LIBRARY Michigan State University PLACE II RETURN BOX ‘0 roman this chockout from your "cord. TO AVOID FINES mum on or baton duo duo. WHITE-TAILED DEER MOVEMENTS, HABITAT USE, AND BROWSING EFFECTS ON VEGETATION IN THE UPPER PENINSULA OF MICHIGAN By Teresa Mackey A THESIS Submitted to Michigan State University in partial fulfillment of the requirements for the degree of MASTER OF SCIENCE Department of Fisheries and Wildlife 1996 ABSTRACT WHITE-TAILED DEER MOVEMENTS, HABITAT USE, AND BROWSING EFFECTS ON VEGETATION IN THE UPPER PENINSULA OF MICHIGAN By Teresa Mackey Habitat use and movement patterns of 61 white-tailed deer (W W) were monitored in the Whitefish River Basin (WRB) and Stonington Peninsula (SP) in the Hiawatha National Forest (l-INF) during 1993 and 1994. Home ranges were calculated. Vegetation types used by deer were compared to availability determined with Landsat thematic mapper data and ARC/INFO. Relative productivity of deer in the 2 study areas was compared. A long-term exclosure study was initiated to quantify the effects of deer on the northern white-cedar (Ihuja occidentafis) forest type; baseline vegetation characteristics of the cedar stands were measured. Spring/summer mean home range size for WRB and SP deer was 640.9 ha and 89.8 ha, respectively. Vegetation types were not used in proportion to availability; selected types were aspen/birch, mixed pine, and white- cedar (Ihm'a W3). Types with high percentages (>15%) of use included northern hardwoods, wet hardwood/conifer mix, and lowland conifers. Productivity estimates . were not different (P>0.10) between the 2 study areas. Vegetation types selected by deer should be maintained throughout the landscape to help reduce the possibility of high concentrations of deer and possible impacts on plant communities. ACKNOWLEDGMENTS Thanks to ...... McIntire-Stennis Cooperative Forestry Research Program, US. Forest Service, U.P. Whitetails Association, and the Michigan Agricultural Experiment Station for funding this project. The idea, without the money, would not have gone anywhere. Thanks to ...... my major professor, Dr. Henry Campa III, for his advice and guidance throughout the study. I would also like to recognize my committee members for their help during the project: Dr. Jonathan Haufler, Dr. Scott Winterstein, Dr. Kurt Pregitzer during the initial part of the project, and Dr. Doug Lantagne in the concluding portions of the study. Thanks to......U.S. Forest Service personnel on the Rapid River Ranger District of the Hiawatha National Forest. Thanks to ...... Michigan Department of Natural Resources, Wildlife Division, for providing Landsat thematic mapper satellite vegetation data files. Thanks to ...... Timothy Van Deelen and UP. Whitetails Association volunteers for trapping and radio-collaring the deer; Jason Halifax, Sharie DeLong, and Paul Thompson for assistance in data collection; Bryan Smith for help with ARC/INFO analysis. Thanks to ...... Dr. Scott Winterstein for help in statistical analyses. iii Thanks to ...... my fellow grad students for their help, input, advice, humor, and friendship, especially my office mates: Tim Van Deelen, Meg Clark, John Stribley, and Allison Gorrnley. Thanks to ..... my parents for all their support over the years; words do not seem to be enough to show my appreciation. Thanks to ...... my husband for all his support, patience, encouragement, and love (and for catching those U.P. trout for dinner when he would visit during my field seasons). Thanks to ...... God for allowing it all to happen. iv TABLE OF CONTENTS LIST OF TABLES ............................................................................................................... vii LIST OF FIGURES .............................................................................................................. xii INTRODUCTION .................................................................................................................. 1 OBJECTIVES ......................................................................................................................... 5 STUDY AREA ...................................................................................................................... 6 METHODOLOGY ............................................................................................................... 12 Capturing and Radio-Collaring ................................................................................ 12 General Location Technique ................................................................................... 12 Movements and Home Ranges ................................................................................ l4 Habitat Use ............................................................................................................... 15 Productivity .............................................................................................................. l8 Vegetation Sampling ................................................................................................ l9 Composition and Structure of Northern White-Cedar Stands ................... 19 Browse Sampling ........................................................................................ 22 RESULTS ............................................................................................................................. 23 Capturing and Radio-Collaring ................................................................................ 23 Movements and Home Ranges ................................................................................ 23 Adaptive Kernel Home Range Results .................................................... 25 Minimum Convex Polygon and Harmonic Mean Home Range Results ................................................................... 25 Habitat Use .............................................................................................................. 31 Productivity ............................................................................................................. 35 Vegetation Sampling ............................................................................................... 35 Composition and Structure of Northern White-Cedar Stands ................... 35 Exclosure and Open Area Site Data Combined ............................. 37 Exclosure and Open Area Site Data Separated .............................. 48 Browse Sampling ........................................................................................ 57 DISCUSSION ...................................................................................................................... 58 Movements and Home Ranges ................................................................................ 58 Home Range Estimator Comparison .......................................................... 6O Habitat Use ............................................................................................................... 61 Productivity ............................................................................................................. 64 Vegetation Sampling ................................................................................................ 65 Composition and Structure of Northern White-Cedar Stands ................... 65 Exclosure and Open Area Site Data Combined ............................. 66 Exclosure and Open Area Site Data Separated .............................. 67 Browse Sampling ........................................................................................ 67 CONCLUSIONS .................................................................................................................. 69 MANAGEMENT RECOMMENDATIONS ...................................................................... 71 APPENDIX .......................................................................................................................... 73 LITERATURE CITED ...................................................................................................... 125 vi Table 1 Table 2 Table 3 Table 4 Table 5 Table 6 Table 7 LIST OF TABLES Vegetation type classifications for Michigan's central Upper Peninsula, 1993-1994 ........................................................................ 17 Locations of exclosure and areas open to browsing sites in the Whitefish River Basin-North and -South (WRB-North and -South) and Stonington Peninsula-North and -South (SP-North and -South) study areas in the Hiawatha National Forest in Michigan's Upper Peninsula, 1993-1994 ........................................................................ 20 Number of white-tailed deer radio-collared in the Whitefish River Basin (WRB) and the Stonington Peninsula (SP) in the Hiawatha National Forest in Michigan’s Upper Peninsula, 1992-1994 ............ 24 Mean spring/summer home ranges (ha) (and standard errors) of white-tailed deer in the Whitefish River Basin (WRB) and Stonington Peninsula (SP) using adaptive kernel with 95% contours in the Hiawatha National Forest in Michigan's Upper Peninsula, 1993-1994 ........................................................................ 26 Mean fall home ranges for white-tailed deer in the Whitefish River Basin (WRB) and Stonington Peninsula (SP) using adaptive kernel with 95% contours in the Hiawatha National Forest in Michigan's Upper Peninsula, 1993-1994 ............................................................. 27 Mean spring/summer home ranges (ha) (and standard errors) of white-tailed deer in the Whitefish River Basin (WRB) and Stonington Peninsula (SP) using minimum convex polygon (MCP) and harmonic mean (HM) with 95% contours in the Hiawatha National Forest in Michigan’s Upper Peninsula, 1993-1994 ............ 29 Mean fall home ranges (ha) (and standard errors) of white-tailed deer in the Whitefish River Basin (WRB) and Stonington Peninsula (SP using minimum convex polygon (MCP) and harmonic mean with 95% contours in the Hiawatha National Forest in Michigan’s Upper Peninsula, 1993-1994 ........................... 