which emphasizes flowering periods and highlights flowering order each year. Ordered data matrixes for each plant association are included in Appendix one. 27 Ordering by flowering time allowed for inclusion of these species into appropriate groups based on the phenological stages that are easiest to recognize in the field. Species with patterns that could place them in more than one phenological group were placed in the group that emphasized the similarity of their flowering periods. The final group membership was based both on the placement of species during the clustering process and where the species appeared in the ordered data matrix. 28 Table 3. Combined phenological code used for statistical analysis, Bunchgrass and Sub-Boreal Spruce study areas, British Columbia, 1995/1996 Combined code Equivalent vegetative Equivalent generative code code 0 No new growth 0 0 1 First shoots showing 1' 0 2 First leaves unfolding 2 0 3 2-3 leaves unfolded 3 0 4 Middle leaves open; 4 0 adding leaves and size 5 0 5 Inflorescence recognizable; 6 1-2 adding leaves and size 6 Mature flower buds; 6 3 plant nearly full size 7 10-30% in bloom; 7 4-5 plant full size 8 >30% in flower; leaves 7-8 6-7 may just start to yellow 9 Flowers+faded flowers+ 7-8 8-9 green fruit 10 Green fruit; plant 8-9 10 up to 50% yellow 11 Ripe fruit; plant 8-9 11 up to 50% yellow 12 Dispersal and/or plant yellow>50% 10 12 13 Fruit dispersed; plant 11 13 dead or dormant 14 Re-growth 12 13 29 erigcom5 ta r a o f f 5 andrsep5 arabsp5 lomamac5 arabhol5 antedim5 ^ a r e p e t 5 ^ , r ~\ zigaven5 penspro5 l i t h r u d 5 geumtri5 anemmulS e r i g l i n 5 anteparS sedulan5 V. J /calomac^i o r t h l u t 5 erioher5 tragpra5 comaumb5 rosawoo5 galibor5 achimil5 a l l i c e r S V J FIGURE 6. -I -I I I - I Spring 0 . 816 1.446 0.471 2 . 576 E a r l y Spring 0 . 851 ( e a r l i e s t flowering) 0. 624 1. 031 9 .163 1.095 0.471 0 . 834 0.667 1.122 0. 745 1.214 27.845 2 .197 1.328 0 .667 3 .335 1. 713 0.708 0 . 624 1.600 5 . 856 Mid-Summer Tree diagram: dry grassland association, BGxw s i t e , Junction Sheep Range Park, B r i t i s h Columbia, 1996 3 0 /^agrospi 5^ cirshooS e r i g f l a 5 koelmac5 agosglaS lepidenS poasecS crepatrS stipcomS f e s t a l t S tragdub5 linuper5 \creptec5 J ^solispa5~^ artecam5 chrynau5 a r t e f r i 5 a r t e t r i 5 V J FIGURE 6 -- I + --I - II E a r l y Summer + --I 0.825 0.660 0.408 0. 667 3 .297 1.080 2 . 989 1.133 0 . 816 1.771 0 . 985 0 . 816 15.252 0 . 667 1. 872 Late Summer 1. 014 ( l a t e s t flowering) 0.577 (Continued) Tree diagram: dry grassland association, BGxw s i t e , Junction Sheep Range Park, B r i t i s h Columbia, 1996 3 1 IS Q. 0) Ol c ra 01 a. o £ w c o o c 3 s x O OQ c o +3 ra o o I/) in ra T3 c ra in in co k_ O) 2" n l m co c o C O co C M C M C M f C O C O C O C O C O C M C M C M 1 C O C O C O C M C O C M C M C M • I C O CO CM CM CO CM CM CM I I f O t O C M C M C O C M C N j ' -E *1 •a E •a = £ e at C o ? 2 (4 CD C M C O C M C M C M C M C O C M C M C O C O C M C O C M C M C M C M C M ^ - C M C O C O C M C O C M C M C M C M C M O C M C O C O C M C O C M C M C M C M C M O C M C O C O C M C O t - C M T - C M C M O C M C O C O C M C M T - C M T - T - T - O C M C M C M C O C O C M C M C O C M C M * - C M C M o - r - nm™ 9 ™ (*) C M r- — C O C M C n I O C O C M ^ - T - C O T - C M O !<M C M T - O C M C M O C M — O C M O C M O l T - O . 0 ) C M O l » - O l 3 0 0 0 ) 0 0 ) 0 1 0 ) o o o i o o o a o i c o >lo O l O) CO ( O CO 00 s >|co co r-- p— r— co co to c o c o r - c o t o m c o m Q. in a) o_ CO W ' c 2 CO CO y i n „ m i n u> | £ g E E * E -8 2 B & £ re 2 CO © « E a, § C O C O C M C M C O C M C M C O C M C M C M CM CO CM CM CM CM C M C O C M C M O C M C O C M C M C M C M C M C O C M C M O C M C M C M C M C M C M CM CO CM CM O CM CM CM CM O CM CM CO CM CM O CM CM CM T - O O C M C M T - T - a > T - O O ' - ' - O O O I ' - O I N 0 0 0 > 0 0 0 0 0 ) 0 0 ) C M 0 ) 0 0 ) 0 0 0 0 ) C O t J ) W O cncnoQOGiaicoGioocn cococncncococos co co co l ^ | ^ | N . 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The total degree days for April through June, 1996, however, were only 71% of average at the BG sites. April, 1996 totals were average at the SBS site, but May and June had totals only 75% of average. July and August temperatures were average in 1996 at all sites but the degree day totals remained below average because of the low spring totals. The BGxw was consistently slightly cooler than the BGxw/h site and had degree day totals 97% of those at the BGxw/h site. The SBS degree day totals averaged only 80% of those at the BG sites in spring. July and August temperatures at the two study areas were similar both years, but the degree day totals of the Sub-Boreal Spruce site were only 85% of the BG sites both years because of the lower spring totals. 4.2 Soil Moisture and Precipitation 4.2.1 Plant Associations Related to Soil Moisture and Tree Canopy Gradients The four riparian plant associations at the BGxw site were very similar in terms of cover and soil moisture levels to the respective riparian associations at the BGxw/h site (Tables 5 and 6). Each association defined a different level of soil moisture along a wet to dry gradient. The open water at the BGxw site was deeper than at the BGxw/h site, but both basins retained some standing water in 1996, the year with average precipitation. The wet riparian associations of both sites had saturated soils (>100%) all season and standing water until 33 34 35 August in 1996. The contours of the basin of the BGxw/h riparian area were shallower than those of the BGxw site and the soils remained saturated for longer each year in the dry riparian and transition associations at this site. Soil moisture remained higher all season in the top 10 cm than at the 10-20 cm depth in the riparian associations at both sites. There were three upland plant associations at each BG site (Tables 5 and 6) related to soil moisture levels. The soil moisture levels were similar at both sites (less than two percent different) except for the deep swales. A deep Ah horizon (10-20 cm) remained moist at both 0-10 cm and at 10-20 cm throughout the growing season in 1995 and 1996 in the deep swale at the BGxw site. The deep swale at the BGxw/h had a thin Ah horizon (2-5 cm) and gravel just below this. Persent soil moisture was lower in this deep swale than in the dry grassland association both years. The shallow swale associations at both sites were more moist than the dry grassland associations throughout the growing season. The dry grassland associations which dominated each site had the driest soils (except for the BGxw/h deep swale). Levels of soil moisture remained lower at the 10-20 cm depth until mid to late summer when the levels in the top 10 cm dropped below those at the lower depth in all upland associations except for the BGxw/h deep swale. This reversed again in most upland associations after the heavy rainfalls in August in 1995 and in September in 1996. The BGxw/h deep swale, however, always was drier in the top 10-20cm. Two riparian associations and an open edge or transition association related to soil moisture levels grew at each of the wetlands that were part of the SBS site (Table 7). Each wetland had a centre of open water (the centre of the floating mesic fen had a floating island 36 Table 5. Plant associations, BGxw site, Junction Sheep Range Park, British Columbia, 1995/1996 Plant association Description Dominant species Cover % Riparian Associations 1 Open Water 2 Wet Riparian 3 Dry Riparian 4 Transition Upland Associations 5 Dry Grassland 6 Shallow Swale 7 Deep Swale 75% open water Carex exsiccata 25-50 remains all season Carex atherodes 25-50 dries to puddles by Juncus balticus 50-75 mid summer Calamagrostis stricta 5-25 standing water early Carex praegracilis 25-50 spring only, then Juncus balticus 5-25 saturated soils Agropyron spicatum 5-25 saturated soils early Carex praegracilis 5-25 spring only; Agropyron spicatum 5-25 moist to dry soils Tragopogon pratensis 5-25 in summer Aster ericoides 5-25 dominant upland Agropyron spicatum 25-50 association; soils Koeleria macrantha 25-50 dry by mid summer Stipa comata 5-25 Artemisia frigida 5-25 soils remain more Stipa curtiseta 25-50 moist than in dry Agropyron spicatum 5-25 grassland association Koeleria macrantha 5-25 Carex petasata 5-25 remains very moist Populus tremuloides 5-25 all season Pseudotsuga menziesii 5-25 grasses and forbs 25-50 Shrubs 5-25 37 Table 6. Plant associations, BGxw/h site, Junction Sheep Range Park, British Columbia, 1995/1996 Plant association Description Dominant species Cover % Riparian Associations 1 Open Water 75% open water Carex exsiccata 25-50 remains all of 1996 Carex atherodes 25-50 2 Wet Riparian dried to puddles by Juncus balticus 50-75 mid summer 1996 Calamagrostis stricta 5-25 3 Dry Riparian standing water early Carexpraegracilis 25-50 spring only, then Juncus balticus 5-25 saturated soils Agropyron spicatum 5-25 Phalaris arundinacea 5-25 4 Transition saturated soils early Carex praegracilis 5-25 spring only; Agropyron spicatum 5-25 moist to dry soils Tragopogon pratensis 5-25 in summer Leymus cinereus 5-25 Upland Associations 5 Dry Grassland dominant upland Agropyron spicatum 25-50 association; soils Koeleria macrantha 25-50 dry by mid summer Stipa comata 5-25 Artemisia frigida 5-25 6 Shallow Swale soils remain more Stipa curtiseta 25-50 moist than in dry Agropyron spicatum 5-25 grassland association Koeleria macrantha 5-25 Carex petasata 5-25 7 Deep Swale soils drain quickly Stipa comata 5-25 and are very dry S. curteseta 5-25 all season Agropyron spicatum 5-25 Koeleria macrantha 5-25 38 Table 7. Plant associations, SBS site, Horsefly, British Columbia, 1995/1996 Plant association Description Dominant species Cover % Riparian Associations 1 Open Water 75% open water a Carex lasiocarpa 5-25 (fen associations) water remains all Carex aquatilis 5-25 a-slender sedge/moss season; no shade Carex utriculata 5-25 b-buckbean/slender b Menyanthes trifoliata 5-25 sedge Drepanocladus aduncus 25-50 c-beaked sedge/water sedge c Nuphar polysepala 25-50 2 Riparian standing water a Drepanocladus aduncus 25-50 (surrounding for most of season Carex utriculata/ aquatilis 25-50 3 fens) can dry to puddles; Drepanocladus aduncus 5-25 no shade Triglochin maritimum 5-25 b Menyanthes trifoliata 5-25 Carex lasiocarpa 5-25 c Carex utriculata /aquatilis 25-50 Carex lasiocarpa 5-25 Calamagrostis canadensis 5-25 Upland Associations 3 Open edge standing water early Betula glandulosa 5-25 spring only; Lonicera involucrata 5-25 6-8 hours no shade Rosa acicularis/woodsii 5-25 Calamagrostis rubescens 5-25 4 Forest moist soils all season; Populous tremuloides 5-25 shade full day Picea englemannii x glauca 25-50 Pinus contorta 25-50 Lonicera involucrata 5-25 Calamagrostis rubescens 5-25 5 Path moist soils all season; Populous tremuloides 5-25 no shade 3-6 hours Picea englemannii x glauca 5-25 Pinus contorta 5-25 Lonicera involucrata 5-25 Calamagrostis rubescens 5-25 6 Clearcut moist soils all season Poa pratensis 5-25 no shade 10-12 hours Agrostis gigantea 5-25 Epilobium angustifolium 5-25 young trees 5-25 39 of vegetation surrounded by a band of water), and each was surrounded by a riparian zone with saturated soils and some standing water. The transition zone was narrow and affected by standing water early in the spring only. The soil remained very moist the rest of the growing season but not saturated as in the riparian zone. The upland plant associations related to a tree canopy gradient rather than wet to dry soil moisture gradient. Soil moisture levels of the three upland plant associations differed very little. All three associations were very moist in spring (averaging 42%) and showed a steady decline in soil moisture throughout the growing season. Soil moisture never went below 22%. They remained more moist in the top 10 cm than at the 10-20 cm depth the entire season. 4.2.2 Changes in 1995 and 1996 Levels of Precipitation and Soil Moisture Precipitation at the BG sites did not conform closely with 20-year averages based on data taken at the Wineglass Ranch Environment Canada Weather Station (1973-1993) in either 1995 or 1996 (Figure 9). The 1995 winter precipitation (snow) averaged only 48% of normal. Spring was even dryer with 30% of the average precipitation (April to May). Rains came in July and August and the total precipitation for these months reached 192% of the average. Precipitation for the winter of 1996 was 83% of the average. Spring had average precipitation, but summer (July and August) precipitation equalled only 47% of average. Precipitation levels at the SBS site conformed to 30 year averages in 1996 but not in 1995 (Figure 10) based on data taken at the Ochiltree Miocene Environment Canada Weather 40 Station (1963-1993). Winter 1994-1995 precipitation (snow) reached only 48% of the average. Spring precipitation (April-June) was 117% of average and summer precipitation increased to 153% of average for July and August. The lower levels of winter precipitation in 1995 than in 1996 meant that both wetlands in the BG study area had spring water levels that were less than half in 1995 compared to 1996. In 1995, the BGxw site open water went from spring highs of 50 cm to just a 6 cm depth in puddles by early July, and the BGxw/h open water went from a 30 cm depth to being completely dried by this time. The growing season of 1996, the BGxw open water decreased from a high of 100 cm to 50 cm, and the BGxw/h went from 80 cm to 45 cm by early September. Both wet riparian associations also retained standing water throughout the summer (60 cm down to 30 cm at the BGxw and 30 cm down to 8 cm at the BGxw/h). Even the dry riparian associations at both sites had standing water until June in 1996. The transition and upland associations also reflected the increased winter precipitation. They had spring soil moisture levels that were 1.5-1.9 times higher in spring 1996 than in 1995. In late July of 1996, soils dried to a level equal to that found in 1995. The deep swales did not follow this pattern. A stream ran through the BGxw deep swale until July, 1996, and the soil moisture remained high throughout the growing season. Soil moisture levels differed very little between years in the BGxw/h deep swale. Because the upland associations at the two grassland sites had very similar soil moisture levels, I have summarized only the BGxw soil moisture in Figure 11. The wetlands at the SBS site also reflected the differences in total winter precipitation each year. The standing water was four times deeper in the spring of 1996 (1.25 m) than in 41 FIGURE 11. Percent soil moisture of each plant association, BGxw site, Junction Sheep Range Park, British Columbia, 1995/1996. 1995 Percents * 43 spring 1995 and remained twice as deep the entire summer (47 cm in August). In 1995, the riparian association had only four centimetres of standing water in spring and by mid summer patches of it dried completely. It averaged 20 cm of standing water all of 1996. Two centimetres of water covered the open edge association in April, 1995 and 8 cm in April, 1996. The upland forested associations soil moisture levels were higher in 1996 than in 1995 only in spring. 4.3 Vegetation Coverage in Each Plant Association 4.3.1 Grassland Sites Vegetation coverage of the three riparian associations was very similar at the two BG sites (Tables 5 and 6). One or two Carex species dominated the open water and wet riparian associations each with a cover of 25% or more. Ten other species with covers of less than 5% (complete species lists and associations for all sites are included in the Appendix 3) grew in these two associations. The dry riparian associations were dominated by C. praegracilis but had a greater variety of species with 5-25% coverage than the other riparian associations. A number of species usually considered upland species (each with a coverage of less than 5%) grew in this association. The only bare ground in these three riparian associations appeared when the water receded and was then quickly covered with seedlings. The transition associations at the two sites had less distinct boundaries than the other riparian associations and no single species covered more than 25% of the association. Field sedge, bluebunch wheatgrass, and meadow salsify (Tragopogon pratensis) together covered 50% of this association at both sites. Most species common to the uplands grew in this association along with species common to the dry riparian association. Giant wild rye 44 (Leymus cinereus) and quackgrass (Elymus repens) covered more of this association at the BGxw/h site than at the BGxw site. The dry grassland association interrupted by narrow bands of shallow swale association covered much of the uplands of both sites. The dry grasslands dominated by bluebunch wheatgrass and Junegrass also had a variety of less dominant grasses and forbs. Bare ground with some cover of cryptogams was common (28% at the BGxw site and 30-40% at the BGxw/h site) in this association. The shallow swale dominated by porcupine grass (Stipa curtiseta) had a thick mulch of dried grass exposing very little bare ground. The mulch was thicker in this association at the BGxw site than at the BGxw/h site. Other grasses and forbs found mainly in the areas of thinner mulch or bare ground were more common at the BGxw/h site. The deep swales were very different at each site in terms of cover. Trees dominated the BGxw deep swale. Shrubs (5-25% cover) and forbs and grasses common to the riparian and upland associations made up the understorey of this association. A number of species found only rarely in the other associations including sticky purple geranium (Geranium viscosissimum), arrow-leaved balsamroot, and Canadian goldenrod (Solidago canadensis) also grew here. Less than 5% of the ground was bare. The deep swale at the BGxw/h site had large patches of bare gravelly ground. Only two large Douglas-fir trees grew in this swale. Aspens at this site grew next to the riparian areas not in the deep swale. The shrub cover was low (<5%) and the dominant grasses grew less thick than at the BGxw/h site. Needle-and-thread grass was more common than porcupine grass in the BGxw/h deep swale association. 45 4.3.2 Sub-Boreal Spruce Site Although the dominant species at each of the three fens at this site differed, one or two species dominated (covers of 25-50%) each riparian plant association of each fen (Table 7). There were a number of rarer species common to all the fens and a few like Rocky mountain cow-lily ( Nuphar polysepalum) and buckbean found at only one fen. Upland associations had a greater variety of species than the riparian associations and the understorey showed less dominance by one or two species. The same species dominated the open edge association at each fen. This association covered a two to three metre band around each fen and was dominated by a mix of shrubs. A variety of riparian and forest forbs and grasses were common. A thick moss layer covered 25% of the ground. The forest association dominated by a mixed stand of mature trees and shrubs, had a wide variety of forbs and a few grasses with cover of 5% or less each in the understorey. Cryptogams including freckled lichen (Peltigera apthosd) haircapped mosses (Polytrichum spp.), and ragged mosses (Brachythecium spp.) covered 50% of the ground in this association. The same species of trees and shrubs dominated the path association but grew further apart in this association. I recorded a variety of other grasses including spike trisetum (Trisetum spicatum) and blue wildrye (Elymus glaucus) with covers of less than 5%. A wider variety of forbs grew in the path association and made up between 25 and 30% of the cover. Cryptogams covered 10-25% of the ground in the path association. Mature trees surrounded the clearcut, and young aspen and lodgepole pine trees less than 0.5 m tall grew in the clearcut. Fewer shrubs grew in the clearcut than in the other upland associations. A mix of 46 grasses and fireweed (Epilobium angustifolium) dominated the association. The cryptogams covered less than 10% of the clearcut. 4.4 Phenology 4.4.1 Identifying Phenological Groups and Indicators The initial hierarchial clustering of all species in each plant association produced 8-15 small groups for each association. The species in each small group had almost identical phenologies, in most cases, and represented a distinct overall phenology that included flowering and dispersing fruit at the same time. These small groups were joined into three to six larger groups by the clustering process that clarified the more general phenologies found in each association. Two of these larger groups were consistently distinct and represented the earliest and latest flowering species. The group of earliest flowering species was in all upland associations, and the group with latest flowering species was in all associations. The other large groups formed when smaller groups were linked by the clustering process represented species that flowered mid season (late spring to mid summer). These groups, at times, included species that did not flower at the same time as other species in the group. Species with unique phenologies such as very short flowering periods like sagebrush Mariposa lily (Calochortus macrocarpus) or species with a long flowering period and early fruit dispersal like prairie pepper grass (Lepidium densiflorum) were grouped with species that did not have the same flowering period (Figure 6, underlined species). Generally, the clustering process grouped most species so that each larger group represented five general phenologies. I named the five general phenologies for group flowering times; Early Spring, Spring, Early Summer, Mid-Summer, and Late Summer (described in Section 4.4.2). These general phenologies 47 were consistent between sites although not all groups were represented in all plant associations. The time between my phenological observations varied from five to nine days depending on conditions. The dates used in the data matrixes mean the phenology was observed in the week of the listed date. In addition, the code I used detailed the phenology of a plant. The accuracy of subjectively deciding whether the modal stage of a species is 10-30% in flower or greater than 30% in flower, for example, by observing as few as 10 plants per association was questionable. Therefore, I did not consider differences of less than two weeks to be significant when comparing the phenologies of species or groups of species. To clarify and compare phenology found in the riparian and upland areas of each site, I needed to reduce the number of species to be included in the process. I did this by choosing one species (the indicator species) from each of the small groups of species with nearly identical phenologies to represent each of these phenologies. Indicators were chosen that both represented the small group and remained within this same group in both 1995 and 1996. The number of indicators per plant association varied because the number of species and small groups included in each general phenological group varied within each association. All the indicators from the riparian associations at each site were used in the clustering process and combined into new ordered data matrixes. The same was done with the indicators for each upland association. This step delineated four to five major phenological groups. In most cases, these groups had very similar phenology including flowering and fruit dispersal dates as the associations from which the indicators originated. I combined the upland and riparian indicators from each site for clustering and in ordered data matrixes. In 48 each case, the resulting groups matched the phenologies of the upland associations of each site but not the riparian associations. 4.4.2 Phenological Groups I separated species into phenological groups based on the date when they were 10-30% in flower. These group flowering patterns are summarized in Figure 12. Figure 12. Average dates for initiation of flowering for each phenological group, B G and SBS sites, Junction Sheep Range Park, and Horsefly, British Columbia, 1995/1996. Phenological Group April May June July August Bunchgrass Sites Early Spring Spring Early Summer Mid Summer Late Summer Sub Boreal Spruce Early Spring Spring Early Summer Md Summer Late Summer 49 The species in the Early Spring group usually had full sized leaves by April 30 in both years. The plants initiated flowering from early April to the last week in May and initiated dispersal of fruit by the end of June at the BG sites. At the SBS site, all species in this group initiated flowering between the end of April to the end of May and most started dispersing fruit from mid to late July (Androsace septentrionalis; Figure 13a). This Early Spring flowering cluster had only one or two species in each of the BG and SBS riparian associations and the SBS upland associations. The Spring group was leafed partly out by April 30 and some species had recognizable flower buds by this time at the BG sites. These species initiated flowering from mid May until early June at the BG sites. Flowering started and ended later at the SBS site by about two weeks. Fruit dispersal occurred in the first half of August at the B G sites and the last half of August at the SBS site. Most species showed a regular development pattern with each stage lasting one to two weeks until fruit dispersal which lasted longer (e.g., Geum triflorum; Figure 13b). The Spring group included a variety of other phenological patterns. The Carex species in this group had steady development until the green fruit stage and remained at this stage for four to six weeks ( C. atherodes; Figure 13c). Most species in this group at the SBS began growing in late April and then developed quickly to the flowering stage (Betula glandulosa, Figure 13d). The phenology of the species in the Early Summer group varied and flowering times overlapped those of the Mid Summer group making these two groups difficult to separate in some associations. The species at the BG sites in the Early Summer group generally had first leaves unfolding by April 30, initiated flowering early to mid June and began dispersing fruit 50 by late July to mid August. Most species in this group at the SBS site had no new growth until early May, flowered mid to late June and dispersed fruit in the last half of August. Flowering periods showed a variety of patterns from very short and quick to fruit development ( Stipa comata; Figure 14a) to longer flowering and late fruit development (Astragalus agrestis; Figure 14b ). At the SBS site, most species in the Early Summer group had extended flowering periods and did not ripen fruit until late August (Smilacina stellata ;Figure 14c) Many of the Mid Summer species at the BG sites including parsnip-flowered buckwheat and annual hawksbeard (Crepis tectorum) had no growth showing until early May, initiated flowering from mid June to early July, and most had ripe fruit by mid August (later in the riparian associations). A few species had a late start, grew quickly and flowered in a short burst (Orthocarpus luteus; Figure 14d ). Others started earlier, but developed and flowered at a slower pace ( Galium boreale; Figure 15a ). At the SBS site, most species in this group did not initiate growth until mid to late May, began flowering late June to mid July, and, except in the forest gradient, more than half had ripe fruit by the end of August (Rosa acicularis; Figure 15b). A few species including yarrow (Achillea millefolium) and Canada goldenrod initiated flowering with this group but had extended flowering periods (six to eight weeks) that completely overlapped the Late Summer species' flowering period Species in the Late Summer group were distinguished by generally late growth initiation, slow development, and flowering that began late (early July in BG sites and late July in SBS sites) and continued into early September at both the BG and SBS sites (Solidago spathulata and Aster ciliolatus Figure 15c and d). A few species in both study areas followed this 51 a. co a. co a. oo ro LU a. W 52 CD £ E oo CO LU <u *i (/) O CD .2 S '8 (13 (Q I ' O </> ro oi a N 00 => CM < T 3 CO < CN O 00 CD CN o CD E E oo 3 0 0 as LU 3 0 0 53 54 development pattern until flowering, which lasted only one or two weeks and this gave them time to disperse fruit by early September (sagebrush Mariposa lily, and fireweed for example). The Late Summer group had the most consistent phenology and dates of flowering and fruit dispersal across gradients and between sites. 