30 vii Table 8 Table 9 Table 10 Table 11 Table 12 Table 13 Table 14 Table 15 White-tailed deer spring/summer habitat use and availability in the Whitefish River Basin (WRB) and Stonington Peninsula (SP) study areas in the Hiawatha National Forest in Michigan's Upper Peninsula, 1993-1994 ........................................................................ White-tailed deer fall habitat use in the Whitefish River Basin (WRB) and Stonington Peninsula (SP) study areas in the Hiawatha National Forest in Michigan's Upper Peninsula, 1993-1994 ............. Vegetation types with highest percentage of use by white-tailed deer by time period in the Whitefish River Basin and Stonington Peninsula study areas in Michigan’s Upper Peninsula, 1993-1994.. Mean productivity and standard error (SE) of white-tailed deer in the Whitefish River Basin (WRB) and Stonington Peninsula (SP) study areas in the Hiawatha National Forest in Michigan's Upper Peninsula, 1994 ................................................................................. Mean percent vertical cover (and standard error) for height strata in the 4 study areas (Whitefish River Basin-North and -South [WRB-North and -South] and Stonington Peninsula-North and -South [SP-North and -South]) in the Hiawatha National Forest in Michigan's Upper Peninsula, 1993-1994 ........................................... Mean percent vegetation horizontal cover (and standard error) for height strata in the 4 study areas (Whitefish River Basin-North and -South [WRB-North and -South] and Stonington Peninsula-North and -South [SP-North and -South]) in the Hiawatha National Forest in Michigan's Upper Peninsula, 1993-1994 ........................... Mean stem densities per hectare (and standard error) of dominant tree species in the 4 study areas (Whitefish River Basin-North and -South [WRB-North and -South] and Stonington Peninsula-North and -South [SP-North and -South]) in the Hiawatha National Forest in Michigan's Upper Peninsula, 1993-1994 ........................... Mean stem densities per hectare (and standard error) of non- dominant woody species in the 4 study areas (Whitefish River Basin-North and -South [WRB-North and -South] and the Stonington Peninsula-North and -South [SP-North and -South]) in the Hiawatha National Forest in Michigan's Upper Peninsula, 1993- 1994 ......................................................................................... viii 32 33 34 36 38 39 4o 41 Table 16 Table 17 Table A1 Table A2 Table A3 Table A4 Table A5 Table A6 Herbaceous species summary for the 4 study areas (Whitefish River Basin-North and -South [WRB-North and ~South] and Stonington Peninsula-North and -South [SP-North and -South]) in the Hiawatha National Forest in Michigan's Upper Peninsula, 1993-1994 .......................................................................................... Mean relative frequencies (and standard errors) of herbaceous species in the 4 study areas (Whitefish River Basin-North and -South [WRB-North and -South] and the Stonington Peninsula- North and -South [SP-North and -South]) in the Hiawatha National Forest in Michigan's Upper Peninsula, 1993-1994 ............................... Seasonal home ranges (ha) of white-tailed deer in the Whitefish River Basin (WRB) and Stonington Peninsula (SP) using minimum convex polygon (MCP), harmonic mean (HM) with 95 % contours, and adaptive kernel (AK) with 95% contours in the Hiawatha National Forest in Michigan's Upper Peninsula, 1993 ...... Seasonal home ranges (ha) for white-tailed deer in the Whitefish River Basin (WRB) and Stonington Peninsula (SP) using minimum convex polygon (MCP), harmonic mean (HM) with 95% contours, and adaptive kernel (AK) with 95% contours in the Hiawatha National Forest in Michigan's Upper Peninsula, 1994 ...... Mean percent vertical cover (and standard error) for height strata in exclosure and areas open to browsing sites in the Hiawatha National Forest in Michigan's Upper Peninsula, 1993-1994 ............. Mean percent horizontal cover (and standard error) for height strata in exclosure and areas open to browsing sites in the Hiawatha National Forest in Michigan's Upper Peninsula, 1993- 1994 ................................................................................................... Mean stem densities per hectare (and standard error) of dominant tree species in exclosure and areas open to browsing sites in the 4 study areas (Whitefish River Basin-North and -South [WRB-North and -South] and the Stonington Peninsula-North and -South [SP-North and -South]) in the Hiawatha National Forest in Michigan's Upper Peninsula, 1993-1994 ........................................... Mean stem densities per hectare (and standard error) of non- dominant woody species in exclosure and areas open to browsing sites in the Whitefish River Basin-North study area in the Hiawatha National Forest in Michigan's Upper Peninsula, 1993- 1994 ................................................................................................... ix 49 50 73 77 81 82 83 85 Table A7 Table A8 Table A9 Table A10 Table A1 1 Table A12 Table A13 Table A14 Mean stem densities per hectare (and standard error) of non- dominant woody species in exclosure and areas open to browsing sites in the Whitefish River Basin-South study area in the Hiawatha National Forest in Michigan's Upper Peninsula, 1993- 1994 ................................................................................................... Mean stem densities per hectare (and standard error) of non- dominant woody species in exclosure and areas open to browsing sites in the Stonington Peninsula-North study area in the Hiawatha National Forest in Michigan's Upper Peninsula, 1993-1994 ............. Mean stem densities per hectare (and standard error) of non- dominant woody species in exclosure and areas open to browsing sites in the Stonington Peninsula-South study area in the Hiawatha National Forest in Michigan’s Upper Peninsula, 1993-1994 ............. Mean stem densities per hectare (and standard error) of non- dominant woody species that were significantly different (P<0.10) among study areas (Whitefish River Basin-North and -South [WRB-North and -South] and Stonington Peninsula-North and -South [SP-North and -South]) in the Hiawatha National Forest in Michigan's Upper Peninsula, 1993-1994 ........................................... Mean absolute (AF) and relative frequencies (RF) (and standard errors [SE]) of herbaceous species in exclosure and areas open to browsing sites in the Whitefish River Basin-North study area in the Hiawatha National Forest in Michigan's Upper Peninsula, 1993-1994 .......................................................................................... Mean absolute (AF) and relative frequencies (RF) (and standard errors [SE]) of herbaceous species in exclosure and areas open to browsing sites in the Whitefish River Basin-South study area in the Hiawatha National Forest in Michigan's Upper Peninsula, 1993-1994 .......................................................................................... Mean absolute (AF) and relative frequencies (RF) (and standard errors [SE]) of herbaceous species in exclosure and areas open to browsing sites in the Stonington Peninsula-North study area in the Hiawatha National Forest in Michigan's Upper Peninsula, 1993- 1994 ................................................................................................... Mean absolute (AF) and relative frequencies (RF) (and standard errors [SE]) of herbaceous species in exclosure and areas open to browsing sites in the Stonington Peninsula-South study area in the Hiawatha National Forest in Michigan's Upper Peninsula, 1993- 1994 ................................................................................................... 89 93 96 99 102 106 111 115 Table A15 Relative frequency (and standard error) of herbaceous species that were significantly different (P<0.10) among study areas (Whitefish River Basin-North and -South [WRB-North and -South] and Stonington Peninsula-North and -South [SP-North and -South]) in the Hiawatha National Forest in Michigan's Upper Peninsula, 1993-1994 .......................................................................................... 119 xi Figure 1 Figure 2 Figure 3 Figure 4 LIST OF FIGURES Location of the western zone of the Hiawatha National Forest in Michigan's Upper Peninsula. ..................................................................... 7 Mean monthly temperature at Manistique, Michigan, during the study (1993-94) and the 30-year average (1951-80) .......................................... 9 Total monthly precipitation at Manistique, Michigan, during the study (1993-94) and the 30-year average (1951-80) .......................................... 10 Location of the Whitefish River Basin (WRB) and Stonington Peninsula (SP) deeryards in Michigan's Upper Peninsula ........................ 13 xii INTRODUCTION Forests and white-tailed deer (ngcgjlgns minim) are valuable natural resources in Michigan. Forests cover approximately 7.5 million ha in Michigan and provide recreation, timber products, and habitat for wildlife. Of the 7.5 million ha of forest land, 7.1 million ha has been classified as commercial forest land and is available for the above uses depending on the owner's objectives (Michigan Department of Natural Resources 1983). Forest lands in Michigan are important economically because of their recreation and timber value. Approximately 8.1 million people used state forests in 1976 (Michigan Department of Natural Resources 1977) and Michigan's national forests had 4,916,400 visitor-days in 1990 (U .S.D.A. Forest Service 1990). Michigan's raw timber products were valued at $310.6 million in 1992 (Potter-Witter 1995). Michigan's Forest Resources Plan (Michigan Department of Natural Resources 1983) set targets for forest outputs by 2000 at 138 million user-activity days for wildlife, fish, and other recreational activities and approximately 14 million cubic meters for timber harvests. Forest wildlife, the most well-known probably being the white-tailed deer, has both consumptive and nonconsumptive users throughout the United States. Williamson and Doster (1981) estimated the capitalized value of white-tailed deer in the United States to be approximately $27.3 billion or approximately $1,657 per animal. 