4.4.3 Bunchgrass Phenologies The phenological groups and the flower and fruit dispersal dates for each BG riparian association (open water, wet riparian, dry riparian, and transition) were very similar between sites with two exceptions (data matrixes Appendix 1). The upland species growing in the BGxw/h dry riparian association (e.g., bluebunch wheatgrass) flowered two weeks later than those in the BGxw dry riparian association in 1996. This pattern was reversed for several species including Junegrass and giant wild rye in the transition association. They flowered two weeks later in the BGxw site. Hierarchial clustering of the 1996 phenological data of the riparian indicators at both sites produced nearly identical groups (BGxw cluster diagram is included in Appendix 2). Because of the similarity between the two BG sites, I used only BGxw indicators to form an ordered data matrix (Table 8) and to summarize the riparian phenologies. The phenologies of each group in the summary were very close to the phenologies of the same groups in the individual associations (Figure 16 and data matrixes in Appendix 1). In addition, the indicators placed in the same phenological groups in both the associations they represented and in this summary of the riparian area. Flowering dates of each phenological group were generally later in 1996 than in 1995 in all the riparian associations, but this was not uniform. Some groups showed no change (one 55 week or less) and others flowered two weeks later. Species in the Early Spring, Spring, and Early Summer groups generally dispersed fruit later in 1996 than in 1995. Species in the Mid and Late Summer groups dispersed fruit at the same time both years. The flowering order of species in each riparian association and in the riparian summary changed from 1995 to 1996 (Table 9). At the BGxw site, 14% of the species flowered later in 1995 than in 1996 and 50% flowered later in 1996 (Table 10). Only four species in the three associations, however, changed phenology enough to be in a different phenological group in 1995 than in 1996. Fourteen percent of the species changed fruit dispersal times between years. The species in the Late Summer group showed the least change in flowering and fruit dispersal times between years. There were 36 species in the riparian area. Some were in more than one gradient. Graminoids dominated the cover of the riparian associations, but there was a greater number of different species of forbs (Table 11). More forbs than graminoids flowered in late summer. Species were divided between four phenological groups: 17% Spring, 31% Early Summer, 25% Mid-Summer, and 25% Late Summer. The respective upland plant associations of the two BG sites (Dry grassland, shallow swale, and deep swale) were also similar but showed more variation between sites in terms of phenology than the riparian plant associations did (data matrixes Appendix 1). The dry grassland phenological groups identified for each site and their flowering and fruit dispersal dates were very close. There were several species in the three earliest phenological groups, however, that were two to four weeks earlier in their phenology at the BGxw/h site including 56 I (M N W T -CM CM T -T - CM T - O | T - (N » - O >> E CM CM CM CM CM CM CM ( N CM CM CM C\fl CM CM CM T - T -O - - O O O O O[0) O O O 01 O O 01 Ol T - CM o o q o o T - o o q O O T - o> o of co co r- co S N rf CO CO *T *T T ^ T T CO T CO Q. o. n 0) 0) 0) o) oi o co co cq 01 oi 01 r^ - r-l Q CO CO S N I^J r-— r— r— i— r^ - i r- co co co ci r~i 1- -M" 1 T 1 co T n 3 8 8 § ^ 2 i £ 8. .1.1 •6 E E I T— f M O J C N O C M O C M c - T - r - ^ O N O ) ' -O O O ' - O O i O l O * E 3 o o> <n r-. 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N C D T C O i n i O ^ i n .2 o I ai o> oilr-i|a> <n oilr-i— i-_ i— •—1 in m • ' t-~;'in co co col t in T CD "T • » 'fl'* «> £ <i> co ool in in T ^  C o> cs 2> E i * CO 3 £ o <o o .5 w |^co t ^| £ SI E * -=r ^ | ^ T ^ <=r co co C O | C M -r T | | -*r in ^ ^ ^ ^ i c M C M C O ^ ^ ^ " " ^ " ^ 0 " ^ ^ " ^ 8 c T 5 „ „ „ _ _ C > . ^ ^ C M C O C O C O O O C V C M C O C O ^ 1 CO a. aj Si co 1-"2 ^  "Slco •* 9 E £ 1 ! i l l S i Q. w 58 TABLE 10 . Number of species reaching flowering and fruit dispersal later in 1996 than in 1995, BGxw site, Junction Sheep Range Park, British Columbia, 1995/1996. Site association Flowering initiation Fruit dispersal Number of # Species and > 2 weeks later Initiation > 2 species that in group phenological in 1996 weeks later change phenological present group in 1996 group 1995/96 Riparian Spring 3 2 0 7 Early Summer 9 2 2 11 Mid-Summer 5 1 0 9 Late Summer 1 0 2 9 Total species/association 36 Drv Grassland Early Spring 1 3 0 7 Spring 7 4 3 12 Early Summer 3 3 1 8 Mid-Summer 4 5 0 9 Late Summer 3 0 0 6 Total species/association 42 Shallow Swale Early Spring 0 1 0 4 Spring 6 4 3 12 Early Summer 2 3 1 10 Mid-Summer 8 5 0 12 Late Summer 2 0 0 6 Total species/association 44 Deep Swale Early Spring 4 1 0 4 Spring 3 0 1 8 Early Summer 3 2 3 10 Mid-Summer 9 7 0 14 Late Summer 1 0 2 8 Total species/association 44 59 TABLE 11 . Number of life forms found in each phenological group, BGxw site, Junction Sheep Range Park, British Columbia, 1996. Phenological groups Life form Early Spring Spring Early Summer Mid-Summer Late Summer Riparian Forbs 0 3 6 2 8 Graminoids 0 3 4 6 1 Shrubs 1 0 1 1 0 Total 36 Uplands Forbs 8 12 12 8 6 Graminoids 0 2 4 5 0 Shrubs 3 0 0 1 4 Total 65 60 Alkali bluegrass (Poa secunda), and Junegrass. Fruit dispersal varied between sites, but was generally earlier in the Early Spring and Spring species at the BGxw/h site. The shallow swale plant association at the two BG sites varied more in phenology than the dry grassland associations. The Early Spring and Spring groups generally flowered earlier at the BGxw/h site. In addition, four species common to both sites, Junegrass, yarrow, western blue flax (Linum perenne), and parsnip-flowered buckwheat (Eriogonum heracleoides), flowered with the Early summer species at the BGxw/h site and with the Mid Summer species at the BGxw site. The phenologies of the groups found in the shallow swales were very close to those found in the dry grassland associations. At both sites, however, several species common to the dry grass and shallow swale plant associations ( Junegrass and parsnip-flowered buckwheat for example) flowered two weeks or more later in the shallow swale than in the dry grassland association. As for soil moisture and cover, phenology differed between the deep swales at each BG site. The phenological groups, the flowering and fruit dispersal times, and the species growing in the BGxw/h deep swale were very close to those of the dry grassland association of this site. Half of the species growing in the deep swale at the BGxw site, however, grew only in the deep swale or rarely in the other upland associations. The phenologies of the species in the five phenological groups of this association varied more therefore (ordered matrixes Appendix 1). The flowering dates for each phenological group were similar to those in the other upland associations with the exception of the Early Spring group which flowered three weeks later in 1996 than in the other associations. Species in this association generally dispersed fruit later than in the other upland associations. Also, the deep swale Early Spring 61 and Spring groups included only 27% of the total species compared to 40% in the other upland associations. I used 1996 phenological data of indicators from each plant association at each site to summarize upland phenologies. Although the upland associations were not as similar in phenology between sites as the riparian associations, clustering of the indicators did produce similar groupings between sites (BGxw cluster diagram is included in appendix 2). For the remainder of the upland summary I used only the BGxw site's indicators, because it included a greater number of the species common to these grasslands. When the 1996 indicators from the three plant associations were ordered by 1996 dates of 10-30% in flower in a data matrix, the groups formed were very close to those formed by the clustering process (Table 12). In addition, plant indicators placed in the same phenological groups in this summary as they did in the plant associations they came from and group dates of flowering and fruit dispersal were close to those found for each group in the individual associations (Figure 16 and ordered data matrixes Appendix 1). Forty-three percent of all species in the upland associations flowered later in 1996, and 25% dispersed fruit later (Table 10). Flowering order changed somewhat between years (Table 13), and four species were in earlier phenological groups in 1995 than in 1996. Sixty-five different species grew in the upland plant associations. Some were in more than one association. Although grasses dominated the uplands, there was a greater number of species of forbs (Table 11). More forbs flowered in the spring and early summer, and more graminoids in early summer and mid summer. Species were divided between the five phenological groups: 16% Early Spring, 21% Spring, 25% Early Summer, 21% Mid Summer, 62 8 I S E i I 2 E J5 E Q. « CO fO IN CM Ol « l tN f O r O r t e N C O O I C N M C N t N C N t N CNlrO CO CO CM CM CM CM CM CM CM CM I to J " bo ro ro CM CM CM rO| r o CO CO CM CM CM col O) t o CO CO CM CM CM COl I N CO fO CM CM CM CM] !d> <T> OT CO) •pi o . ' c n cn o t o ed fcn o.co co co f - r-0 to m ID M i€ I sag . 1 S I 'g 'g i . 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ID ID COl ( O T D I F T I O L O T O I O C O I F T I O L i O t f ) i O ( D t O i D i o i r ) i o < o i o i n l l o i o i o i o i o i o - o i o i o i o i n i o l i o i i O i o o i O i O i o i o m i f -^1 h r o N i n ^ v i o i f l T T i ' |CMCMCM-«r-«3-'>f'q--^--«3-TrTfTj-| • ^ C M T - - t 3 - - r r ^ T 3 - - < J - - g - T T T j - -l o o O ' j r O ' j ' r T T ' r T i l o o o r o o i r o i N c o r o N T r i -O O O C M C M C M C M C M C O C M f O f O l :CD m r— m ^ u o o c r - m c D r - ^ 3 „ , £ £ E u Q. oi oj r r r ; 10 in n t o o « w *e . c t : " o o u c o t o c o c o c a c a u c o 63 CO CO CN CM CO CM CO CM CM i - CM I C O C O C O C N C M C N C N C N C N C N C N C N C N 5 E CO CO CO CO N f T CM IN (N T I C O r O C O C M C M C M C M C M C M C M C M C M C M 1 t C O C O C N C N C N C N C N C N C N C N C N C N C N C ^ C O C O C N C N C N C N C N C N I C N C N C N -C M f O C O r O ' i - C M - > - C M C M C M C M * - C M ; CO CO CM CO C M ^ f ^ r C M T - C M - s r COCMCMCO (SI 1 - ^ M O <N (N CO (N CM CN C S I T - C M C M O C M C M COCMCMCM CJ i - CM C M C O C O C O i - C M - ^ C M i - C M C M O O O • C M C M C M T - C M I - T - T - C M C M CM CO (M CM T - CM CM CO CM CM O CM CN CO CM CM O CM CN CO CM CM O CM I - I> * CM CM CM CM i O - ^ - ^ - ^ O ' - ' - ' - O O C N (\| r r r ( M O i " C N I - O T - IN O >- O o o o o o o o o o #)) *a> o& C i j O O O O ' O T O O O O C CM O O •»- i - O o , t o f o a j r v ao4«oj CO O l N lO CO -COl O «2 <D Q) CO O O O O);0)j 0i]« to CD ^^^^^^^^^^^mM eoSio •* T * r * . J c o m C M C M C M t - V C M C M C O C N C M T j - C M C M C M O ^ r C N C M C O C M ' ^ -CO CO CO O O INjCft O CO h « i W Oi Qjj O JCD <3J CO. h- CO < ^ T C N C M C M O ^ - C M C M C M C N O C M C M C N i c O C J J O l B O C f t N - r -C M C M T - C M ^ | C N C M ' ^ C M T - 0 O T- O ( O O O t31 OTj T - O |Oni Iii W O O O l CO o j m c n o ) cn co co en co co o i O |d> CO CJ) eft C f t *o ffl c o j o j e o eo Gi «o • et) f ~ co h» H co t o K ' co (D col to i o m o m f i o i o | m l O C O l O if) T if) lO f i f l to l CD i f CD T t >J 1 f iT - J t f l .1 CO t CO CN CO CM CO CN COl ) r O f O f O V 1 - 1 - C O ^ C O C O C N C N q » O *T O T - C O C O O C O O C N ) l l S S l l l l l l l & l i l i t t f l s S ' i ' s Hil l&££i8 313*1 8 -a li.SISiVS'SSS'S'SS' I S 2^ £ &SS £ e & 8 E 'S S E i 3 3 8 8 8 8 8 8 & 5 is cd t o co eo !*• r*> 10 c h - cO I**- f»* t** II 10 t v m i n col * 0 ) J O | C BC O f ^ C D t D C O ^ f ' T ^ t f ' ^ J - l f ) r - 01 01 or> co r--j IJJ r - i> r—'co ca to co. co to co to s e o o j i v M o c o u D i n i C f t C f t i ^ r ~ c O r ^ . h - i f j i o t D i o 00 CO 1 -^ <£> r*- f*- CD iT> lO lO ID | H i t> K fcD 3 10 ;h*j'cD co i n ^ i - u C D C D C O C O i o i D C O i O l f l ^ i O i O C O l O C O O l f l ' t t i O ' • " • ' • i f l i o i n i n ^ f t ' C f i f l l O f i D l f ) T t f T 1 , 3 ' m i o t i o c O ' r ' M N i - c N ' r ] T J - ^ ^ ^ M C O ^ C N O C M ' C O C N C M C M O C M ^ f O O O ' C N O C N O O C N ^ O O O C O l . .• >i M -J ^ i f ' O O O ^ C N I - f O f T C N ' T O O O C O C N t O C O C O C O C N f C f l ; £ E g n a u D E c E a 64 65 66 and 15% Late Summer. Periods of flowering and fruit dispersal for each phenological group were generally earlier in the upland associations than in the riparian associations (Figure 16). 4.4.4 Sub-Boreal Spruce Phenologies I combined species from the open water association of the three fens included in the SBS site into one association for categorizing into phenological groups. The same was done for the riparian associations for each fen. Clustering of the indicators for each phenological group in each of these associations resulted in four groups (cluster tree diagram in Appendix two). The ordered data matrix of these indicators showed flowering and fruit dispersal times that overlapped those for each riparian association. Also, the species used as indicators were in the same phenological groups in this summary as they were in the individual plant associations they represented (Table 14 and ordered data matrixes Appendix 1). Thirty-one percent of the riparian species flowered two or more weeks later in 1996 than in 1995 (Table 16). Flowering order changed slightly between years, but no species changed phenological groups (Table 15). There were more forbs in the Mid and Late Summer groups than in the earlier groups, and more graminoids in the Spring and Early Summer groups (Table 17). Only willow flowered in early spring in the riparian associations. The other species were divided between the other four groups: 28% Spring, 22% Early Summer, 25% Mid Summer, and 22% Late Summer. The phenological groups in each upland association including the open edge had very similar flowering and fruit dispersal dates. I included the transition association (open edge) with the uplands at this site because its phenologies matched the upland phenologies more closely than the riparian phenologies. The phenology differed only in spring of 1996 when 67 both Early Spring and Spring flowering dates of the open edge were two to three weeks later than in the other upland associations. Differences between upland associations were more apparent in the changes individual species made in phenology growing in more than one association. Six species found in the forest association including heart leaved arnica (Arnica cordifolia) and arctic lupine (Lupinus arcticus) flowered two weeks later in this association than in the path association both years. Forth-eight percent of the species found in the clearcut association and in the path association flowered two weeks earlier in the clearcut. Six of these including wild strawberry (Fragaria virginiana), yarrow, and pinegrass (Calamagrostis rubescens) were in different phenological groups in the two associations. The upland associations also differed in the proportion of species included in each phenological group. The clearcut association had 31% of its species in the Early Summer group and the other associations averaged only 19% in this group (Table 17). The summary cluster of the upland indicators formed five phenological groups (cluster tree diagram is in appendix 2). The groups defined by the matrix of indicators had phenologies very similar to those of the individual associations included in this summary. Most indicator plants fell in the same phenological group in this summary as in the original plant association matrixes (Table 18 and ordered matrixes Appendix 1). Flowering and fruit dispersal dates were slightly later in 1996 for the groups formed in this summary, but the dates did not show the differences of two weeks that were indicated in some of the individual associations. Overall, 42% of all upland species flowered later in 1996, but 68% of the edge species did (Table 16). Flowering order changed, and six species 68 changed phenological groups between years (Table 19). There were more shrubs at this site, and they were found in all but the Late Summer group (Table 17). Graminoids were evenly distributed in all but the Early Spring group. Most forbs flowered in early to late summer. Species were divided unevenly between the five phenological groups: 4% Early Spring, 13% Spring, 26% Early Summer, 31% Mid Summer, and 25% Late Summer. Species in respective upland and riparian phenological groups generally flowered at the same time, but riparian species in all groups dispersed fruit two or more weeks later than the upland species in the same phenological groups (Figure 17). 