2 In Michigan, there were an estimated 1.6 to 1.8 million deer in October, 1992 (Michigan Department of Natural Resources 1992). Approximately 1,250,000 deer hunters spent over $400 million during all 3 deer hunting seasons in 1992 (Michigan Department of Natural Resources 1992). Nationally, the values received by hunters is estimated at $1.8 billion (Williamson and Doster 1981). Langenau (1979) found 3 times more people in Michigan participated in nonhunting activities than the number who hunted deer. The estimated value of benefits received by nonhunters from the national deer herd is substantial-approximately $5.4 billion annually (Williamson and Doster 1981) Forest lands and white-tailed deer numbers have undergone dramatic changes during the past 150 years (Blouch 1984). Mature forests covered the state until the mid- 19th century; correspondingly, white-tailed deer numbers were very low. Extensive logging in the Great Lakes states in the last half of the 19th century created more favorable habitat for deer and their numbers increased. Excessive hunting and repeated wildfires resulted in low deer numbers by the early 19005. Regeneration of large cutover and burned areas began by the 19303 and 19405 and deer numbers once again increased. This cycle of forest-white-tailed deer interaction has provided biologists with valuable information for the management of forest ecosystems. As forests mature, setting back succession is required to maintain optimal habitat conditions for white-tailed deer throughout its range. With the passage of the Multiple Use-Sustained Yield Act of 1960, national forest managers are required to manage for multiple uses of "recreation, range, timber, 3 watershed, and wildlife and fish purposes” (Hunter 1990). With these policy requirements, land management decisions on national forest lands must be oriented toward maintaining wildlife and timber resources to meet multiple-use demands without detrimental effects to either resource. Local concentrations of deer in forested areas may impact forest vegetation by affecting tree regeneration (Dahlberg and Guettinger 1956, Case and McCullough 1987), growth and development (T ilghman 1989), and reduced stocking (Marquis 1974). Providing high quality summer range is important to the over-winter survival of deer because of the role of summer forages in fat accumulation (Mautz 1978). Reducing locally abundant deer numbers may help maintain the general welfare of the deer herd, habitat quality, and the forest ecosystem composition and structure. The recent goal for deer herd size in Michigan is 1.3 million animals (Michigan Department of Natural Resources 1992). Once the goal is reached, management must include accurate harvest quotas to maintain herd size in all areas that will not detrimentally impact forest vegetation and agricultural lands. Currently, deer population numbers and distribution are estimated from deer check station data, highway counts, and field reports (Michigan Department of Natural Resources 1991). However, because of seasonal habitat use and movement patterns, harvest quotas set for regions of Michigan may not reflect the number of deer which should be harvested. Quantification of white-tailed deer spring, summer, and early fall habitat use and movement patterns will provide information to help attain more accurate estimates of herd demographic and habitat requirements in various regions of Michigan's Upper 4 Peninsula. Management across the landscape for vegetation types used by deer can reduce concentrations of deer but still provide a population to meet recreational demands. Managing Michigan's forest lands to meet the demand for timber and wildlife is a complex problem. A study to investigate the deer-forest land relationship in Michigan’s Upper Peninsula was initiated in 1992. A concurrent project with this study quantified deer population dynamics and winter habitat use (Van Deelen 1995). The focus of this project was to quantify forest vegetation types used by white-tailed deer during spring, summer, and fall; determine deer seasonal movement patterns and home ranges; assess the impacts of white-tailed deer on forest vegetation, specifically northern white-cedar (Ihm'a Wis); and provide management recommendations for optimal use of the deer herd and timber resources. OBJECTIVES Specific objectives for this project were to: 1. Determine quantitative estimates of white-tailed deer spring, summer, and early fall habitat use patterns in the central portion of Michigan's Upper Peninsula. 2. Determine deer seasonal movement patterns and home ranges. 4. Evaluate effects of deer browsing on the composition and structure of northern white-cedar stands. 5. Quantify deer browse use of selected tree species. 6. Attain productivity estimates of deer. 7. Provide management recommendations to enhance the ability to manage forest-deer relationships to achieve multiple-use objectives for forest ecosystems. STUDY AREA The study area was centered in the Whitefish River Basin (WRB) and Stonington Peninsula (SP) in the Hiawatha National Forest (HNF) in the central portion of the Upper Peninsula of Michigan (Fig. 1). The HNF lies within Delta, Alger, and Schoolcraft Counties and encompasses approximately 4050 km2. Lake Michigan and Lake Superior border the area to the south and north, respectively. Approximately 90% of the study area is wooded, primarily owned by federal and state governments and several large corporations (Bemdt 1977). Recent (1991) Landsat thematic mapper data (MacLean Consultants Ltd.) estimates that approximately 14% of the land in the SP is comprised of agricultural and herbaceous openland vegetation types compared to approximately 3.5% of the WRB. Major industries in the area are timber, especially pulp production, and recreation. Modern physiography and soils are a result of post-glacial erosion and soil formation processes acting on the glacial deposits (Albert et a1. 1986). Low elevations (207 to 235 m) dominate the flat, glacial lake plains and consist of poorly drained sand and clay soils, exposed limestone and dolomite bedrock, or thin soils over bedrock (Albert et a1. 1986). Soils on the SP are primarily the Nahma-Ensley-Cathro and the Rubicon associations. The majority of the WRB consists of the Tawas-Carbondale- Roscommon, Kiva-Chippeny-Summerville, Rubicon, and Kalkaska associations. The Figure 1. Location of the western zone of the Hiawatha National Forest in Michigan's Upper Peninsula. 8 remaining portion of the study area in the HNF is primarily Dawson-Tawas-Rousseau and the Kalkaska-Tawas-Carbondale soil associations (Berndt 1977). The climate is dominated by lacustrine influences (Albert et al. 1986). Prevailing westerly winds result in a quasi-marine climate near the Great Lakes changing to a semi- continental climate over the inland areas. Spring is delayed because of the cooling of warm southerly air by Lake Michigan. Summers are cool because of lake breezes (Fig. 2) (National Oceanic and Atmospheric Administration 1993-1994). The growing season averages 120 days (Berndt 1977). Winter (November to March) averages 19 days of -17.8 C or below and summer temperatures are rarely (once every 2 years) higher than 32.2 C (Berndt 1977). Precipitation (Fig. 3) (National Oceanic and Atmospheric Administration 1993- 1994) is greatest during the growing season; 60% of annual totals fall from April to September (Berndt 1977). Snow flurries are frequent with snowfall averaging <152.4 cm annually in the southern region to 355.6 cm annually near Lake Superior (Eichenlaub et al. 1990). Vegetation on the SP is both deciduous and evergreen, such as balsam fir (Ahies balm), sugar maple (Age: W), paper birch (Henna W), and hemlock (Isnga W5). In the WRB, vegetation consists of evergreen stands dominated by tamarack (Larix lamina), black spruce (Ricea madam), and white-cedar; broadleaf deciduous forests composed of sugar maple, yellow birch (Henna lntga), beech (Ems grandifolia); along with hardwood-conifer mixes (Kuchler 1964). Vegetation on the well-drained end moraine and ground moraine ridges is dominated by northern 30 20" 10 — . Q \‘ / ' Temperature (Deg C) \ (10) ” I e ’ 19931994 30-Yr.Avg ale -o- (20) r 1 l l i l 4 1 l 1 r J F M A M J J S O N Month Figure 2. Mean monthly temperature at Manistique, Michigan, during the study (1993-94) and the 30-year average (1951-80) (National Oceanic and Atmospheric Administration 1993, 1994). Precipitation (cm) 10 16 I 14 12 1 1 fl \ I 10 5 A a - 0 ’a . ¢ 9 o 2 " 6 1993 1994 30-YrAvg. He ' O l r l r r r l 1 r J F M A M J J A s o N Month Figure 3. Total monthly precipitation at Manistique, Michigan, during the study (1993-94) and the 30-year average (1951-80) (National Oceanic and Atmospheric Administration 1993, 1994). 11 hardwoods. Eastern hemlock, red pine (Emu: resingsa), and white pine (Finns W) are species whose abundance has been altered from cutting and fire (Albert et a1. 1986). Conifer swamps are primarily white-cedar, balsam fir, and white spruce (Eicea glam) (Albert et a1. 