69 °> CL •5 e S E V 3 •6 E E | » E I E CM CN CM CN CN I I CM CM CM CM CM l CM T - O O C 5 I CM CN i - i - CN CN CN T - T—I - - O O O C O ) CN I 5-1 I CM T - O O O O < • > CQ'r~ f o o < d o;oi N f eg la E 3 O o to >; d) in (0 OQ CO ra E cs CO •o dl •a k_ o £• ra E E 3 in cs 41 a. UJ _ l m < - - - o o . O O O ' CO o o o o o a o i o i c n CD CQiCD CD '(.(*- t>->;CD if) CO CO CO co l ID ID m f I CO » nj N- N | 'I o t in t ^1 I o M - o co col O f O IN IN I C 0) g CD CD roc ra ra O CD O Ol *r *r ^- *r CO CO o o l CN CN O O l CN CN O O • - CN i - CN cr o" CD CD E p] ro ro -2> t= o o . K | CD j K , I f in ^t in in i m t f [ i > CD i o col f T CO COl • •^ T CN CO CN ( *r o o ol < v o o a < f O O O i l ,3 E • 70 •o o> <J> ra~ ! a E 3 O o m 5= CD co m co ro E co ra T3 T3 & •a o CO E E 3 in co c .2 co a . bs id LU - I m < S E a . OJ a> —1 CO CM CM CM CM I CM CM CM I « CM H I CM. O <- r -O O O O < i|o> eo"a>| Ij CO (O lO 3^"I ' 8 8-tr cr a> p El •o E E | CM CM o ol i - T - o O ci ,oi oy en 0) o> 01 o> i l p t l tj- -<H CO ^ J - "*t\ N * n co rav q t o N (NJco q f o o q ro o l ^ o o C J o c j O 5 D) y » f ; £ E 2 E 71 TABLE 16 . Number of species reaching flowering and fruit dispersal later in 1996 than in 1995, SBS site, Horsefly, British Columbia, 1995/1996. Site association Flowering initiation Fruit dispersal Number of # Species a n d > 2 weeks later Initiation > 2 species that in group phenological in 1996 weeks later change phenological present group in 1996 group 1995/96 Riparian Early Spring 1 1 0 1 Spring 3 2 0 13 Early Summer 4 0 0 9 Mid-Summer 2 2 0 9 Late Summer 2 1 2 6 Open edge Early Spring 2 1 0 2 Spring 2 1 0 6 Early Summer 8 2 4 10 Mid-Summer 6 0 4 10 Late Summer 5 1 1 6 Forest Early Spring 1 1 0 2 Spring 1 3 0 5 Early Summer 1 3 1 5 Mid-Summer 7 3 0 9 Late Summer 3 0 2 10 Path Early Spring 1 1 0 2 Spring 2 1 0 5 Early Summer 4 3 0 14 Mid-Summer 5 4 1 12 Late Summer 4 0 3 14 Clear cut Early Spring 0 0 0 1 Spring 4 4 0 5 Early Summer 3 3 1 10 Mid-Summer 2 1 0 8 Late Summer 0 1 0 8 72 TABLE 17. Number of life forms found in each phenological group, SBS site, Horsefly, British Columbia, 1996. Phenological groups Life form Early Spring Spring Early Summer Mid-Summer Late Summer Riparian Forbs 0 3 2 5 7 Graminoids 0 5 4 3 0 Shrub/tree 1 1 1 0 0 Total 32 Open edge Forbs 1 1 9 10 5 Graminoids 0 3 2 2 2 Shrub/tree 1 2 1 1 0 Total 40 Forest Forbs 1 1 4 10 9 Graminoids 0 2 0 0 1 Shrub/tree 1 2 1 1 0 Total 33 Path Forbs 2 4 14 18 17 Graminoids 0 2 1 2 4 Shrub/tree 1 2 2 2 0 Total 71 Clear Cut Forbs 0 4 10 6 7 Graminoids 0 0 0 4 2 Shrub/tree 1 1 2 1 0 Total 38 73 Ol CM CO CO CO Ol CM CO CO CM D) CM CO CM CM J «— " < O) CM CO CM CM 3 CM CO CM CM co CO la E D O o CQ 4) in i _ o X cf 4-1 'cn CO CQ CO TO E a ro T3 "D V TJ o > CO E E 3 cn re oj k_ re "D C ro Q oo Ul -I ffl < I-; CM CO CM CM C M C N C M C M C M T - C M C M C M C M C N ; C M C M C M C M C M T ~ C N * - C M C M C M C M C M C M C M C M T - C N ^ - C N C M C M C M C M C M T - C M O C M T - C M C M C M T - C M O C M O C M T - C M C M O C M O C M O 0 * - C M O * - 0 * - 0 0 0 0 j CM CO CM CM CM CM • C M C M C N C M C M C O C M C M CM CM CM CM CM * - CM CO CM CM CM CM CM CM CM O CM CM CM T - CM CM CM j •6 E 6 I N O O I ( M f M M O ( N r O ' - r ( N O O ' - ( N O O ' - O I ( N O O C N C N I - C N O C M ^ - O ^ - I - O O O I - T - O r O)l O C N CN O O O ^ - T - T - C N O ^ - O O O O O O O O O O (OJ" O O ^ - T - T - ' - O - ' - O O O O O O O O C N O O O O O o o o o CN;O> o o o O ^ — C N O T — O O O ; 0 3 O O O T - o o o p)| o <o>| o o o o o o i o i a a i o c o a i o ) 5 O ' C O CO CT> O ' C p C R O J C B C J O C P O J c O c O C O CO CDl 03 CO N 1 N N ^ 00j If) CD if) if)| i->. co h» i*- m 'h»* T m co co i ^ m CD co if) co h-^ T T CN CN - CP h»; CD If) 6 Wmlm £ K 3 £ ? l • O j O o co o) a cn co oV a a c o c o c o c o c o s r o c o c o r - r--tr>,,rv. s ' c o co to co co m | O O i - O O O O O O O O O O l o T- T - oi O T - Q ) C O O) o> o T ' c j j j o O Q O ) C J ) 0 > C I ) 0 ) t p c O C O C O t O t O t O * t j ^ C O t © W C » O } O > O 3 C 0 C 0 C # C O ^ r ^ r - - h - t O CO j CD CO CO CO if) CD i^ . r- c- c- C O C D C D C O C D C D C O l f ) C D l f > l O r - » l t - in co T T T T IT) if) >J 1 N~: i n ^ i j o i n ^ c v ' i O ' r c N i f N T T t f ) N - 0 " I ' - N - 0 " J > *D 7; i2 -r; O) t0 ^ C c C ro S 0 1 S i a y i5 £ := := S. & ] CO > £3 t f l i - Q O > — — m c D c o i n c Q i r t m i f ) m (D 1 C N i o i n i r ) i f ) C N C N * t f m 10 C N C N ^ r T T C N C N C O T T O f N r - ' N ' T r l f M C N C N T CO O o ^ - ^ r c o c o o o o T O O O ^ - T O C N O O O T O O T T T l O C ^ t f ) l f ) T T l / ) T T T C O C O l f ) T T T T T T C O C O C N T i n C N M T T T T T T C O O O T T T T C O ' t r T ' C N C N O T T C N C N C O T T C O T T C N O O T T T T C N C O ' T O l O O C O C O ^ C N C N T C O C N C O C O O O O C O T T C O O O C O O ] O O O C O O O C N T ^ C N i - C N l O O O C N C O T O O O C O O O O O O O O ^ - T O t - O O O O O C N C N - C O O O O O > > . 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CN tf» o o o o o • o ra E 3 o o CD in o i o cn"o^cD o o o o o < CO CM dOtOtulltUfuiuiui w| Q • • : • " : • <• H l i c ^ H B l ^ s coco Oct o v o oi j <Mii! l l lM - N N :c cr- cn OI CO CM ^ ^^Plli^^^^ -5 CO i n —j ; lO *T « C ^ N j ^ CO V 'in W m ra E S ra T5 •o L_ a> •a k_ o >< ra E E 3 in ra <D ra T3 C ra a. =3 S" CD 3 C o o CO f -Ol _ l m < 3 " CD (O (D ^ 1 T c ^ ^ i D i n m c o i ' j ^ CO CN —> CN 3 CO 5 > f ( N c o ^ r ^ i c M c o ' r > , CO C N C N ^ ' C ' C C N C O ^ f > O C N ( N C 0 C 0 C 0 O C N « CO 5 OOrtNtNCMOOCN o 8 o E S 2 2 5P 8 a <i) 2 a ^ ro ro ro o 5 J ^ Q - o w o c c c o c a c n i o - c t O t f l r o r o o o o r o r o o . 77 78 79 5.0 Discussion 5.1 Phenological Changes Related to Growing Degrees Over 40% of the species in both riparian and upland areas at both the BG and SBS sites flowered two or more weeks later in 1995 than in 1996. This clearly reaffirms that calendar date (day length) cannot be used to predict flowering dates (Blaisdell 1958, Fitter et al. 1995, Diekman 1996). Spring (up to May 30) total degree days averaged slightly lower than 20 year norms in 1995 at both sites (Figure 7). End of May totals averaged much lower in 1996 (69% of 20 year norms at the BG sites and 73% at the SBS site). Plant phenology between years reflected these differences in spring temperatures. At the BGxw site, 43% of the upland species and 50% of the riparian species flowered two or more weeks later in 1996 than in 1995. At the SBS site, 31% of the riparian species and 42% of the forested upland species flowered later in 1996. Fitter et al. (1995) compared the phenology of 243 species over 36 years in England and similarly found that 60% of the species flowering in spring (January to April) had first flowering dates that were strongly affected by temperatures one to two months before flowering. Twenty-five percent of the species flowering in summer had flowering dates that were strongly affected by temperatures up to four months before flowering. Blaisdell (1958) summarized phenological data for eight species over 16 years growing in an area with winters similar to those of the BG sites. Later than average plant development was always associated with mean temperatures lower than average. Beaubien's (1991) study of species growing in Edmonton, Alberta over a two-year period similarly showed later flowering related to lower spring total degree days. 80 Although plants generally developed earlier in 1995 related to the higher spring degree day totals, the changes in phenology were not uniform. Fewer than half the species flowered later in 1996 and fewer had later fruit dispersal dates despite the annual differences in total degree days. A larger proportion of the BG Mid Summer species flowered later in 1996 (59%) than Late Summer species (24%). At the SBS site, 63% of the species in the Spring flowered later compared to 32% of the Late Summer species. This variability of changes in phenology related to spring temperatures confirmed those of other studies (Fitter et al. 1995, Sparks and Carey 1995, and Diekman 1996). White (1979) found that 49% of the earliest flowering species in a mixed prairie in Montana flowered at different times with changes in spring temperatures and only 34% of the late flowering species did. Each species responds differently to the sum of abiotic factors affecting development and this makes it difficult to predict how phenology will change each year. In this study, this variability affected the 1995 and 1996 flowering sequence within each phenological group. The order of initiation of flowering within each phenological group changed in all groups in all plant associations between years. Classification of species in the phenological groups, however, remained almost the same between years in all associations. This agrees with Pitt and Wikeem (1990) and Diekman (1996) who found changes in flowering sequence but consistent membership in phenological groups. In addition, changes in sequence of flowering between 1995 and 1996 may have resulted from my observation techniques. I recorded phenological observations once per week and I could miss when a species reached each stage. Also, without counting actual blossoms, accuracy of labelling a species 10-30% or more than 30% in bloom is questionable. Beaubien (1991) stated that during the peak of flowering, 81 observations need to occur almost daily and that the flowering order sequence for an area should be based on 10-13 years of data. 5.2 Effects on Phenology of Soil Moisture and Precipitation Changes Between 1995 and 1996 Winter precipitation totals affected the depth of the standing water and soil moisture levels in the riparian areas at both SBS and BG sites more than growing season precipitation did. The greater amount of winter precipitation (snow) in 1995/1996 than in 1994/1995 raised the water levels at all the riparian areas well above those of 1995. Even with less growing season precipitation in 1996 than in 1995, these higher standing water levels persisted through the 1996 growing season. The higher 1996 water levels in the riparian area meant that a larger portion of each riparian zone was under water for longer in spring 1996. Some species at all three sites that developed in the wet riparian zone after the water receded in 1995 did not appear at all in 1996. Examples from the BG sites are common spike rush (Eleocharispalustris) and little meadow foxtail (Alopecurus aequalis). At the SBS site, purple-leaved willowherb (Epilobium ciliatum) and northern grass-of-Parnassus (Parnassia palustris) grew throughout the riparian zone in 1995 but only in isolated high spots in 1996. The transition areas between riparian zones and uplands at all sites also reflected these higher water levels in 1996. These areas showed the greatest difference in spring soil moisture levels and the largest shifts in phenology between years. Other studies have suggested that soil moisture and precipitation affect later stages of phenology. Blaisdell (1958) stated that late development was more closely related to precipitation than to temperature. Dickinson and Dodd (1976) varied the level of moisture 82 reaching a section of short grass prairie over two years. They found that species that flowered from late May to late June had different phenologies depending on moisture levels. Pitt and Wikeem (1990) found that five of the eight Spring Ephemerals (my Early Spring group) completed growth one month earlier in the year with lower precipitation (growing degree day differences were negligible). In my study, changes in phenology related to soil moisture were not as clear. Fruit dispersed earlier in 33% of the Early Spring and Spring species at the BGxw site and 35% at the SBS site in 1995 (the drier year). Higher spring temperatures in 1995 or increased soil moisture in 1996 could have caused these differences. Many species in the later flowering phenological groups reached flowering and fruit dispersal later in 1996 though this part of the growing season was drier in 1996 than in 1995. Some Mid-Summer species (e.g. yarrow, northern bedstraw (Galium boreale), and dune goldenrod/ Solidago spathulata) at the BG sites flowered for longer periods in 1995 after the rains in July and August. Sauer and Uresk (1976) saw a similar trend of extended flowering periods in later flowering species related to precipitation. The grasslands at the Junction have a more even distribution of precipitation throughout the growing season than the British Columbia southern interior grasslands studied by Tisdale (1947) and Pitt and Wikeem (1990). In July 1995 and in August 1996 at the Junction sites, precipitation came just as soil moisture levels reached their lowest level in each association. In both dry grassland associations and the deep swale at the BGxw/h site, soil moisture did drop below 10% at the 10-20 cm depth, a level that can cause growth to stop (Brady 1990), but that is dependent on the moisture holding capacity of a soil and the depth of roots of 83 each plants that I did not consider in this study. Soil moisture at the SBS site never fell below 20%, which should have had no effect phenological development (Brady 1990). 