1986); red pine and jack pine (Pinus hanksiana) grow on dry sands (Berndt 1977) METHODOLOGY CAPTURING AND RADIO-COLLARIN G White-tailed deer were live-trapped using Stephenson (McBeath 1941) and Clover (Clover 1954) traps from January through mid-April in 1992, 1993, and 1994 in the WRB and SP deeryards (Fig. 4). Trapping was conducted in cooperation with U.P. Whitetails Association's program and field assistants. Traps were baited with shelled corn. Deer were manually restrained, ear-tagged, and radio-collared (Telonics Inc., Mesa, Ariz. and Lotek Engineering Inc., Ontario, Canada). Radio-collars were equipped with mortality switches that doubled the pulse rate if collars remained still for 12 hours. Collars were distributed to each sex in 3 age classes (adults, yearlings, and fawns) in each deeryard. Age of fawns and yearlings was determined through tooth development and wear criteria developed by Severinghaus (1949); age of adult deer was determined by canine tooth extraction and analysis of the cementum annuli (Gilbert 1966, Van Deelen 1995). GENERAL LOCATION TECHNIQUE Seasonal movement patterns and habitat use of deer were determined using a portable TR-2 receiver (Telonics Inc., Mesa, Ariz.) with a hand-held 2-element yagi antenna. Deer were located throughout the winter for another component of the study investigating winter habitat use and population dynamics (Van Deelen 1995). 12 METHODOLOGY CAPTURING AND RADIO-COLLARING White-tailed deer were live-trapped using Stephenson (McBeath 1941) and Clover (Clover 1954) traps from January through mid-April in 1992, 1993, and 1994 in the WRB and SP deeryards (Fig. 4). Trapping was conducted in cooperation with U.P. Whitetails Association's program and field assistants. Traps were baited with shelled corn. Deer were manually restrained, ear-tagged, and radio-collared (Telonics Inc., Mesa, Ariz. and Lotek Engineering Inc., Ontario, Canada). Radio-collars were equipped with mortality switches that doubled the pulse rate if collars remained still for 12 hours. Collars were distributed to each sex in 3 age classes (adults, yearlings, and fawns) in each deeryard. Age of fawns and yearlings was determined through tooth development and wear criteria developed by Severinghaus (1949); age of adult deer was determined by canine tooth extraction and analysis of the cementum annuli (Gilbert 1966, Van Deelen 1995). GENERAL LOCATION TECHNIQUE Seasonal movement patterns and habitat use of deer were determined using a portable TR-2 receiver (Telonics Inc., Mesa, Ariz.) with a hand-held 2-element yagi antenna. Deer were located throughout the winter for another component of the study investigating winter habitat use and population dynamics (Van Deelen 1995). 12 13 Schoolcrafi Delta 93° N Figure 4. Location of Whitefish River Basin (WRB) and Stonington Peninsula (SP) deeryards in Michigan’s Upper Peninsula. 14 Radio-telemetry data for spring/summer and fall were collected from mid-May through September 22 for spring/summer and September 23 through December 20 for fall. Mid- May was the approximate time when all radio-collared deer appeared to be on their spring/summer home ranges as determined by having 2 or more consecutive weekly location points in the same vicinity. For home range analysis, deer were grouped by the area where they established their spring/summer home range; for habitat use analysis, deer were grouped by where they were trapped because their availability area was based on trap location (e. g., a deer trapped on the SP but moved to the WRB for its spring/summer home range was grouped with the WRB deer for home range analysis and with SP deer for habitat use analysis). MOVEMENTS AND HOME RANGES Seasonal movement locations were obtained weekly for all deer using triangulation techniques. If a weekly location point was located in a vegetation type adjacent to the road according to Forest Service compartment maps, the data was recorded for habitat analysis. The order in which deer were located was alternated each week to obtain varied times for individual deer. Triangulation bearing error angle was estimated with 50 sets of 3 bearings obtaining an overall standard deviation using LOCATE II (Pacer 1993). Seasonal home ranges were calculated using the adaptive kernel with 95% contours (Worton 1989) with CALHOME (Kie et a1. 1994) and the minimum convex polygon (Mohr 1947) and the harmonic mean with 95% contours (Dixon and Chapman 15 1980) with TELEM88 (Coleman and Jones 1988). Home ranges were separated by year and season (spring/summer and fall) for analysis. Prior to home range comparisons between study areas, a parametric analysis of variance was performed on the ranked data (Conover and [man 1981) using SAS (SAS Inst, Inc. 1993) to detect possible interactions between and among the 3 factors: sex, age, and area. When interactions between main effects exist, significant differences detected between the main effects would not be meaningful (Sokal and Rohlf 1981); in this project, a comparison of study areas was of primary interest. If interactions were not significant (P>0.10) and if sample size allowed (r210), the Mann-Whitney U test was used to test for significant differences between the study areas. Spearrnan rank correlation coefficients were calculated using SYSTAT (1992) to compare home range sizes estimated by the adaptive kernel, minimum convex polygon, and harmonic mean methods. HABITAT USE Habitat use data were collected during 3 time periods: 0800 to 1559, 1600 to 2359, and 0000 to 0759. The 24-hour sampling technique was used to avoid potential bias involved in sampling just during daylight hours (Beyer and Haufler 1994). Sampling deer was alternated within and among time periods to obtain unbiased data and equal sampling intensities. Vegetation types used by deer were determined by triangulating along edges of vegetation types or walking around deer along the perimeter of stands a 16 minimum of 3 sides to pinpoint locations. Habitat use data points were included with weekly location data for movement analysis. Percent availability of each vegetation type was determined with a circle centered at a central trap site coordinate for the 2 trapping areas (WRB and SP). The circle, with a radius equal to the 85th percentile of the maximum distance moved by a single deer from that trapping area, was overlayed on Landsat thematic mapper vegetation data (Michigan Department of Natural Resources, Wildlife Division, Lansing, Mich.) using the geographic information system ARC/INFO (Environmental Systems Research Institute, Redlands, Calif); the 85th percentile included 95% of the deer from each trapping area. Satellite vegetation classifications (Maclean Consultants Ltd. 1991) were combined into 12 categories for project purposes (Table 1). Satellite areas designated as water were not included in the total land area available; areas designated urban and non- vegetative were grouped into the "other" category. Agricultural-cropland is comprised of row crops only; hay-related crop fields would fall into the herbaceous openland designation. The red, jack, and other (mixed) pine satellite vegetation categories were grouped into the mixed pine category. Tamarack, black spruce, white spruce, balsam fir, and mixed conifer vegetation types were combined into the lowland conifer category. White-cedar was kept as a separate category because the focus of part of this project quantified possible impacts deer have on the composition and structure of cedar stands. Five satellite vegetation types did not have any land area in the 2 study areas and were not included in the project list (Table 1). Table 1. Vegetation type classifications for Michigan's central Upper Peninsula, 1993- 1994. Michigan DNR Deer Habitat Project Deer Habitat Use Thesis Classificationsll Project Classifications Non-Coniferous Non-Coniferous Urban Agricultural-Openland Non-Vegetative Herbaceous Openland Agricultural-Cropland Shrubland Herbaceous Openland Northern Hardwood Shrubland Aspen/Birch Northern Hardwood Dry Hardwood/Conifer Mix Oak” Wet Hardwood/Conifer Mix Aspen/Birch Wetlands Lowland Hardwoodsb Other Dry Hardwood/Conifer Mix Wet Hardwood/Conifer Mix Coniferous Wetlands Mixed Pine Water Lowland Conifers White Cedar Coniferous Pines Red Pine Jack Pine White Pineb Other (Mixed) Pine Tamarack Hemlockb <70% Crown Closure >70% Crown Closure Black Spruce <70% Crown Closure >70% Crown Closure White Spruce <70% Crown Closure >70% Crown Closure Balsam Fir” <70% Crown Closure >70% Crown Closure White Cedar <70% Crown Closure >70% Crown Closure Mixed Conifer <70% Crown Closure >70% Crown Closure 'Data obtained from 1991 Landsat thematic mapper (Maclean Consultants Ltd.). bVegetation type with no land area in the 2 study areas (Whitefish River Basin and Stonington Peninsula). 18 Habitat use analysis combined data from all animals for both years. A chi-square goodness-of-fit test was used to determine if deer used vegetation types in proportion to their availability as described by Neu et al. (1974). In this analysis, the observed value is the number of data points in a vegetation type; the expected value is the proportion of total acreage of that vegetation type times the total number of deer observations. These partial chi-square values for each vegetation type are summed for a total chi-square to be compared to the table value. A confidence interval is then constructed around the proportion observed in each vegetation type to determine which types are being used more than, less than, or as expected. Use is considered to be more than, less than, or as expected if the proportion of the vegetation type available to the deer is lower than, higher than, or within the confidence interval, respectively, built around the proportion of use of that vegetation type. PRODUCTIVITY Productivity estimates were initially determined through direct observation of radio-collared females in 1993. After locating individual deer in specific vegetation types, animals were observed closely to determine if they had fawns. Due to the difficulty in directly observing individual radio-collared deer, 3 standardized driving surveys were conducted at dusk in both study areas during mid-summer in 1994. The number of deer observed in each area was recorded by sex and age. A Mann-Whitney U test was used to compare the fawnzdoe ratio in the SP and WRB using SYSTAT (1992). 19 VEGETATION SAMPLING To assess the impacts of deer browsing on the composition and structure of mature cedar stands, 2 paired areas within selected stands were delineated and 1 was randomly chosen for exclosure construction and the other to be left open to browsing. Twelve mature, well-stocked (70%) stands were selected in sets of 3 on a north-south snow depth gradient (Eichenlaub et a1. 1990) resulting in 4 study area gradients in the WRB and SP (WRB-North and -South and SP-North and -South). Snow depth has been shown to be related to deer use of tamarack swamps (Beier and McCullough 1990), with lowest deer densities believed to be in the WRB-North area and highest numbers in the SP-South area. One stand selected in the SP-South area was not used due to inaccessibility; no replacement stand could be located that met established criteria. Exclosures are 30 m x 30 m x 2.4 m. Three exclosures were built in the WRB- North study area in 1993; l exclosure was constructed in each of the 3 remaining study areas in 1994. Remaining exclosures will be constructed by the US. Forest Service and Michigan State University personnel potentially by 1996. All site locations are listed in Table 2. For the remainder of this document, the sites chosen for exclosure construction will be referred to as exclosure sites even if construction has not been completed. Vegetative sampling for baseline structural and compositional components was conducted on exclosure sites and their respective paired areas open to browsing. A 2-m 20 Table 2. Locations of exclosure and areas open to browsing sites in the Whitefish River Basin-North and -South (WRB-North and -South) and Stonington Peninsula-North and -South (SP-North and -South) study areas in the Hiawatha National Forest in Michigan's Upper Peninsula, 1993-1994. Exclosure Site Study Legal Location Open Area Site Description Compt. Stand (UTMs)‘ Areab WRB- l T43N,R20W 143 27 511025- South North Sec. 19 5106651 2 T43N,R20W 143 36,37 511109- North Sec. 30 5104876 3 T43n,R21 W 143 20,21 510068- Southeast Sec. 24 5105394 WRB- 1 T41N,R19W 64 69 520494- Northeast South Sec. 19 5085756 2 T41N,R21W 94 3 505685- East Sec. 3 5091006 3 T42N,R20W 103 25 512165- South Sec. 20 5095398 SP- 1 T40N,R20W 28 27 517259- West North Sec. 33 5072983 2 T40N,R20W 39 31,32 515563- East Sec. 20 5076573 3 T40N,R20W 39 33,34 515864- Southeast Sec. 20 5076637 SP- 1 T39N,R21W 9 2 509216- Northwest Southc Sec. 27 5065470 2 T39N,R21W 10 22 507210- Northwest Sec. 33 5064156 aUniversal Transverse Mercator. bOpen area direction is in relation to the exclosure site and distances are 25-30 m. c’Ihird site in SP-South was not used due to inaccessibility; no replacement site could be located. 21 buffer was established along the inside perimeter of all sites, so the area disturbed by exclosure construction would not be included in data collection. Vertical cover of vegetation was quantified using the line intercept method (Canfield 1941) and was recorded in 3 height strata: <0.5 m, 0.5 to 2.0 m, and >20 m. Line intercepts were systematically located within exclosure and areas open to browsing sites. Downed woody material cover was also recorded for descriptive purposes. Horizontal cover was determined using a profile board described by Nudds (1 97 7) at randomly selected points in each sample area. The standard observing distance was 4 m determined by recording cover at different distances (3, 4, and 5 m) and choosing the one with the greatest variation (Gysel and Lyon 1980). The height strata for the board were <05 m, 0.5 to 1.0 m, 1.0 to 1.5 m, 1.5 to 2.0 m, and 2.0 to 2.5 m. The stem densities of dominant tree species (northern white-cedar, balsam fir, and sugar maple) were determined by conducting complete counts at each site of each species. Other woody stem densities and frequency of herbaceous species were determined using randomly located nested quadrats 1 x 8 m and l x 4 m, respectively. Densities of woody species were determined using the same 3 height strata used for vertical cover. The height strata used for the above measurements is based on the growth forms of vegetation and structural requirements of deer (Alverson et a1. 1988). Due to exclosure construction only being partially complete, analysis was conducted 2 ways: 22 l. combining exclosure site and open area data within study area gradients for comparisons among gradients. 2. keeping the exclosure site and open area data separate for comparisons within and among study area gradients. Kruskal-Wallis one-way analysis of variance (Siegel and Castellan 1988) was used to compare study area gradients with both combined and separated data using SYSTAT (1992). A Kruskal-Wallis multiple comparison test (Siegel and Castellan 1988) was used to detect where the differences occurred among the study areas. The paired t-test (Steel and Torrie 1980) was used to determine significant differences (P<0.10) for all vegetation characteristics between exclosure and open area sites within a study area. Quality control of all vegetative sampling was assured by determining statistically adequate sample sizes (F reese 1978). An 80% confidence level was used to determine sample sizes. Allowable error was set at 20% of the mean. E . 5 1° Browsing estimates were conducted in the 2 study areas in spring 1994 using 12 randomly established 25-m belt transects in vegetation types adjacent to mature cedar stands used as wintering areas. To allow for sufficient sampling area, stands were selected based on the length of the perimeter of the stand adjacent to the cedar stand. The number of current annual growth stems available of the dominant tree species and the number browsed of the dominant tree species only < 2 m in height were recorded. RESULTS CAPTURING AND RADIO-COLLARIN G One hundred one white-tailed deer were radio-collared in the WRB and SP during the 3 years of trapping (Table 3). Due to mortality occurring prior to spring/summer, location data for this portion of the study were gathered on only 61 of these deer, 22 (36%) males and 39 (64%) females. Of the 61 deer, 17 (28%) were radio-tracked during both seasons and years of the study. MOVEMENTS AND HOME RANGES The median date for movement of wintering deer from the 2 deeryards was March 29 in 1993 and April 4 in 1994 (Van Deelen 1995). Two-thousand four-hundred sixty- five locations were obtained during the 2 years, including 790 (39%) habitat use data points. Maximum migration distances during the 2 years by a single deer from the 2 trapping areas were 54.4 and 52.9 km from the WRB and SP, respectively. In 1993, 9 (28%) of 32 deer with summer ranges in the WRB had been trapped in the SP; in spring/summer 1994, 6 (26%) of 23 WRB deer were SP-trapped deer. Mean telemetry triangulation bearing error angle standard deviation for observers was 8 degrees. 23 24 Table 3. Number of white-tailed deer radio-collared in the Whitefish River Basin (WRB) and the Stonington Peninsula (SP) in the Hiawatha National Forest in Michigan’s Upper Peninsula, 1992-1994. Study Area Year Sex Age WRB SP Total 1992 Male Adult 2 0 2 Yearling 2 1 3 Fawns 5 5 10 Female Adult 7 10 17 Yearling 1 0 1 Fawn 5 5 10 1 993 Male Adult 0 0 0 Yearling 4 0 4 Fawn 5 5 10 Female Adult 8 5 l 3 Yearling 1 0 1 Fawn 6 5 l 1 1994 Male Adult 0 0 0 Yearling 0 0 O Fawn 5 5 10 Female Adult 0 0 0 Yearling 0 0 0 Fawn 4 5 9 Total 55 46 101 25 W Analysis of variance of home range data showed an interaction between area and sex for spring/summer 1993 (P<0.10) for the adaptive kernel (AK) method. Due to the interaction of these 2 factors, the test to determine a significant difference between the 2 study areas for spring/summer 1993 had to be separated by sex first. No interactions were evident for either season in 1994. CALHOME (Kie et a1. 1994) was not able to produce an AK home range with 95% contours for 4 deer; 80% contours worked for 3 of these deer and 50% contours worked for the other deer (Tables A1 and A2, Appendix). In 1994, mean spring/summer and fall home ranges of deer in the WRB were significantly larger (P<0.01) than home ranges of deer in the SP (Tables 4 and 5). WRB female home ranges were significantly larger than SP female home ranges in spring/summer 1993 and 1994 (Table 4). Low sample size did not allow testing of females in fall 1994 or males for all seasons. Male mean spring/summer home ranges ranged from 76.0 ha to 1354.7 ha in the SP and WRB, respectively (Tables A1 and A2, Appendix). Male mean home ranges for SP deer were smaller than WRB deer home ranges during the study, except fall 1993 when a mean for SP male deer could not be determined with only 1 male deer being monitored (Tables 4 and 5). Analysis of variance of home range data showed an interaction between area and sex for spring/summer 1993 (P<0.10) for the harmonic mean (HM); in fall 1993, an interaction between age and area for the minimum convex polygon (MCP) and HM 26 Table 4. Mean spring/summer home ranges (ha) (and standard errors) of white-tailed deer in the Whitefish River Basin (WRB) and Stonington Peninsula (SP) using adaptive kernel with 95% contours in the Hiawatha National Forest in Michigan's Upper Peninsula, 1 993-1 994. vvarl SP ‘uan SP Females 219.6‘ 90.9 817.1' 78.7 (49.4) (22.6) (655.5) (7.8) Males 1354.7 283.3 223.6 76.0 (1147.4) (94.0) (80.7) (17.7) Study Area 645.3 96.7 636.5” 82.8 (431.1) (31.6) (455.7) (8.2) ‘Significantly different from SP (P<0.05) with Mann-Whitney U test using SYSTAT (1992). l’Significantly different from SP (P<0.01) with Mann-Whitney U test using SYSTAT (1992) 27 Table 5. Mean fall home ranges for white-tailed deer in the Whitefish River Basin (WRB) and Stonington Peninsula (SP) using adaptive kernel with 95% contours in the Hiawatha National Forest in Michigan's Upper Peninsula, 1993-1994. 1993 1994 WRB sp WRB SP Female 1252.2 109.2 2147.7'II 39.2 (657.4) (24.0) (1746.0) (12.2) Male 238.4 4...." 