5.3 Variations in Species Phenology Between Plant Associations The plant associations of this study were related to microclimates that differed in several ways. In both study areas plant associations had different levels of soil moisture. Soil temperatures most likely varied with the moisture levels, because wetter soils take longer to warm in spring (Brady 1990). Canopy cover was an additional factor at the SBS sites that affected the microclimate of each plant association. The shallow swales in the BG sites had more dense cover of drying grasses from the previous year than the other upland associations and this also may have kept the soil cooler longer in spring in this association. In addition, except for the deep swales, the associations at the two BG sites were very close in soil moisture levels and ground cover (Table 5 and 6) but differed in air temperature (Figure 7). Some species in this study that grew in more than one association differed in flowering and fruit dispersal dates between associations (e.g., Junegrass and bluebunch wheatgrass). These grasses grew in the transition associations and all three upland associations at both BG sites. In 1996, Junegrass began flowering and ended flowering two weeks earlier in the dry grass association than in the other three associations at the BGxw site. Timing of fruit dispersal did not differ between associations. In 1996, bluebunch wheatgrass showed the same change in flowering dates as Junegrass, but fruit dispersal varied more. Dispersal began on August 7 in the dry grassland association and on September 7 in the shallow swale. These two species also showed a shift in phenology between the BGxw and BGxw/h sites flowering an average of two weeks earlier at the BGxw/h site both years. 84 Other species, like arctic rush, showed almost no change in phenology between associations.. It grew in all the riparian associations at the BG sites. It flowered and dispersed fruit at the same time in all associations except one (deep water association of the BGxw site) where it dispersed fruit two weeks earlier. Overall, most species that were common to several associations in the grasslands of this study showed some shift in phenology between associations. Those that did change flowered and dispersed fruit earlier in the dry grassland associations than in other associations at the BGxw site and earlier in the dry grassland and deep gully association at the BGxw/h site. Species also shifted phenologies between upland associations at the SBS site. Northern bedstraw flowered three weeks later in the open edge association in 1996 than in the path and clearcut associations. Yarrow flowered three weeks earlier in the clearcut association than in the other upland associations and pinegrass was five weeks earlier in this association. Forty-eight percent of the species common to both the path and the clearcut changed their phenologies between these associations. This range of phenological response to microclimates are similar to those described in other studies that related phenology to micro sites (Jackson 1966, Ratcliffe and Turkington 1989, Beubien 1991). The phenological shifts for species that grew in several associations at each site indicated that the microclimates of these plant associations affected phenology and these variations in phenology should be included in a phenological description of these ecosystems. 85 5.4 Phenological Groups Identified There were no completely discrete phenological groups and I classified species into five phenological groups in this study based on flowering times of individual species. I did not consider other stages of development for placement in a group unless the flowering period was long and overlapped the flowering period of two phenological groups. The earliest flowering and latest flowering species were usually the easiest to classify. The clustering process consistently grouped some species in each association into the Early Spring or Late Summer groups, because these groups had generally distinct uniform phenologies. The phenologies of species that flowered mid season varied more and were less distinct. The groups formed reflected this variation. In order to focus on an easily identified phenological stage, I chose to divide the mid season into three flowering periods and classified species based on when each was 10-30% in flower. This grouping seemed arbitrary, but when the species in each association were placed in order of flowering dates, mid-season flowering species did divide into three semi distinct groups (Spring, Early Summer, and Mid-Summer) that began flowering approximately two to three weeks apart but with flowering periods that overlapped (Figure 12). Except for species with unique phenologies, these groups also matched the groups formed by the clustering process. Pre-flowering stages of each group were not consistent within each group, but the post flowering stages showed more similarity. The phenological groups identified in this study were similar to those used by Dickenson and Dodd (1976) in a study of a shortgrass prairie with similar precipitation levels and temperatures as the Junction grasslands. They divided the species into six phenological groups with similar flowering periods as the five of my study except that the Mid-Summer 86 group of my study was split into two groups. Their mid summer groups also showed variation in pre flowering phenologies. Beubien (1991) divided the Edmonton growing season into seven phenological groups. Again, the groupings were similar to my study but with an extra division in spring and late summer. Tisdale (1947) and Pitt and Wikeem (1990) classified species by over-all phenological patterns rather than flowering times. Pitt and Wikeem's Spring Ephemeral group had more species but corresponded to the Early Spring group of this study in its phenology. Their Protracted Growth group corresponded to the phenology of the Late Summer group of my study but with a later flowering period. Species that flowered early and midsummer in their study also overlapped in flowering periods. Pitt and Wikeem defined two mid season groups, Summer Mature and Summer Quiescent. Summer mature species cease growth to avoid drought and may show regrowth in response to fall rains. Summer Quiescent become only semi dormant during drought and continue growth when soil moisture levels increase. I recorded regrowth for only three species in 1995 (Junegrass, needle-and-thread grass, and bluebunch wheatgrass) and none in 1996. I did not observe semi-dormancy in any species. Five species in the Early Summer and Mid-Summer groups of the dry grassland and shallow swale associations were dormant by mid August, but this was not a general characteristic of these groups. This lack of observed dormancy or semi-dormancy may have been a result of the precipitation that came just as soils dried. I was not, however, observing changes in foliar cover and may have missed a reduction in growth. For any ecosystem there will be a number of species along a phenological continuum that will be difficult to separate into groups with distinct phenologies. A method of 87 classification will need to be used that both simplifies and best represents the phenological progression in the ecosystem. In this study, the five general phenological groups (Early spring, Spring, Early Summer, Mid Summer, and Late Summer) could be used to summarize and represent the phenological progression of the BG and SBS study areas, but this summary missed the diversity of phenological adaptions to the varied habitats that made up each study area. To relate phenological progress in the varied habitats that were part of each study area required comparing the specifics of the phenological groups identified in the riparian and upland areas and in the individual plant associations of each area. 5.5 Bunchgrass Riparian and Upland Phenologies Differences in phenological patterns occured between most associations in the BG sites in the dates of flowering or fruit dispersal and/or in the size of each phenological group in an association. I identified the most significant differences in phenological patterns between upland and riparian habitats. A single riparian species flowered in early spring compared with the uplands where 17% of the species were in the Early Spring group. By early summer (mid to late June), only 50% of the riparian area species had flowered, but 63% of the upland species had. In late summer, 25% of the riparian species remained in flower compared to 18% of the upland species. Species included in each riparian phenological group dispersed fruit three weeks later than species in the respective upland groups. These differences in phenological patterns between upland and riparian habitats may be explained by the differences in microclimates of these two habitats. The riparian areas at the BG sites sat in depressions and surrounded by grasslands. Cold air drainage lowers the temperature of depressions and killing frosts occur later in the spring and earlier in the fall 88 than in adjacent uplands. The wet soils of riparian areas warm more slowly than the dryer upland soils. In addition, cold temperatures both reduce decomposition and nutrient availability that also slows growth in plants (National Wetlands Working Group 1988). The cooler temperatures and slower development could explain the smaller number of species in the earlier flowering groups in the riparian area. Longer flowering periods and later fruit development could be related to both the cooler temperatures and lower levels of available nutrients. In both 1995 and 1996, only a few plants of the Carex species in the riparian area dispersed fruit and this was done late in the season. Others have also reported low levels of seed production in Carex species (Bernard and Gorham 1978). Because Carex species commonly reproduce vegetatively, the low levels of seed production have not been considered important to maintaining riparian habitat. Shaver (1979), however, showed in his study that genetic diversity resulting from sexual reproduction was important to Carex adaptation to the varying nutrient levels commonly found in riparian areas. This emphasizes the need for riparian plants to reach maturity and dispense fruit to maintain the diversity of the riparian areas. The two BG sites whose microclimates differed mainly by slight differences in degree day totals had common species that flowered two or more weeks earlier at the BGxw/h site than at the BGxw site especially in 1995. The Range Management Guidebook (MOF 1995a) suggests that grassland range is ready for grazing when Junegrass and arrow-leaved balsamroot are in flower. In this study, these two species began flowering one month apart in 89 1996. It appears that even slight shifts in microclimates can result in changes in phenology. These results emphasize the need to assess each site separately in terms of phenology. In addition, the upland associations and species showed smaller but important variations in phenology. A wider range of species grew in the BGxw deep swale and each phenological group of this association included more varied phenologies than in other associations. The delayed flowering in spring 1996 in the deep swale may have resulted from the much wetter soils in this association that year (Figure 11). The higher water levels in spring 1996 of the BGxw/h dry riparian association compared to the same association at the BGxw site may have caused the later flowering dates of the upland species growing in the BGxw/h association. Some upland species that grew in the transition associations (e.g., western blue flax and yarrow) at both sites had the same phenology across the associations. Other upland species growing in the transition associations including Junegrass, bluebunch wheatgrass, and NuttalPs pussytoes (Antennaria parvifolia) flowered and dispersed fruit later both years. Species growing in the transition associations of both sites also showed the greatest shifts in phenologies between years perhaps related to the wetter, cooler soils in spring 1996. The different phenological patterns of the transition and deep swale areas combined with the wider variety of species found in these areas suggest that these areas may be playing an important role in maintaining the diversity of the Junction grasslands. Grassland species that require moisture to establish and/or to reach fruit dispersal commonly grow in the deep swales and transition areas and rarely in the other upland associations. Some examples are sticky purple geranium (Geranium viscosissimum), star-flowered Solomon's seal (Smilacina stellata), and northern bedstraw. In years that have enough soil moisture, these species 90 spread out to the dryer upland associations. They may not survive or reproduce in these associations in drought years, and the swales and transition areas, therefore, act as a refuge for these species in these years. Some species common to the upland associations such as bluebunch wheatgrass have erratic and unpredictable seed production (Quinton et al. 1982). They can stop growth and may not reach the stage of fruit dispersal if soil moisture is limiting and temperatures high (Wilson et al. 1966). In drought years, bluebunch wheatgrass growing in the deep swales and transition associations continues to grow and disperse fruit and help to maintain the populations of these species in the grasslands. The differences found in phenologies between associations at the BG sites emphasized the need to see a management site as a combination of habitats with plant associations that are not all at the same stage of development. In this study, I summarized each riparian and upland area using indicator plants from the BGxw riparian associations and from the BGxw upland associations separately. The resulting phenological groups for each area and their phenological patterns matched those described for each association included in the summaries. When I combined indicator plants for the entire site into a single summary for each site, however, the resulting phenological groups did not match the phenological patterns of the riparian areas. This indicated the need to identify and compare phenological patterns in at least the two predominant habitats of the BG study area to summarize a site's phenology in a manner that will help manage for diversity. 