297.6 31.5 (99.2) ..-.. (216.8) (8.8) Study Area 914.3 107.7 1563 .4c 37.8 (445.9) (21.9) (1198.3) (10.0) IN too small (<10) to conduct test. I’Only one animal in this category; no mean available. cSignificantly different (P<0.01) from SP with Mann Whitney U test using SYSTAT (1992). 28 methods was detected. Due to the interactions of these factors, tests to determine a significant difference between the 2 main effects of interest (study areas) were not conducted for these seasons and methods. No interactions were evident for either season in 1994. TELEM88 (Coleman and Jones 1988) was not able to produce an HM home range with 95% contours result for 3 deer; 80% contours were used for these deer (Tables A1 and A2, Appendix). Results for the MCP and HM methods were similar to the AK results. Mean spring/summer home ranges for deer in the WRB were significantly larger than for deer on the SP using MCP for 1993 and 1994 and HM in 1994 (Table 6). Mean fall home ranges for deer in the WRB were significantly larger than deer in the SP in 1994 (Table 7) with both methods. In spring/summer 1993 and 1994, WRB female home ranges were significantly larger than SP female home ranges (Table 6) for the MCP and HM home range methods. Low sample size did not allow testing of females in fall 1994 or males for all seasons. Spearman rank correlation coefficients comparing the 3 home range methods ranged from 0.546 to 0.979 for the SP and from 0.852 to 0.989 for the WRB. The low coefficient for the SP was spring/summer 1994, HM versus AK method. Disregarding study area, the coefficient ranged from 0.817 to 0.979 for the 3 sets of correlations (MCP vs. HM, MCP vs. AK, and HM vs. AK). 29 Table 6. Mean spring/ summer home ranges (ha) (and standard errors) of white-tailed deer in the Whitefish River Basin (WRB) and Stonington Peninsula (SP) using minimum convex polygon (MCP) and harmonic mean (HM) with 95% contours in the Hiawatha National Forest in Michigan's Upper Peninsula, 1993-1994. 1993 1994 MCP HM MCP HM WRB SP WRB SP WRB SP WRB SP Females 136.7'I 48.9 343.3. 65.7 185.8. 52.6 490.4h 71.5 (32.0) (10.1) (171.6) (14.5) (76.0) (6.3) (318.3) (10.6) Males 159.2 124.8 693.5 294.0 150.0 56.8 204.9 98.5 (43.5) (42.2) (511.7) (1 15.7) (37.6) (14.9) (49.8) (35.2) Study 145.1' 64.1 474.6 1 l 1.3 174.9c 53.4 366.7° 76.9 Area (25.4) (13.4) (216.9) (33.3) (53.6) (5.6) (194.4) (10.7) ‘Significantly different (P<0.05) from SP with Mann Whitney U test using SYSTAT (1992) l'Significantly different (P<0.10) from SP with Mann Whitney U test using SYSTAT (1992). °Significantly different (P<0.01) from SP with Mann Whitney U test using SYSTAT (1992) 30 Table 7. Mean fall home ranges (ha) (and standard errors) of white-tailed deer in the Whitefish River Basin (WRB) and Stonington Peninsula (SP) using minimum convex polygon (MCP) and harmonic mean with 95% contours (HM) in the Hiawatha National Forest in Michigan's Upper Peninsula, 1993-1994. 1993 1994 MCP HM MCP HM Group WRB SP WRB SP WRB SP WRB SP Females 354.7 62.3 613.4 87.3 371.6 21.7 1727.9 27.9 (230.6) (1 1.4) (314.6) (19.0) (162.7) (7.5) (1 140.3) (9.8) Males 98.8 122.8 ---- -...-- 137.1 22.8 207.5 34.2 (38.0) (54.3) (----) (----) (96.5) (6.8) (133.8) (10.3) Study 269.4 60.3 449.9 89.1 297.6' 21.9 1247.7" 29.1 Area (154.6) (10.6) (213.7) (17.5) (116.4) (6.2) (789.0) (8.0) ‘Significantly different (P<0.05) from SP with Mann Whitney U test using SYSTAT (1992). 1’Significantly different (P<0.01) from SP with Mann Whitney U test using SYSTAT (I 992). 31 HABITAT USE Habitat availability was based on the maximum migration distance by a single deer from the 2 trapping areas. The circle radii, centered at a central trap site coordinate, for the WRB and SP were 46.2 and 45.0 km, respectively. The dominant vegetation types for the 2 areas were northern hardwoods for the WRB and wet hardwood/conifer mix for the SP area, respectively, averaging 24.7% of the land. Northern white-cedar was the least available (<1%) in both study areas. Habitat use in the 2 areas was not in proportion to availability for spring/summer (Table 8). Aspen/birch and mixed pine were used significantly more than expected in the WRB; aspen/birch, mixed pine, and northern white-cedar were used significantly more than expected by SP deer (Table 8). Vegetation types used less than expected were agricultural-croplands and other by both WRB and SP deer and northern hardwoods and wetlands by WRB and SP deer, respectively. All other vegetation types were used as expected in the 2 study areas (Table 8). Agricultural-croplands and “other” were not used by deer in either area, but an observed value of 0.0001 was used in the analysis so a result of being used less, more, or as expected could be determined. Fall habitat use for the 2 areas was similar to spring/summer use. Approximately 87% of the habitat use data points occurred in the same 5 vegetation types in the WRB and SP (northern hardwoods, aspen/birch, wet hardwood/conifer, mixed pine, and lowland conifer) (Table 9). The highest use was lowland conifers--21.77% and 26.85% 32 Table 8. White-tailed deer spring/summer habitat use and availability in the Whitefish River Basin (WRB) and Stonington Peninsula (SP) study areas in the Hiawatha National Forest in Michigan's Upper Peninsula, 1993-1994. WRB SP n'=27 n=33 mb=246 tn=277 Vegetation Type %Avail %Use %Avail %Use Agricmm'cmpland 9.29 0.00004c 12.86 0.00004“ Herbaceous Openland 4.74 4.88 5.95 10.47 Shrubland 1.90 4.47 2.04 5.05 Northern Hardwoods 27.31 20.33‘I 17.75 18.41 Aspen/Birch 4.57 11.79“ 2.92 13.36“ Dry Hardwood/ 5.11 9.35 5.98 3.61 Conifer Mix Wet Hardwood/ 21.46 15.85 23.38 19.49 Conifer Mix Wetlands 2.13 0.81 2.25 0.00004“ Mixed Pine 5.96 14.23“ 5.55 11.19“ ‘ White Cedar 0.20 1.63 0.25 3.25“ :1] . . 1 1‘ ) Lowland Conifers 15.96 16.67 19.30 15.16 Other 1.38 0.00004c 2.10 0.00004“ Chi-square value 134.83" 295.32" ‘Total number deer monitored. l’Total number of locations. cUsed significantly less than (P<0.01) percent available with Bonferroni normal statistic (Neu et a1. 1974). (Used significantly less than (P<0.10) percent available with Bonferroni normal statistic (Neu et a1. 1974). °Used significantly more than (P<0.10) percent available with Bonferroni normal statistic (Neu et a1. 1974). "Significantly different (P<0.0001) from availability by chi-square analysis (Neu et al. 1974) 33 Table 9. White-tailed deer fall habitat use in the Whitefish River Basin (WRB) and Stonington Peninsula (SP) study areas in the Hiawatha National Forest in Michigan's Upper Peninsula, 1993-1994. % Use WRB SP n'=27 n=23 Vegetation Type rn"=124 m=149 Agricultural-Cropland 0.00 0.00 Herbaceous Openland 6.50 2.68 Shrubland 0.81 1.34 Northern Hardwoods 14.50 15.44 Aspen/Birch 16.94 10.74 Dry Hardwood/Conifer 6.45 2.01 Mix Wet Hardwood/Conifer 12.90 19.46 Mix Wetland 0.00 0.00 Mixed Pine 18.55 16.10 White Cedar 1.61 5.37 (Ihuia occidentalis) Lowland Conifers 21.77 26.85 Other 0.00 0.00 'Total number of deer monitored. bTotal number of locations. 34 for the WRB and SP, respectively. Agricultural-croplands, “other,” and wetlands were not used in either study area. Because of the low sample size for fall, no analyses were conducted. A breakdown of the habitat use data points by the 3 time periods used to record data had similar results as the total percentages. The 2 vegetation types with the highest percentage of use during the spring/summer (disregarding time periods) were northern hardwoods (20.33%) and lowland conifers (16.67%) by WRB deer and wet hardwood conifer mix (19.49%) and northern hardwoods (18.41%) by SP deer. Northern hardwoods, lowland conifers, and wet hardwood/conifer mix had the highest percentage of use by time period (Table 10). Table 10. Vegetation types with highest percentage of use by white-tailed deer by time period in the Whitefish River Basin and Stonington Peninsula study areas in Michigan’s Upper Peninsula, 1993-94. Study Area Time Period Whitefish River Basin Stonington Peninsula 0800-1559 Northern Hardwoods Wet Hardwood/Conifer Mix 1600-23 59 Lowland Conifers Wet Hardwood/Conifer Mix 0000-0759 Lowland Conifers Wet Hardwood/Conifer Mix, Northern Hardwoods, and Lowland Conifersal 1'All 3 vegetation types with same level of use. 35 PRODUCTIVITY During the first year of the project, the attempts to estimate productivity through direct observation of radio-collared females resulted in only 8 of 19 deer being observed. The difficulty in observing radio-collared females at close range led to the use of road driving surveys in the 2 study areas to estimate productivity. The highest fawn:doe ratio for the WRB and SP was 0.44 and 0.42, respectively; the lowest fawn:doe ratio for the WRB and SP was 0.0 and 0.07, respectively (Table 11). The mean fawn:doe ratio was not significantly different (P>0.10) between the WRB and SP in mid-summer 1994 (Table 11). VEGETATION SAMPLING C .. 15 [ll] 30.4215 1 The mature cedar stands selected for the exclosure study differed among study area gradients in ways that were evident by direct observation. The 3 paired sites in the WRB-North were wetter (i.e., more standing water) than the sites in the other 3 study areas; the understory was also much more dense in the WRB-North sites. The majority of the surrounding vegetation in the WRB-North was northern hardwoods, primarily maple. These factors may contribute to some of the differences found in the vegetation data among the study areas. The 2 sets of analyses (exclosure and open area site data combined and exclosure and open area site data separated) are reported; all tables for the exclosure and open area data separated are contained in the Appendix. 36 Table 11. Mean productivity and stande error (SE) of white-tailed deer in the Whitefish River Basin (WRB) and Stonington Peninsula (SP) study areas in the Hiawatha National Forest in Michigan's Upper Peninsula, 1994. Number Observed Study Area Survey Replicate Does Fawns Fawns:Doe SP 1 12 5 0.42 2 8 2 0.25 3 14 1 0.07 Mean 11.3 2.7 0.251‘ (S.E.) (1.8) (1.2) (0.10) WRB 1 15 0 0.00 2 9 4 0.44 3 13 2 0.15 Mean 12.3 2.0 0.20 (SE) (1 .8) (1.2) (0.13) 'No significant difference (P>0.10) between study areas with Mann Whitney U test using SYSTAT (1992). 