5.6 Sub-Boreal Spruce Riparian and Upland Phenologies Riparian and upland areas at the SBS site differed very little in the size of phenological groups and dates of flowering. The Spring group included a greater proportion (28%) of the 91 riparian species than the same group in the uplands (13%), but other phenological groups differed by only a few percentage points. As in the BG sites, species began dispersing fruit later in the season in the riparian associations than in the upland. Riparian and upland phenologies may have been more similar at the SBS site than at the BG sites because of the different plant associations involved and/or because these two habitats at the SBS site shared similar early spring air and soil temperatures. The riparian areas at the SBS site were not in a deep depression, and were surrounded by forest on all sides. These two factors lessened the effects of cold air drainage. In addition, shade in the upland associations (except the clearcut) kept air temperatures cooler than in the unshaded riparian associations. Snow remained in patches throughout the uplands including the clearcut later in spring than in the riparian area resulting in more similar soil temperatures between these areas.. The riparian areas in the SBS, therefore, warmed more quickly allowing riparian species to begin growing as early or earlier than some upland species. The later fruit dispersal of the riparian areas may be related the slower growth associated with lower levels of available nutrients that are common in fens. As in the grasslands, to maintain the diversity of a riparian area the late season fruit dispersal in riparian areas must be considered in any management plan that involves these areas. The upland associations at the SBS site represented microhabitats that varied in canopy cover and soil moisture and the phenologies of each association reflected these variations. Forty-eight percent of the species growing across the uplands flowered earlier in the clearcut and were included in an earlier phenological group in the clearcut than in the other upland 92 a associations. These results agree with the findings of Jackson (1966) and Dale and Causton (1992) who found that different levels of shade affected flowering dates and phenology. Phenological groups in the open edge association showed the largest shifts in phenology from 1995 to 1996 of all the SBS upland associations. All groups in this association had flowered two or more weeks later in 1996. These changes in phenology may have been caused by the higher water levels in the riparian area in 1996. The edges of riparian areas are the section of the forest ecosystem that receives more sun and where snow melts first in spring allowing the soil to warm more quickly. In 1996, however, the edge area remained under water until the middle of June delaying the warming of the soil. The later phenologies of the forest species growing in the edge association reflected this. In 1995, plants of several spring and early summer species including Saskatoon (Amelanchier alnifolia), wild strawberry, and kinnikinnick (Arctostaphylos uva-ursi) flowered at the same time in the edge association as in the clearcut, but in 1996 they all flowered later than the clearcut. The riparian species growing in the edge association did not shift phenologies between years. The open edge or transition areas associated with each riparian area at the SBS site may serve to maintain diversity in the forest ecosystem. I noted during the two years of observations that most species common across the forested uplands flowered much less or not at all under complete canopy cover. Dale and Causton (1992) also found that shading reduced the prevalence of sexual reproduction in some species. Species common in the forested association and also rarer species grew successfully in the edge association and usually produced fruit growing in this association. The edge of each riparian area, therefore, allows forest species the opportunity to reproduce sexually. 93 A single summary of the site using plant indicators from uplands and the riparian areas did not represent the variety of phenologies in all the plant associations. The summary of the riparian areas using plant indicators matched the individual associations closely, and the upland summary was very close to all but the clearcut association. The forested study area like the grassland study area had plant associations related to a variety of microhabitats with different phenologies that need to be clarified when describing this type of area and its level of range readiness. 6.0 Summary To identify patterns of phenology in each plant association that made the BG and SBS study areas, I used hierarchial clustering which grouped species with similar phenologies, but did not clarify how phenology differed among associations or changed from 1995 to 1996. To focus on a single developmental stage, I then placed species found in each association in an ordered data matrix based on the date each species was 10-30% in flower. This highlighted changes in phenology from 1995 to 1996 within each plant association and made comparisons among associations possible. Differences in spring degree day totals appeared to affect phenology. More than 40% of the species at the three sites flowered two or more weeks later and less than 2% flowered earlier in 1996 the year with much lower spring degree day totals. Twenty-four percent of the species dispersed fruit later in 1996. Changes in phenology related to yearly differences in soil moisture were obvious only in the transition areas. More species growing in the transition areas between riparian and upland areas flowered two weeks later in 1996 than species growing in other areas. This was 94 probably related to the higher levels of winter precipitation in 1996 that increased the levels of standing water in all the riparian areas and therefore the time that the transition areas were subject to saturated soils. Changes in phenology were variable. Late summer flowering species shifted phenologies the least between years at all sites. Other generalizations can not be made with only two years of data. This short study suggests that the effects of yearly differences need to be considered when describing the phenology of an area. I categorized species into five general phenological groups based on flowering dates (Early Spring, Spring, Early Summer, Mid Summer, and Late Summer). The Early Spring and Late Summer groups each had distinct, uniform phenologies. The three mid season groups (Spring, Early Summer, and Mid Summer) overlapped in both flowering and fruit dispersal times and included a variety of phenologies. Although the five groups divided a continuum of phenologies and did not represent exclusive groups, they did help clarify how phenological patterns within each plant association differed from other associations. Most associations differed in at least one aspect of their phenological patterns related to dates of flowering and fruit dispersal and/or number of species in each phenological group. In addition, the phenology of some species common in several associations varied among associations. These differences suggested that species in an association had phenological adaptations related to the habitat and microclimate where each association grew. The uplands and riparian areas were successfully summarized by the clustering process and in the ordered data matrixes using species (indicators) that best represented the phenological progression in each upland association and in each riparian association (Tables 8, 95 12, 14, and 18). This process clarified the differences in upland and riparian phenologies. At the BG sites, only one riparian species flowered in early spring compared to 17% of the upland species. By early summer, only 50% of the riparian species had flowered compared to 63% of the upland species. In each riparian phenological group species dispersed fruit three weeks later than upland species in respective groups. At the SBS site, flowering times were more similar in the riparian areas and uplands than at the BG sites. As in the grasslands, species in each group dispersed fruit two to four weeks later in the riparian area than species in the upland groups. Phenological patterns also differed between upland associations. The BGxw deep swale and the BGxw and BGxw/h transition associations had the highest levels of soil moisture and delayed phenology compared to the respective groups of the dominant dry grassland association. In addition, many species common across the uplands flowered up to two weeks later growing in these associations. Changes in phenology at the SBS site related to tree canopy cover and to soil moisture. Species generally flowered earlier in the clear cut (low levels of canopy cover) than in the other upland forest associations (higher levels of canopy cover). Species growing in the open edge association (forest/riparian edge) and in the clearcut flowered and dispersed fruit at the same time in 1995. In 1996, with much higher water levels in the riparian area, phenologies in the edge association lagged two weeks behind the clearcut. Further studies need to be done to clarify how the delay in phenology of the transition areas of both study areas and the grassland's deep swale compared to the phenology of the dominant associations at each study area might relate to maintaining both species and habitat diversity in these two ecosystems. 96 This study has characterized phenological progression in riparian areas and associated uplands in the Junction grasslands and a Sub-Boreal Forest. It also showed that phenology differs between the plant associations that make up these ecosystems and that range readiness criteria can not be based on readiness of a single plant association. Understanding these phenological differences can aide in the development of range use plans that successfully meet many of the objectives stated in the Range Management and Riparian Management Area Guidebooks (B.C. Ministry of Forests 1995 a and b). 7 Management Recommendations 7.1 Phenological Summary of a Site as Part of a Range Management Plan This study described a method for identifying the phenological patterns that are found in an ecosystem with a variety of habitats including riparian areas. Understanding the phenological patterns of a range area is an important step in designing a grazing management plan that will maintain the diversity of habitats within the range area. I feel the method I used can be modified to take less time and still provide a phenological picture of the varied habitats within an area. Modifications could include: 1. Use a less detailed phenological code. The code could include the five main and easiest to observe developmental stages. For example: first growth, flower initiation, full flower, fruit dispersal, and dormancy. The manager should choose those stages that highlight developmental stages important to the site being managed (e.g., Fruit distribution in the riparian areas). 97 2. Focus on just a single stage of development. Recording only when each species reaches a single stage of development (usually first flower or full flower) can give a general picture of how the phenology varies in the associations in a site. 3: Record phenological information on only a few species in each association. Choose species with phenologies representative of the phenological patterns of the association. 4. Include any rare species that deserve special consideration. Relate their phenologies to the indicator plants chosen so as to have enough information to protect plants at vulnerable stages of development and to allow seed production to maintain their populations. 5. Choose indicators from both the dominant and rare species that can represent the phenological progression of only the main plant associations. This is usually the riparian and dry grassland associations in the grassland range and the riparian and open forest or clearcut associations in the forested range. These modifications can shorten the observation time needed to produce a phenological description of a management area, but because not all species respond in the same way to yearly climate changes, it is necessary to have two or more years of observations to choose plant indicators that consistently represent the phenology of a group of plants. These indicators can then be used to produce a phenological site summary like the ones of the BG and SBS sites in Table 20 and 21. 98 7.2 Using Phenology to Design Range Management Plans This study has shown that plant associations within a site have important differences in their phenological patterns. While this study did not focus on provincial guidelines for range readiness, the results point out that the present guidelines do not allow for this variation. The term readiness needs to be broadened to the seasonal time period when a site can be grazed with the least damage to the the health of all plant associations involved not just the time when grazing can begin. In order to define this idea of readiness using the phenology of plants, it is necessary to focus on the phenological patterns of all plant associations of a site. The phenology of the dominant association in the uplands of the grasslands is considered when choosing range readiness criteria and managing these areas. Plants of species growing in the dominant upland association flowered two weeks earlier than plants located in areas with higher levels of soil moisture. These more moist areas are often subjected to heavy cattle use (Nicholson et al. 1991), because the vegetation in these areas remains greener and more palatable later in the season and is usually near water. A plan based only on the species in the dominant dryer associations could place pressure on the other associations and negate the advantages of the extended growing season they offer to both common and rarer grassland species. To maintain diversity in the grasslands the phenologies of these habitat need to be considered. Patterns of use should allow for seed production in all associations in some years. This is especially important in very dry years when many plants may not reach ripe seed stage. Range readiness criteria in the SBS (commonly length of new growth of Kentucky bluegrass or pinegrass) are usually based on the phenology of species found in upland open areas (often in clearcuts). In 1996, in my study, the phenologies of the transition associations 99 at the edges of riparian areas lagged behind the clearcut by two weeks in this study. These associations are where the rarer species are found and where the forest species in general more consistently set fruit. Cattle utilize these associations more than other parts of forest range (Knezevich pers.comm.). It is important to design patterns of use that allow for seed production in these associations to maintain their diversity and the diversity of forested areas in general. In the SBS, this would require moving cattle out of these edge associations on a regular basis in late summer and fall. Range readiness criteria should be based on the phenology of species found in each major association in a range area and should identify readiness in each plant association. For example; in the BG sites, bluebunch wheatgrass in the transition and deep swale associations could be used as an indicator of spring readiness of a site where it is necessary to protect these areas. Brown-eyed Susan might be used to indicate late summer and fall readiness in these same associations in terms of allowing species to disperse seed (Table 20). In the SBS study area, mid to late summer use may be indicated by ripe fruit on twinberry bushes as an indicator that the early fruiting edge species are dispersing seeds. Ripe rose hips indicate that late fruiting edge species have dispersed fruit (Table 21). This study showed that the phenology of riparian areas is different from the phenology of the surrounding uplands. Grassland riparian species began growing later in the spring than species growing in the uplands, and more riparian species flowered mid to late summer compared to spring and early summer flowering in the uplands. SBS riparian and upland species did not show this difference in early growth and flowering phenology. Seed production was later in the riparian areas than in the upland areas of all sites. 100 Seed production in riparian areas is sporadic and tied to natural fluctuations in water levels. Many riparian species need a drought period to germinate seeds. These drought periods can be rare and seed must be available to take advantage of it. Any added stress to a plant including grazing pressure can interfere with its ability to produce seed in the shorter growing season typical of riparian areas. Since many riparian species reproduce vegetatively, an area may not show the affects of grazing on seed production. Colonies of riparian plants age and die, however, and new plants from seeds are needed to replace them and to maintain genetic diversity and the long term stability of the riparian associations. Wildlife dependent on riparian areas rely on seed production. Waterfowl and other birds and mammals eat the seeds of a wide variety of riparian species (Interior Wetlands Program 1998). In addition, healthy stands of emergent plants (dependent on seed production) provide important cover and nesting habitat to a wide variety of wildlife. The fact that riparian areas are especially sensitive to range use practices (MOF 1995b) together with the delayed phenology and importance of seed production means range plans need to be designed to specifically meet riparian area habitat diversity objectives. If this can not be done within the general range area plan, cattle use in riparian areas will need to be restricted (MOF 1995b). As suggested by Hooper and Pitt (1995), riparian areas within a range area need to be ranked based on their importance to plants and wildlife and to the habitat diversity of an area. Riparian areas with high importance will need some form of protection to restrict livestock access to those areas. Fencing so that only part of an area is used and allowing the rest to have a full season of growth may be one alternative. 101 Riparian areas ranked less important could be managed using adjustments of levels and/or timing of use. Spring grazing of riparian areas is often considered to cause less stress to riparian plants. It will, however, cause a delay in phenology which may result in no seed production that year. Therefore, repeated spring use of small riparian areas should be avoided. Deferred grazing until after seed ripens in uplands can result in over use of the associated riparian areas (Elmore 1989), little or no riparian seed production that year, and delayed growth the next season. In addition, many Carex species produce flowers on two-year-old tillers and only a small percentage of tillers survive naturally to produce flowers (Bernard and Gorham 1978). To lessen these affects, a grazing plan should allow years when the riparian area is rested from livestock grazing. In addition, time of use could change from one year to the next. In the grasslands, grazing could be deferred until after seed dispersal in the riparian areas. The alternative to timing changes is to keep level of use low enough that there are plants left ungrazed to produce seed each year. Plant indicator species for each site should be chosen for their consistent ability to indicate both early and late season range readiness within the riparian association. For example; flowering of many Carex spp. in spring and dispersal of fruit in the fall indicate readiness at both ends of the season. Season long continuous grazing should not be used in riparian areas (MOF 1995b), but it is used in the forested range found in the SBS. To prevent overuse of small riparian areas and to allow for seed production may require bringing cattle in from range earlier in the fall or consistently moving cattle to keep level of use low and to allow plants time for development and dispersal of fruit. 102 TABLE 20. Summary of the phenological progress, BGxw site, Junction Sheep Range Park. BC. 1996. Duration Phenological Date of Group flowering* or flowering fruit dispersal* begins Association Riparian Dry grassland shallow swalea Deep Swale EARLY SPRING April 21 April 30 May 7 May 14 "Salix spp SPRING May 21 May 28 EARLY June7 SUMMER June 14 June 21 MID June 28 SUMMER July 7 LATE SUMMER July 14 July 21 July 28 Aug 7 Aug 14 Aug 21 Aug 28 Sept 7 Mid-late Sept "Lomatium macrocarpum 'Taraxacum officinale Agropyron 18cm new growth 'Erigeron compositus Stioa 12 cm new growth "Amelanchier alnifolia 'Balsamorhiza sagittata "Carex praegracilis "Zigadenus venenosus "Penstemon procerus "Erigeron linerais Agropyron boot "Lithospermum ruderle "Smilacina stellata "Carex atherodes "Smilacina stellata "Erigeron flagellaris it-Taraxacum officinale "Koeleria macrantha "Agoseris glauca "Tragapogon pratensis "Linum perenne "Agropyron spicatum "Erigonum hercleoides "Stipa comata "Geranium viscosissimum "Gaillardia aristata "Koeleria macrantha Agropyron boot "Juncus Balticus "Koeleria macrantha "Agropyron spicatum "Calamagrostis stricta "Rosa woodsii "Achillea millefolium "Galium boreale "Allium cernuum "Agropyron spicatum "Artemisia campestris "Solidago candensis #Balsamorhiza sagittata 'Symphoricarpus albus "Mentha arvensis "Leymus cinerus "Aster ericoides "Sonchus asper #Koeleria macrantha ItCarex praegracilis ikluncus balticus #Leymus cinerus #Calamagrostis stricta #Agropyron spicatum #Carex atherodes #Sonchus asper HSmilacina stellata #Tragopogon pratensis #Aster ericoides "Calochortus macrocarpus itErigeron linearis #Lithospermum ruderle #Erigeron compositus #Stipa comata ttZigadensus venenosus #Agoseris glauca "Solidago spathulata #Agropyron spicatum "Chrysothamnus nauseosus MLinum perenne MKoeleria macrantha ttAnemone multifida "Artemisia frigida #Calochortus macrocarpus ^Achillea millefolium MTragopogon pratensis #Rosa woodsii #Erigonum hercleoides #Galium boreale #Artemisia frigida #Artemisia campestris "Aster campestris #Erigeron flagellaris #Stipa comata #Agropyron spicatum #Gaillardia aristata #Smilacina stellata #Achillea millefolium #Aster campestris * = 10-30% in flower; # = fruit dispersal a If a species is found in the dry grassland, shallow swale, and deep dates in all three associations, it is listed only once in this column. If swal  and has the same flowering and fruit dispersal the dates are different, it is listed in both columns. 103 TABLE 21. Summary of the phenological progress, SBS site, Horsefly, BC. 1996. Duration of flowering Phenological Group Date flowering* or fruit dispersal# begins Association Riparian Forested uplands Clearcut EARLY SPRING April 14 April 30 May 7 May 14 May 21 "Salix spp. 'Menyanthes trifoliata "Populous tremuloides "Calypso bulbosa "Petasites frigidus "Oryzopsis asperifolia "Arctostaphylos uva-ursi "Taraxacum officinale SPRING "Eriophorum angustifolium May 28 "Carex lasiocarpa "Betula glandulosa June 7 "Ranunculus uncinatus "Arctostaphylos uva-ursi "Viola adunca "Fragaria virginiana "Galium boreale EARLY SUMMER June 14 June 21 "Carex aquatilis "Agrostis scabra "Antennaria neglecta "Lonicera involucrata "Fragaria virginiana #Populous tremuloides "Smilacina stellata "Cornus canadensis "Aquilegia formosa "Vaccinium myrtilloides "Castilleja miniata "Arnica cordifolia "Antennaria microphyllla "Lupinus arcticus "Calamagrostis rubescens MID-SUMMER June 28 July 7 "Triglochin maritimum "Mimulus guttatus "Nuphar polysepalum July 14 July 21 "Potentilla palustris "Parnassia palustris "Glyceria borealis "Calamagrostis canadensis "Cirsium vulgare "Clintonia uniflora "Arnica cordifolia "Lilium columbianum "Lupinus arcticus "Vaccinium myrtilloides "Poa pratensis "Galium boreale "Castilleja miniata ttPetasites frigidus 'Rosa acicularis "Achillea millefolium "Poa pratensis "Hieracium albiflorum "Rosa acicularis it-Taraxacum officinale UFragaria virginiana "Epilobium angustifolium LATE SUMMER July 28 Aug 7 Aug 14 Aug 21 Aug 28 Sept 7 "Mentha arvensis "Stachys palustris "Polygonum amphibium #Menyanthes trifoliata #Ranunculus uncinatus #Carex lasiocarpa #Agrostis scabra #Triglochin maritimum #Glyceria borealis ttEriophorum angustifolium #Potentilla palustris "Calamagrostis rubescens "Aster ciliolatus "Solidago candensis #Fragaria virginiana #Lonicera involucrata  b H-Ameianchier alnifolia "Agoseris glauca "Aster conspicuus #Viola adunca #Shepherdia canadensis itCornus canadensis #Lupinus arcticus ^Arctostaphylos uva-ursi #Clintonia uniflora KAquilegia formosa #Galium boreale ULilium columbianum itSolidaao candensis ttAmelanchier alnifolia "Agrostis gigantea "Agoseris glauca "Phleum pratense "Aster conspicuus "Arnica cordifolia ttArctostaphylos uva-ursi it-Vaccinium myrtilloides #Rosa acicularis #Poa pratense #Epilobium angustifolium ttPhleum pratense #Calamaarostis rubescens * = 10-30% in flower; # = fruit dispersal a If a species is found in the forested, path, and clearcut associations and has the same flowering and fruit dispersal dates in all three associations, it is listed only once in this column. 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Ordered data matrixes for each plant association, BGxw and BGxw/h sites, Junction Sheep Range Park and SBS site, Horsefly, British Columbia, 1996 Notes: • Species in each matrix are placed in order of the week in 1996 that each species was 10-30% in flower. • Species are listed by their shortened code (see appendix 3) • The number following the species is the plant association the species was growing in. • The flowering period for each species is shaded. • The phenological code that the numbers refer to is on page 29 of the text and is included here for easier reference. 113 Phenological code 0 No new growth 1 First shoots showing 2 First leaves unfolding 3 2-3 leaves unfolded 4 Middle leaves open; adding leaves and size 5 Inflorescence recognizable; adding leaves and size 6 Mature flower buds; plant nearly full size 7 10-30% in bloom; plant full size 8 >30% in flower; leaves may just start to yellow 9 Flowers+faded flowers+green fruit 10 Green fruit, plant up to 50% yellow 11 Ripe fruit; plant up to 50% yellow 12 Dispersal and/or plant yellow>50% 13 Fruit dispersed; plant dead or dormant 14 Regrowth 114 o o t»| N r~..,t>| „,.£ in £ S o E E •a E I § c c (0 3 o o o o « p i 0) OJ CO <04 co.wco'coi co co-'r- r*-T t CD in N" T IO T T CM T CO CO O CM CM ^ n ^ 2 I s 11 W 05 O C o co ef o ra 'o 0 in •a co tjl a> < 1X1 i m < co co! n - 3 SJ 5 E | CM CN CM CN CN T - CN CN CN | C N O CN CM T - O O T - 0 ) ; O iO> 3 O O C> CO CO o o o|aco„,o> 0> CO CO 0> CO CO CO oo cojoj'OCrco co r-i: i ' tr , i r r 1 co m ID m ix •j T o "C v *^ *c O O O C N C M C O C O < 3 -O O O C N C N C N C N C O co V> P <2 co co ,8 g §•= 2 2 S2 '•e a " - D D . 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L O L n m C D C D C D C D ^ LO|N- h-^ • T i o c D c D i n i o ^ r ^ m i o • T T T N r ^Tmmmin ^ r ' « ru - ) \ r CD in in LO m m ^ CN in rr CN *<T mcNTrcOT^ r ^ CM co CM TJ-cn I O m CO CO O JO C Olffi <£ D_ O C £ C Jo 5 CO Qj (0 (0 CO *3 <*= • ^ • • ^ • ^ • C M C N ^ - '^J" CN ^  CN CN CO t T - n o o n ^ o ^ o o o o CN o o o ^ r o ^ r o o o CD i n IO CO (O 2 "C . £ « • § « <ecn.oce.gCg S E E S I « O 8 E £ O | I -G » E 5 = c f J ! = | l V > C 0 * 0 o " = 3 * 0 c 9- C CT £ " o > u o d cn — ^ ro nj co > ^ ^ c N i n ' ^ - c N ' ^ - ^ T ' ^ - ^ r T j -o ^ - c N ^ r ^ r c N c o ^ - ' ^ T r - T CMCOOCO-^-OCNCOCOCO^r O O O C O C N O C N C N O C N C O OOOOOO -^OOCNCN CD CO CD CO CD CD m CO <£ o 2 o o. ra c o c o-M is o £ « a 2 is = * :s So 8 m T ( D i n , < r m i r ) t \ r T ^ ^ - c N ^ ^ r i n ^ ^ r r^^ cNo^ rcNCNco-^ -^ C O C N O C N O C N O C O C N C O O O C N O O O C N OOOOCNOOOO CD CD CD ^ (O (D (O CD l l U l f l ' l l l co o. o i c cue: co "5 -C c o v r a c o c o c o c o c o Q . 131 APPENDIX 2. Hierarchial cluster tree diagrams for riparian and upland summaries, BGxw site, Junction Sheep Range Park and SBS site, Horsefly, British Columbia, 1996 Note: • Species are listed in code (see appendix 3). • The number following each species is the plant association the species was growing in. • Species placed in a different phenological group by their flowering dates are underlined. 132 ( s a l i s p e l ] ^ a r e p r a J ^ carepra4 c a r e a t h l smilste4 V. J I<rinuper4~^ linuper3 poapral juncbal4 j u n c b a l l ^ u n c b a l 3 ^ ''agrospi^ agrospi3 c a l a s t r l trappra3 hierumb3 elymcin4 c i r s a r v l I^spargra3^ ^ s o l i s p a 4 ^ mentarvl aste e r i 3 ^ a r t e f r i 4 ^ E a r l y Spring + - -+ -- - I I - - - - I + -Spring E a r l y Summer Mid-Summer Late Summer 3 . 940 0 . 624 1.467 1.247 12.066 0.000 1.333 3 .367 0 . 000 0 . 000 1.654 0 . 846 0 . 577 1.029 8.250 1. 027 1. 414 0 . 850 2 . 754 1.196 0 . 850 1.599 c l u s t e r diameters Figure A.2.1 Tree diagram from clustering of rip a r i a n indicators, Bgxw s i t e , Junction Sheep Range Park, B r i t i s h Columbia, 1996 1 3 3 j^andrsep"^ taraoff5 erigcom5 erigcom6 balssag7 lomamac5, V. J 'carepetSNj amelain7 - - I lithrud5 anemmul5 lithrud6 penspro5 anemmul6 anemul7 er i g l i n 5 zigaven5 zigaven7 zigaven6 ^agosglaS^ agosgla5 agosgla7 ^geravis7^ e r i g f l a 6 ^erigf la7 - I ++ Early Spring Spring Early Summer Spring erioher5 1 1 g a i l a r i 7 + | | + - j | tragpra6 -1 1 1 + J | astragr6 -+ | | j I I I 1 Early linuperS 1 1 1 1 -+ | | | I I I 1 linuDer6 I I I 1 0.816 1.865 0. 333 1. 091 1. 000 8 . 773 0 .471 2 .455 1. 041 0 .707 1 .282 0.703 0 . 000 0.904 1.938 0 . 875 0 .707 38 . 555 0 . 000 0 . 000 2.460 0. 889 0 . 000 4.806 0 . 577 0 .695 1. 373 1. 054 1. 121 Figure A.2.2 Tree diagram from clustering of upland indicators, BGxw site,Junction Sheep Range Park, B r i t i s h Columbia, 134 'stipcomS~^ ^ t i p c u r T ^ agrospi5 koelmacS koelmac7 agrospi6 > J rosawoo5 achimil7 galibor7 a l l i c e r 6 a l l i c e r 7 ^agrospi7J //calomac7~Ni calomac6 calomac5 siledru6 solican7 solispa5 asteeri6 asteeri7 artecam5 chrynau5 a r t e f r i 7 a r t e f r i 5 Mid-Summer Early Summer Mid-Summer Late Summer 3 . 745 0 . 000 1. 877 0.600 0.408 0. 698 3 .150 0 . 667 0 . 816 1 .326 0 . 527 0 . 737 8 . 830 0 . 000 1. 133 1.587 22.225 0.626 0 . 517 0.333 0.818 2 . 773 0 . 979 0 . 544 0 . 000 .artefri6 c luster diameters Figure A.2.2 (Continued) Tree diagram from clustering of upland indicators, BGxw site, Junction Sheep Range Park, B r i t i s h Columbia, 1996 135 r . ^ m e n y t r i l erioang2 salispe2 carechol carelas2 \ c a r e d i a l j ^urtidic-2^ t r i g r a a r l careaqul careaqu2 ( / g l y c b o r l ^ potepal2 calacan2 mimugut2 ^ c i r s v u l 2 y polyampl mentarv2 — I + " I — I I + - -+ - -I H I I Spring 1.312 2 . 003 0 . 745 1.277 0.G24 S . 891 0 . 524 2 .165 | 0.235 I | 2.468 I + 14.649 I | 0.881 |Mid-Summer | 0.745 I | E a r l y Summer 2 . 260 1.128 0 . 