37 Exclosure and Open Area Site Data Combined Vertical cover was significantly different (P<0.10) among the study areas in all 3 strata (Table 12). The largest difference was seen in the 0.5-2.0 m stratum where the mean percent vertical cover in the WRB-North was substantially higher than found in the other 3 study areas (Table 12). Downed woody material cover ranged from 2.1% to 5.5%. Mean percent horizontal cover was significantly greater (P<0.10) in the WRB-North than the other 3 study area gradients for the upper 3 strata (1-1 .5 111, 1.5-2.0 m, and 2.0-2.5 m) (Table 13). For the <05 and 0.5-1.0 m strata, mean percent horizontal cover in the WRB-North was significantly greater (P<0. 10) than cover in the SP-North and -South and SP-South, respectively (Table 13). Stem densities of 3 dominant tree species (northern white-cedar, balsam fir, and sugar maple) were significantly different (P<0.10) among the study areas in the 4 strata except for balsam fir and sugar maple in the >2.0 m, >12.67 cm dbh stratum (Table 14). Northern white-cedar stem densities in the <05 m stratum ranged from 987 to 12,475 stems/ha but from 0 to 202 stems/ha in the 0.5-2.0 m stratum. Forty-four non-dominant woody species were identified in the 4 study area gradients (Table 15). Densities were substantially different depending on the study area gradient and stratum for a few of the species. For instance, black ash (Earrings nigra) had 29,069 stems/ha in SP-North in the <05 m stratum compared to 0 stems/ha in this stratum in the WRB-North. Stern densities were significantly different (P<0.10) among 38 Table 12. Mean percent vertical cover (and standard error) for height strata in the 4 study areas (Whitefish River Basin-North and -South [WRB-North and -South] and Stonington Peninsula-North and -South [SP-North and -South]) in the Hiawatha National Forest in Michigan's Upper Peninsula, 1993-1994. Study Area Stratum WRB- WRB- SP- SP- (m) North South North South <0.5 89.5A' 78.9AB 73.53 72.43 (1.8) (4.2) (2.2) (5.3) 0.5-2.0 69.1A 5.23 7.33 1.58 (6.1) (1.1) (3.8) (0.5) >2.0 82.4A 92.8B 88.3AB 92.8B (1 .8) (1.5) (2.4) (1.1) DWMb 5.5 2.1 4.3 3.4 (1.1) (0.4) (1.2) (0.4) IIMeans with different letters within a stratum were significantly different (P<0.10) among study areas with the Kruskal-Wallis one-way analysis of variance (Siegel and Castellan 1988) using SYSTAT (1992) and the Kruskal-Wallis multiple comparison test (Siegel and Castellan 1988). bDowned woody material; descriptive only, no tests conducted. 39 Table 13. Mean percent horizontal cover (and standard error) for height strata in the 4 study areas (Whitefish River Basin-North and -South [WRB-North and -South] and Stonington Peninsula-North and -South [SP-North and -South]) in the Hiawatha National Forest in Michigan's Upper Peninsula, 1993-1994. Study Areas Stratum WRB- WRB- SP- SP- (m) North South North South <0.5 68.0A' 29.2AB 21.03 18.43 (6.7) (3.0) (2.7) (2.7) 0.5-1.0 48.6A 17.4AB 17.9AB 9.93 (6.7) (4.4) (4.4) (1.3) 1.0-1.5 50.4A 15.03 12.53 9.43 (4.5) (3.2) (2.7) (2.6) 1.5-2.0 37.6A 14.73 13.03 8.53 (9.7) (3.2) (3.1) (2.6) 2.0-2.5 44.0A 14.83 17.73 6.03 (4.1) (3.7) (4.6) (3.0) llMeans with different letters within a stratum were significantly different (P<0.10) among study areas by the Kruskal-Wallis one-way analysis of variance (Siegel and Castellan 1988) using SYSTAT (1992) and the Kruskal-Wallis multiple comparison test (Siegel and Castellan 1988). £8: $3.5 use. see 3:986 was Fwommv 3554; we £ng 33.28 £33-?me 05 3 383% mm 803 tween 05 muons 838%. can 860% a £55 3580 BEBE—o Ewe—mama 05 20:3 82% 3 0555 83 A32 §=880 Ea Emommv 68 :SEQES oat—=8 333433.. .932 5:280 one .omoav as 88858 29:2: 3:63-345. o5 es... 38: .255 was: $2 5:890 one enema 85:? eo auras atone £33 rid—mav— ofi .23 32a beam mecca 8..cv& Engage bucmoficwmm 203 838% 93 86on a £53» £83— 88er 5m? «:32. 8V iaxm=br .63. 5:23 Ba 32% as Engages 22.32 £33-.“me 2: 23 a8: :55 was 52 5:230 as 329 8.3:? .3 max—£8 33-25 £33-3me 05 .23 30.8 35% $555 2: .ovmv EEobE 3.585%? 203 833% can 33on a £53 3080— “53$“. .23 mane—2. 2: S. § 2.0 m strata) and open area sites (0.5-2.0 m stratum) (Table A3, Appendix). Downed woody material cover at the sites ranged from 2.0 to 6.3%. No significant differences in horizontal cover were detected between exclosures and open areas for all 4 study areas (Table A4, Appendix). Significant differences (P<0.10) in horizontal cover comparisons were detected among the study area gradients 49 Table 16. Herbaceous species summary for the 4 study areas (Whitefish River Basin- North and -South [WRB-North and -South] and Stonington Peninsula-North and -South [SP-North and -South]) in the Hiawatha National Forest in Michigan's Upper Peninsula, 1993-1994. Study Area Species Richness No. Species With 3 Most Common Single Occurrence Species in Study Area WRB-North WRB-South SP-North SP-South 49 59 56 52 Bunchberry 7 Camus mediums: Canada mayflower Maianthemum mamas: Goldthread 9.912113 mmandica Goldthread 14 Naked miterwort M11911: nude Starflower I . 1° 1 1° Canada mayflower 3 Violet m spp. Starflower Canada mayflower 13 Naked miterwort Starflower 50 Table 17. Mean relative frequencies (and standard errors) of herbaceous species in the 4 study areas (Whitefish River Basin-North and -South [WRB-North and -South] and the Stonington Peninsula-North and -South [SP-North and -South]) in the Hiawatha National Forest in Michigan's Upper Peninsula, 1993-1994. Study Areas Species WRB-North WRB-South SP-North SP-South Anemone spp. OAB' OAB 0.3A OB (0) (0) (02) (0) Arrow arum 0A 0A 0.2A 0A Eelflanshxitainisa (0) (0) (0-2) (0) Am spp. 1.8A 1.3A 2.1A 0A (0.6) (0.5) (0.6) (0) Bedstraw 0A 0.5A 0.2A 0.2A fialinm Spp. (0) (0-5) (02) (0-2) Blunt-lobed woodsia 0A 0A 0.1A 0A mm (0) (0) (0-1) (0) Boots wood fern 0A GA 2.48 0.8AB Dnmtrris 1mm (0) (0) (0-7) (0.6) Bracken fern 0A 0.3A 0.3A 0A Mdiumamminum (0) (0.2) (0-3) (0) Bugleweed 0A 0.5AB 0.8AB 2.7B Lmaus SW (0) (0.2) (0.4) (1.0) Bulbet fern 0A 0A 0. 1A 0A WW (0) (0) (0-1) (0) Bunchberry 5.6A 3.6AB 1.6B 3.1AB Camus madsnsis (0-5) (1 -3) (0-8) (1-0) Buttercup 0A 0A 2.4A 0A Banunszulns SPP- (0) (0) (1 -7) (0) Canada mayflower 4.6A 5.7AB 7.03 5.8AB Maianthsmnmsanadsnse (0-3) (0.4) (0-9) (0-6) Cinnamon fern 0.1A 0.4A 0A 0A Qsmunda cinnamoma (0.1) (0.3) (0) (0) Cinquefoil 0A 0A 0.1A 0A Retentilla SW (0) (0) (0- 1) (0) Clover o" o 0.7 o Irifalium SW (0) (0) (0-3) (0) Club moss 0A 0.6A 0.2A 0A Winn: SW (0) (0-4) (0.2) (0) Club-spur orchid 0A 0.1A 0A 0A Hahmada daxellata (0) (0.1) (0) (0) Table 17(cont'd). Study Areas Species WRB-North WRB-South SP—North SP-South Columbine 0A 0.1A 0A 0A Amnesia SPP- (0) (0- 1) (0) (0) Common wood sorrel 0.1A 0A 0A 0A Qxalismnmana (0-1) (0) (0) (0) Coralroot o" o o 0.5 991311931113 5139- (0) (0) (0) (0-3) Crested wood fem 0.4A 0.6A 0.1A 1.1A Wanted: mm (0-3) (0-4) (0-1) (0-6) Dandelion 0" o 0.7 o Immm 5w (0) (0) (0.3) (0) Dewberry 0A 4.2A 2.2A 5.3A Ruhus 111512151115 (0) (0-6) (1 -1) (0-9) mm spp. 4.4” 0.1 1.2 0.5 (1.3) (0.1) (0.6) (0.5) Dwarf enchanter‘s nightshade 0.1AB 0A l.6AB 3.8B Qirsasulnina (0-1) (0) (1-0) (0.7) mm spp. 0.3A 0.1A 3.1A 0.6A (0.2) (0.1) (1.1) (0.6) False Solomon's seal 0A 0A 0.1A 0A Smilasinarassmasa (0) (0) (0.1) (0) Fragile fern 1.7A 0.1AB OB 0.9AB WWII: (0-5) (0-1) (0) (0.6) Fringed brome 0.1A 0A 0A 0A Bmmnsciliams (0.1) (0) (0) (0) Fringed polygala OB 5.5A OB 0.2AB WW (0) (0-6) (0) (02) Golden ragwort 0A 0.5A 0.1A 0.2A Wm (0) (0-4) (0-1) (0-2) Goldenrod 3.1A 2.2A 3.2A 0.7A Sgfidagg spp. (1.0) (0.7) (0.7) (0.4) Goldthread 5.4A 6.7A OB 4.0AB Coulis malandisa (0-8) (0-5) (0) (0-6) Grass spp. 2.4A 4.0A 4.1A 4.3A (1.1) (1.2) (1.0) (1.8) Grass/sedge spp. 0A 0A 4.9B 1.4AB (0) (0) (1.2) (0-9) 52 Table I7 (cont'd). Study Areas Species WRB-North WRB-South SP-North SP-South Hawkweed 0.6AB 013 4.0A 013 Hiemiumspp. (0.3) (0) (1.9) (0) Hooked crowfoot 0A 0A 0.2A 0A Bannncuhlsmmm (0) (0) (0.2) (0) Intermediate wood fern 0.2" o 0.7 2.9 mm intennedia (0.2) (0) (02) (1.7) Interrupted fern 0A 0.1A 0A 0.3A annnda damniana (0) (0.1) (0) (0.3) Jack-in-the-pulpit 0" 0.5 0 2.5 Ansaemasvp- (0) (0.5) (0) (1.3) Jewelweed 0.3" 0.7 0 0.9 linemen: SW (0.3) (0.7) (0) (0.4) Joe-pye weed 1.3" 0.7 o 0.2 W spp. (0.5) (0.3) (0) (0.2) Large-leaved aster 0.6A 0.4A 0A 0A Aster MW (03) (0.4) (0) (0) Lesser pyrola 0A 0.5A 0A 0A mm]: minor (0) (0.5) (0) (0) Long beech fern 0" 0.1 0 2.0 MW (0) (0.1) (0) (0-8) Manna grass 0.6A 0A 0A 0A 01m 5013- (0-6) (0) (0) (0) Marsh bedstraw 1.8b 0.1 0.3 0 Galinm 0111115112: (06) (0.1) (0.2) (0) Marsh fern 0A 0.3A 0A 0.9A mums aheanatsris (0) (0.3) (0) (0.5) Marsh marigold 0.5A 0.2A 0A 0A (2111111: mm: (0.5) (0.2) (0) (0) Marsh Skullcap 0A 0A 0A 0.2A WW (0) (0) (0) (02) Milkweed 0A 0A 0.1A 0A Ameniasspp. (0) (0) (0.1) (0) Mint 0" o 0.2 0.4 Menrhaspp- (0) (0) (0.1) (03) Moss spp. 6.7A 7.1A 8.1A 7.3A (0.3) (0.2) (1 .0) (0.4) Table 17 (cont'd). 53 Study Areas Sim“ WRB—North WRB-South SP-North SP-South Naked miterwort SAAB 6.2AB 3.3A 7.5B Mltella nuda (1 -0) (0-5) (1 -0) (03) Oak fern 12A 0.3A 2.4A 2.4A Gmnmamiuln 5m (05) (0-3) (1 -0) (12) One-flowered pyrola 0A 0.1A 0A 0A Maltese: nniflnra (0) (0- 1) (0) (0) mm: spp. 0.4A 0.1A 0A 0A (02) (0.1) (0) (0) Ostrich fern 0A 0A 0. 