527 C l u s t e r Diameters Late Summer Figure A.2.3 Tree diagram from c l u s t e r i n g of r i p a r i a n i n d i c a t o r s , SBS s i t e . Horsefly, B r i t i s h Columbia, 1996 1 3 6 r ~\ poputre5 calybul5 petafri3 petafri4 V. J f ~\ fragvir6 < taraoff6 violadu5 oryzasp4 arctuva6 amalain5 V. J ^ragvirS^ anteneg5 antemic6 ^astmin^y ''loniinv^ 1 loniinv4 vacimyr5 betugla3 violadu4 V J /agrosca!^ arnicor5 lupiarc5 vacimyr4 smilste3 aquifor6 lupiarc4 hieralb6 - I - I Early Spring Spring Early Summer Spring Early Summer 1.682 1. 062 0.707 4 . 116 0 . 908 0.667 1.121 0 . 624 1.529 36.607 0 . 577 1.200 0.782 3 . 064 0 . 000 0 .889 1.736 0.850 8 .458 1.225 0 .408 0.830 1.671 0. 957 0. 624 1. 002 2 . 661 Figure A.2.4 Tree diagram from c l u s t e r i n g of upland ind i c a t o r s , SBS s i t e . Horsefly, B r i t i s h Columbia, 1996 137 /^corncan4\ clinuni5 cornser5 ^ l i l i c o l ^ /^chimi!^ galibor3 l i l i c o l 6 l i l i c o l 5 rosasp3 rosaasi5 elymgla6 achimil6 calarub6 qaliborG poapra6 linnbor4 , pyrochl4 C J asteciaS^ 1 astecon4 soldcan5 agrogig5 calarub5 calarub3 calarub4 phlepra6 epilana^; /'hieralb5\ juncvas3 melalin3 agosala4 parnpal3 parnpal5 . - - I + - I -II + -- I + + -I -- I Mid-Summer I i I Mid-Summer Late Summer Mid-Summer 0 . 707 0. 952 1. 030 5. 990 0 . 943 1.705 0 . 000 1.332 0.236 0. 778 2 . 747 0 . 624 1.356 0 . 707 1. 969 1.528 21.437 0 . 972 0 . 913 1.185 2 .602 0. 000 0 . 000 2 . 070 0 . 943 4 .312 0 . 707 1. 606 0. 913 2 .317 0.000 cluster diameters Figure A.2.4 (Continued) Tree diagram from c l u s t e r i n g of upland indi c a t o r s , SBS s i t e . Horsefly, B r i t i s h Columbia, 1996 138 Append ix 3 Plant spec ies found in each plant associat ion BGxw, B G x w / h si tes , Junct ion S h e e p Range Park and S B S site, Horsef ly, Brit ish Co lumb ia , 1996 B G Si tes T a x o n o m i c N a m e C o m m o n N a m e Plant Assoc ia t ions Code Achilea millefolium yarrow 4,5,6,7 achimi l Agoserus glauca pale agoser is 5,6,7 agosgla Agropyron cristatum crested wheatgrass 7b agrocr i Agropyron repens quack grass 4 agrocr i (Elymus repens) Agropyron spicatum bluebunch wheatgrass 3,4,5,6,7 agrospi (Elymus spicata) Agropyron trachycaulum s lender wheatgrass 3,4,6,7 agrotra (Elymus trachycaulus) Agrostis scabra hair bentgrass 4 agrosea Allium cernuum nodding onion 5,6,7 al l icer Alopecurus pratensis m e a d o w foxtai l 1.2 a lopaqu Amelanchier alnifolia Saskatoon 7 amela ln Androsace septentrionalis northern fa i ry -candelabra 4,5 andrsep Anemone multifida cut - leaved a n e m o n e 5,6,7 a n e m m u Antenaria species 5,7b antenna (microphyla or umbrinella) rosy or u m b e r pussytoes Antenaria parvifolia Nuttal l 's pussytoes 4,5,6,7 an tepar Antennaria dimorpha low pussytoes 5,7b an ted im Arabis holboellii Hoelboel l 's rockcress 5,6,7 arabhol Arabis species arabis species 5,6 arabis Arctostaphylos uva-ursi kinnik innick 7 a rc tuva Artemisia campestris northern w o r m w o o d 5,6 a r tecam Artemisia frigida pasture sage 4,5,6,7 artefr i Artemisia tridentata big sagebrush 5 artetri Aster campestris m e a d o w aster 7a as tecam Aster ericoides ssp pansus tuf ted whi te prair ie aster 3,4,6,7 asteer i Astragalus agrestis f ie ld mi lk -ve tch 6,7 astragr Astragalus miser t imber mi lk -ve tch 6,7 ast rmis Balsamorhiza sagittata arrow leaved ba lsamroot 7a balssag Bromus anomalus nodding b rome 7a b romano Bromus tectorum cheatgrass 7b b romtec Calamagrostis stricta s l ims tem reedgrass 1,2,3 calastr Calochortus macrocarpus sagebrush Mar iposa lily 5,6,7 c a l o m a c Carex atherodes awned sedge 1,2,3 careath Carex eleocharis need le- leaved sedge 5,6,7b careele Carex exsciccata inf lated sedge 1,2,3 careeexs Carex lasiocarpa s lender sedge 1,2 care las Carex petasata pasture sedge 5,6,7 carepet Carex praegracilis f ie ld sedge 2,3,4 carepre Chrysothamnus nauseosus c o m m o n rabbi t -brush 5 chrynau Cirsium hookerianum Hooker 's thist le 5,6,7 cirshoo 1=open wate r associat ion, 2=wet r iparian associat ion, 3=dry r iparian associat ion, 4=transi t ion associat ion, 5=dry grassland associat ion, 6=sha l low swale associat ion 7a=deep swale associat ion B G x w site, 7b=deep swale associat ion B G x w / h site T a x o n o m i c N a m e C o m m o n N a m e Plant Assoc ia t ions Code Cirsium undulatum w a v y - l e a v e d thist le 5,6,7 c i rsund Comandra umbellata pale c o m a n d r a 5,6 c o m a u m b Crepis atrabarba s lender hawksbeard 5,6 crepatr Crepis tectorum annual hawksbeard 3,4,5,7 c reptec Descurainia sophia f l i xweed 1,2 descsop Eleocharis palustris c o m m o n spike-rush 1 e leopal Epilobium ciliatum purp le- leaved wi l lowherb 2 epilci l Erigeron compositus cut - leaved daisy 5,6,7 e r igcom Erigeron flagellars t rai l ing f leabane 5,6,7 er igf la Erigeron linearis l ine- leaved daisy 5,6 erigl in Erigeron speciosus showy f leabane 7a er igspe Eriogonum heracleoides sulphur buckwheat 5,6,7 er ioher Festuca altaica northern rough fescue 3,4,5,7 festalt Fragaria virginiana wild s t rawberry 7 f ragv i r Gaillardia aristata brown-eyed Susan 5,6,7 gai lar i Galium boreale northern bedst raw 5,6 gal ibor Geranium viscosissimum st icky purple ge ran ium 7a gerav is Geum triflorum old man 's whiskers 5,6 geumtr i Heuchera cylindrica round- leaved a lumroot 6 heuccy l Hieracium umbellatum nar row- leaved hawkweed 3 h ie rumb Hordeum jubatum foxtai l bar ley 2,3,4 hord jub Juncus balticus balt ic rush 2,3,4,7a juncba l Koeleria macrantha Junegrass 4,5,6,7 koe lmac Lactuca tatarica blue lettuce 2,3 lacttat Lappula redowskii western s t ickseed 5,6 lappred Lepidium densiflorum prairie pepper-grass 5,6 lepiden Leymus cinereus giant wi ldrye 4 leymcin Linum perenne western blue f lax 3,5,6,7 l inuper Lithospermum ruderale l emonweed 5,6,7 l i thrud Lomatium macrocarpum large-frui ted deser t -pars ley 5,6,7b l o m a m a c Mentha arvensis f ie ld mint 1,2,3 men ta rv Orthocarpus luteus yel low owl -c lover 4,5,6 orthlut Penstemon procerus smal l - f lowered pens temon 5,6 penspro Phacelia linearis th read- leaved phacel ia 7b phacl in Phalaris arundinacea reed canarygrass 2,3 phalaru Poa palustris fowl b luegrass 2,3,4 poapal Poa pratensis Kentucky b luegrass 3,4,6,7a poapra Poa secunda alkali b luegrass 5,7 poasec Polygonum amphibium water smar tweed 1,2 po l yamb Populous tremuloides t rembl ing aspen 7a poputre Potentilla anserina s i lverweed 3 poteans Potentilla pensylvanica prair ie c inquefo i l 5,6 potepen Pseudotsuga menziesii Douglas- f i r 7 pseumen Ranunculus cymbalaria shore but tercup 1,2,3 ranucym Ranunculus sceleratus ce lery- leaved but tercup 1,2 ranusee 1=open wate r associat ion, 2=wet r iparian associat ion, 3=dry r iparian associat ion, 4=transi t ion associat ion, 5=dry grassland associat ion, 6=sha l low swale associat ion 7a=deep swale associat ion B G x w site, 7b=deep swale associat ion B G x w / h site 140 T a x o n o m i c N a m e C o m m o n N a m e Plant Assoc ia t ions Code Rosa woodsii/acicularis Prickly/prair ie rose 4,5,6,7 rosawoo Salix species wi l low spec ies 1,2 sal ispe Scirpus validus so f t - s temmed bulrush 1,2 sc i rval Sedum lanceolatum lance- leaved s tonecrop 5 sedulan Senecio canus wooly groundsel 7b senecan Silene drummondii D r u m m o n d ' s camp ion 6 si ledru Sisyrinchium montanum mounta in b lue-eyed grass 4,6 s i symon Smilacina stellata star- f lowered Solomon's-sea l 3,4,7a smi ls te Solidago canadensis C a n a d a go ldenrod 7a sol ican Solidago spathulata dune go ldenrod 3,4,5,6,7 sol ispa Sonchus asper prickly sow-thist le 3 soncasp Spartina gracilis alkali cordgrass 3,4 spargra Stachys palustris s w a m p hedge-net t le 1,2,3 stacpal Stipa comata needle-and- thread grass 5,6,7 s t ipcom Stipa curtiseta shor t -awned porcup inegrass 6,7 st ipcur Symphoricarpos albus c o m m o n snowberry 5,6,7 s y m p a l b Taraxacum officinale c o m m o n dandel ion 4,5,6,7 taraof f Tragopogon dubius yel low salsify 3,4,5,6,7 t ragdub Tragopogon pratensis m e a d o w salsi fy 3,4,5,6,7 t ragpra Verbascum thapsus c o m m o n mul le in 6 verb tha Vicia americana A m e r i c a n ve tch 3 v i c i a m e Zygadenus venenosus m e a d o w death c a m u s 5,6 z igaven 1=open water assoc ia t ion, 2=wet r iparian associat ion, 3=dry r iparian assoc ia t ion, 4=t ransi t ion associat ion, 5=dry grassland associat ion, 6=sha l low swale associat ion 7a=deep swale associat ion B G x w site, 7b=deep swale associat ion B G x w / h site 141 S B S site T a x o n o m i c N a m e C o m m o n N a m e Plant Assoc ia t ions Code Achillea millefolium yarrow 3,4,5,6, ach imi l Agoseris glauca pale agoser is 3,4,5,6 agosgla Agrostis gigantea redtop 5,6 agrogig Agrostis scabra hair bentgrass 3,5 agrosea Alnus sp. alder species 3,4,5,6 a lnus Amelanchier alnifolia Saskatoon 3,5,6 ame la ln Anaphalis margaritacea pearly ever last ing 5,6 a n a p m a r Antennaria microphylla rosy pussytoes 5,6 an temic Antennaria neglecta f ield pussytoes 4,5,6 anteneg Aquilegia formosa red c o m u m b i n e 3,5,6 aqu i for Aralia nudicaulis wild sarsapar i l la 5 ara lnud Arctostaphylos uva-ursi kinnik innick 3,4,5,6 arc tuva Arnica cordifolia hear t - leaved ar in ica 3,4,5,6 arnicor Arnica latifolia mounta in arn ica 5 arni lat Aster ciliolatus f r inged aster 3,4,5,6 asteci l Aster conspicuus showy aster 3,4,5,6 astecon Betula glandulosa scrub birch 2,3 betugla Brachythecium species ragged mosses 3,4,5,6 Calamagrostis canadensis bluejoint reedgrass 2,3 ca lacan Calamagrostis rubescens pine grass 3,4,5,6, ca larub Calypso bulbosa fa i rysl ipper 5 calybul Carex aquatilis water sedge 1,2 careaqu Carex canescens grey sedge 2,3 carecan Carex chordorrhiza cordroot sedge 1,2 carecho Carex concinnoides northwestern sedge 5 carecoc Carex diandra lesser panic led sedge 1,2,3 caredia Carex interior in land sedge 2,3 careint Carex lasiocarpa s lender sedge 1,2 care I as Carex limosa shore sedge 1 care l im Carex utriculata beaked sedge 1,2 careutr Castilleja miniata c o m m o n red paintbrush 3,4,5,6 cas tmin Chimaphila umbellata pr inces pine 4,5 c h i m b u m b Cirsium vulgare bull thist le 2,3 c i rsvul Clintonia uniflora • queen's cup 5 cl inuni Coeloglossum viride long-bracted f rog orchid 3,5 coe lv i r Cornus canadensis bunchberry 3,4,5 co mean Cornus sericea red osier dogwood 5,6 cornser Crepis tectorum annual hawksbeard 5,6 creptec Drepanocladus aduncus c law- leaved fea ther moss 1,2 Elymus glaucus blue wi ldrye 5,6 e lymg la Epilobium angustifolium f i reweed 5,6 epi lang Epilobium ciliatum purp le- leaved wi l lowherb 2,3 epilci l Equisetum species horsetai l 2,3 equiset Eriophorum angustifolium narrow- leaved cot ton-grass 1,2 er ioang 1=open wate r associat ion, 2=r ipar ian associat ion, 3=open edge associat ion, 4=forest associat ion, 5=path associat ion, 6=c learcut associat ion 142 T a x o n o m i c N a m e Festuca occidentalis Fragaria virginiana Galium boreale Galium triflorum Geocaulon lividum Geum macrophyllum Glyceria borealis Glyceria grandis Goodyera repens Hieracium albiflorum Hieracium gracile Juncus vaseyi Lathyrus nevadensis Lathyrus ochroleucus Lilium columbianum Linnaea borealis Lonicera involucrata Lupinus arcticus Melampyrum lineare Mentha arvensis Menyanthes trifoliata Mimulus guttatus Nuphar polysepalum Orthilia secunda Oryzopsis asperifolia Oryzopsis pungens Osmorhiza chilensis Parnassia palustris Peltigera aphthosa Petasites frigidus varpalmates Phleum pratense Picea engelmanniiXglauca Platanthera dilatata Platanthera obtusata Poa palustris Poa pratensis Polygonum amphibium Polytrichum species Polytrichum strictum Populus tremuloides Potentilla palustris Pyrola asarifolia Pyrola chlorantha Ranunculus uncinatus Rosa woodsii/acicularis Rubus pubescens C o m m o n N a m e western fescue wi ld s t rawberry northern bedst raw sweet -scented bedst raw bastard toad- f lax la rge- leaved a v e n s northern mannagrass reed m a n n a g r a s s dwar f rat t lesnake orchid whi te hawkweed s lender hawkweed Vasey 's rush purple peav ine c reamy peav ine t iger lily tw in f lower black twinberry arct ic lupine cow-wheat f ie ld mint buckbean ye l low monkey - f l ower Rocky Mounta in cow-l i ly one-s ided win tergreen rough- leaved r icegrass shor t -awned r icegrass mounta in sweet-c ice ly northern grass-of -parnassus f reck led l ichen sweet col tsfoot c o m m o n T i m o t h y enge lmann/wh i te spruce whi te bog orchid one- leaved rein orchid fowl b luegrass Kentucky b luegrass wate r smar twed hair c a p spec ies bog hair -cap t rembl ing aspen marsh c inquefoi l pink win tergreen green win tergreen smal l - f lowered but tercup prair ie/pr ickly rose trai l ing raspberry Plant Assoc ia t ions Code 5 fes tocc 3,5,6 f ragv i r 3,4,5,6 gal ibor 3 gal i tr f 3.4.5 geoc l iv 5 g e u m m a c 1,2 g lycbor 2 g lycgra 4,5 goodrep 3,5,6 h ieralb 3 hiergra 3 j u n c v a s 3,4,5,6 la thnev 3,4,5,6 lathoch 5,6 li l icol 3,5,6 l innbor 2,3,4,5 loni inv 4,5,6 lupiarc 3,5 mela l in 2 men ta rv 1,2 menyt r i 2,3 m i m u g u t 1 nuphpol 4,5 or thsec 4,5 oryzasp 5 oryzpun 5 osmoch i 2,3,5 parnpal 3,4,5,6 3,5 petapal 6 phlepra 4,5,6 p iceax 3.5 platdil 3,5 platobt 2,3,5 poapal 5,6 poapra 1,2 po l yamp 3,4,5,6 1,2 3,4,5,6 poputre 1,2 potepal 4,5 pyroasa 4,5 pyrochl 3,5 ranuunc 3,4,5,6 rosawoo 3,4,5,6 rubupub 1=open water associat ion, 2=r ipar ian associat ion, 3=open edge associat ion, 4=forest associat ion, 5=path associat ion, 6=c learcut associat ion 143 T a x o n o m i c N a m e C o m m o n N a m e Plant Assoc ia t ions Code Salix species wi l low species 2,3 sal ispe Scutellaria galericulata marsh skul lcap 3 scutgal Senecio pauperculus Canad ian but terweed 5 senepau Shepherdia canadensis soopolal l ie 5 shepcan Smilacina racemosa false So lomon 's seal 5 smi l rac Smilacina stellata star- f lowered So lomon 's seal 2,3,5,6 smi ls te Solidago canadensis Canada go ldenrod 5,6 so l ican Solidago spathulata dune go ldenrod 5 sol ispa Sphagnum angustifolium yel low-green peat moss 1,2 spirbet Spiraea betulifolia bi rch- leaved spi rea 5 Stachys palustris s w a m p hedge-net t le 1,2 stacpal Stellaria longipipes long-stalked starwort 5 stel log Symphoricarpos albus c o m m o n snowberry 5 sympa lb Taraxacum officinale c o m m o n dandi l ion 6 taraof f Thalictrum occidentale western meadowrue 5 tha locc Trifolium repens whi te c lover 3,5,6 t r i f rep Triglochin maritimum sea-side arrow-grass 1,2 t r i gmar Trisetum spicatum spike t r ise tum 5,6 tr isspi Urtica dioica st inging nettle 2 urt idio Utricularia intermedia f la t - leaved bladderwort 1 utri int Vaccinium myrtilloides ve lve t - leved blueberry 3,4,5,6 v a c i m y r Viburnum edule high-bush cranberry 5 v ibuedu Vicia americana A m e r i c a n ve tch 6 v i c i a m e Viola adunca early blue v io let 5 v io ladu 1=open wate r associat ion, 2=r ipar ian associat ion, 3=open edge associat ion, 4=forest associat ion, 5=path associat ion, 6=c learcut associat ion 144 

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