1A 0A Maneusiammmm (0) (0) (0-1) (0) 2mm spp. 0.7A 1.5A 1.1A 0.2A (0.3) (0.5) (0.6) (0.2) Rattlesnake fern 1.3A 1.8A 3.3A 2.6A Wm (04) (0-8) (0-6) (1-0) Rattlesnake plantain 0.1h 0 0 0.7 W SPP. (0- 1) (0) (0) (0-4) Rough bedstraw 0A 3.2B 2.8AB 5.3B fiallumasnmllmn (0) (0.5) (0-8) (05) Royal fern 0.1A l.lA 0A l.lA ernunda malls (0.1) (0.5) (0) (1-1) Sedge 5.9AB 5.8A 3.8AB l.6B Cami spp. (0.1) (0.9) (1.2) (0.7) Self-heal 0.5" 0.4 1.6 0.6 2111112113 inland: (0-3) (0-3) (0-2) (06) Sensitive fern 0A 0.4A 0.1A 0.9A Shingles scnsilzills (0) (0-2) (0- 1) (0-9) Showy lady's slipper 0.3A 0A 0A 0A mm (0-2) (0) (0) (0) Silvery glade fern 0b 0 1.0 0.7A Athxflumthslmtsmides (0) (0) (0-6) (0-7) Skullcap 0A 0A 0.2A 0A Santellaria SW (0) (0) (0-2) (0) Small—flowered cranberry 0.4A 0.6A 0A 0A Macsininmnxxcm (0.2) (0.6) (0) (0) 54 Table 17 (cont'd). Study Areas 59“” WRB-North WRB-South SP-North SP-South Snowberry 3.3AB 5.0A OB OB Whlsnidula (0-9) (0-9) (0) (0) Spinulose wood fern 0A 0A 0.5A 0A WW (0) (0) (0-4) (0) Spurred gentian 0A 0A 1.3A 0A Haleniadsflsxa (0) (0) (1-0) (0) Starflower 3.1A 6.2A 4.5A 6.0A WW1: (1.0) (0-5) (1.1) (1-3) Strawberry 4.9A 1.8AB 12B 0.8AB Emma SPP- (0-5) (1.1) (0-5) (0.3) Sundew 0. IA 0A 0A 0A Dream 5119- (0.1) (0) (0) (0) Sweet coltsfoot 0.9" 0.1 3.2 0 2211151195 mlmanls (0.6) (0- 1) (1-5) (0) Sweet-scented bedstraw 2.4” 0.1 0 0 Cralium Eiflnmm (0-9) (0.1) (0) (0) Tall buttercup 0A 0A 0.5A 0A Rannnsulus am: (0) (0) (0-5) (0) Tall meadow rue 0A 0. 1A 0A 0A malicmlm mlxsamnm (0) (0.1) (0) (0) Thistle 3 .4A 1 .9AB 2.8A 0.2B Cirsium Spp. (0-7) (0-5) (0-7) (0-2) Three-leaved Solomon's seal 4.6A 2.1AB OB OB Smilacinairifclia (0-8) (0-6) (0) (0) Trailing arbutus 0.1A 0.5A 0A 0A Wm (01) (0.3) (0) (0) In'lliurnspp. 0" 0 0 0.2A (0) (0) (0) (0.2) Turtlehead 0A 0A 0A 0.2A Chem: SPP- (0) (0) (0) (0-2) Twinflower 5.5A 3.5AB 0.4B 0.2B Linnasa mm (0-4) (1 .3) (0-2) (0.2) Twisted stalk 0A 0.4A 0.2A 0.3A W amnlsxifczlius (0) (0.4) (0-2) (0-3) 55 Table 17 (cont'd). Study Areas Species WRBoNorth WRB-South SP-North SP—South Violet 5.2A 3.4A 6.2A 5.7A M11213 5w (03) (0-8) (0-5) (07) White adders mouth 0A 0A 0A 0.2A Malaxishmbmda (0) (0) (0) (02) Wild ginger OA 0.1A 0A 0A Ammadsnse (0) (0-1) (0) (0) Wild sarsaparillo 3.3A 2.1A 3.8A 3.5A Aralia nudisaulis (0-6) (1 -1) (1 .0) (0-9) Wintergreen 0.9" 1.1 O 0 Gaulthma when: (03) (0-5) (0) (0) Wood anemone 0.5AB GA 2.28 0.6AB Anemone ill-11115111219113 (0-4) (0) (0.7) (0-6) Wood sorrel 0.6A 0.9A 0A 3.9A Qxalis 599- (0-6) (0-7) (0) (23) Yellow lady's slipper 0A 0.3A 0A 0.7A WW (0) (0.2) (0) (0-4) ‘Relative frequencies with different letters within a species were significantly different (P<0.10) among study areas with the Kruskal-Wallis one-way analysis of variance (Siegel and Castellan 1988) using SYSTAT (1992) and the Kruskal-Wallis multiple comparison test (Siegel and Castellan 1988). eruskal-Wallis multiple comparison test (Siegel and Castellan 1988) was unable to detect where the significant difference occurred within a species among the study areas as detected by the Kruskal-Wallis one-way analysis of variance (Siegel and Castellan 1988) using SYSTAT (1992). 56 for exclosure sites (0.5-1.0 and 2.0-2.5 m strata) and open area sites (1.5-2.0 and 2.0-2.5 m strata) (Table A4, Appendix); however, 4 of the 8 multiple comparisons were not able to detect where the difference occurred using the Kruskal-Wallis multiple comparison test. Stem densities of balsam fir in WRB-North was the only dominant species that showed a significant difference between exclosures and paired open areas in the 0-0.5 m stratum (Table A5, Appendix). Northern white-cedar in the WRB-North study area had significantly more stems in both >2.0 m in height strata than the WRB-South study area (Table A5, Appendix). Eight other Kruskal-Wallis multiple comparisons were not able to detect where the significant difference occurred as indicated by the Kruskal-Wallis one- way analysis of variance. Densities of other woody species showed no significant differences between exclosure and open area sites for the WRB-North and SP-South study area (Tables A6 and A9, Appendix). Densities of ironwood (Carpinns camhniana) and balsam poplar in exclosure sites (0-0.5 m stratum) were significantly greater (P<0.10) than open area sites for WRB-South and SP-North, respectively; density of alder-leaved buckthorn (Rhamnns alnifqlia) in the open area was significantly greater than in the exclosure for SP-North (Tables A7 and A8, Appendix). Densities of 8 woody species in exclosures and 5 species in open areas were significantly different from other study areas (Table A10, Appendix). However, the Kruskal-Wallis multiple comparison test was only able to detect where the difference occurred in 7 of 13 of these cases (Table A10, Appendix). 57 Only a few herbaceous species were significantly more frequent in exclosure or open area sites (Tables A1 l-A14, Appendix). Thirteen species in exclosures and 11 species in open areas showed significantly different relative frequencies (Table A15, Appendix) among the study area gradients with the Kruskal-Wallis one-way analysis of variance. The Kruskal-Wallis multiple comparison test was only able to detect where the difference occurred for 4 species in exclosures and 3 species in open areas (Table 15, Appendix). W The 3 tree species sampled for browse use were black ash (Eraxim nigza), balsam poplar (12me habemifera), and maple (Aeer sp.); the 3 species were the dominant species in stands adjacent to the cedar stands and were common to both study areas (WRB and SP). The mean percent browse use of sugar maple was substantially less than black ash and balsam poplar in both the WRB and SP: 8.7 and 1.2% for sugar maple, 42.3 and 42.5% for black ash, and 45.3 and 31.3% for balsam poplar, respectively. DISCUSSION MOVEMENTS AND HOME RANGES The maximum distance moved from wintering areas to summer range by a single deer was 54.4 and 52.9 km from the WRB and SP, respectively. These results are comparable to 51.5 km found by Venue (1973) in Michigan's Upper Peninsula, but were greater than maximum distances traveled by deer in Wisconsin (Dahlberg and Guettinger 1956, Larson et a1. 1978). Although several deer moved north-northeast off the SP for their summer range both years of the study, the majority (70%) of SP-trapped deer remained on the SP for their summer range and are considered a resident population. Interactions were detected for all home range methods in 1993 in either spring/summer or fall. The interactions involved area and sex (HM and AK in spring/summer) or age (MCP and HM for fall). The main objective was to compare home ranges between the 2 study areas, but with sex or age interacting with area, a comparison of the main effects was not conducted for the above cases. The smallest adaptive kernel (AK) home range during the study was 6.3 ha in fall 1993 (Tables A1 and A2, Appendix) and may be attributed to having few weekly locations (4-5) recorded during the season due to not being able to locate the individual deer because of its movement away from what appeared to be its home range. AK home ranges for 4 WRB deer (2 adult females, 1 yearling female, and 1 yearling male) were 5,000+ ha (5,274.0 to 22,8300 ha), which was 5 to 21 times higher than the study area 58 59 mean for that particular season and year (Tables A1 and A2, Appendix). These large home ranges were due to mid-season travel by the deer, sometimes returning to their original summer or fall ranges, others remaining at their new location. Reasons for the travel may have been attributed to dispersal or some type of disturbance. In South Dakota, Sparrowe and Springer (1970) had an adult doe move 12.8 km from her summer range in September and returned in October. This type of behavior is either uncommon or is not reported in the literature. In addition to the AK home range method, the MCP was used for comparisons to previous deer studies. The mean spring/summer home range (MCP estimator) for WRB deer (160 ha) for the 2 years was comparable to deer home ranges documented in Minnesota (162 ha) (Nelson and Mech 1984) and Wisconsin (178 ha) (Larson et al. 1978) in contrast to the much smaller mean spring/summer ranges of SP deer (58.8 ha). Larger home range sizes for WRB deer than SP deer may be attributed to the smaller percentage of land in herbaceous openings in the WRB (3.5%) compared to the SP (14%), which provides forage for deer during the spring/summer period. Use of this vegetation type was 10.47% in the SP compared to only 4.88% in the WRB. With a higher percentage of land in herbaceous openings and potentially other early successional stages, deer may not have needed to travel as far to meet their nutritional requirements. Female summer home range size was similar to the home range size of the entire sample. This is probably a result of the high percentage of females in the radio—collared sample and averaged 161.3 ha and 50.8 ha for the WRB and SP, respectively. These data 6O compare to Minnesota studies that found female deer summer home ranges of 282.7 ha (Kohn and Mooty 1971) and 120 ha (Nelson and Mech 1984). In the WRB, mean fall home ranges for deer were larger than spring/summer home ranges for both years. These results are in contrast to data reported by Mooty et a1. (1987) in Minnesota where the trend was for ranges during summer intervals to be larger than ranges during November-December and other winter intervals. Supplemental feeding may also affect home range size. Although this topic was not addressed in this project, personal communication with residents on the SP indicates supplemental feeding may be an extensive practice. The degree of feeding during each season is unknown. This practice may impact home range size and habitat use more in the SP than the WRB, because the SP appears to have a higher density of permanent human residents than the WRB. The effects of supplemental feeding and the higher percentage of herbaceous openland in the SP versus the WRB would need to be separated to determine their respective impacts on home range sizes and habitat use. 11 B E . C . In seven of the 8 time periods for the 2 study areas (spring/summer-WRB and SP for 2 years; fall-WRB and SP for 2 years), AK mean home ranges were the largest followed by HM and MCP. Analyzing deer individually, though, only 63% followed this pattern; 27% resulted with HM>AK>MCP. For comparisons to previous studies, MCP was one of the home range methods used. A major disadvantage to this method, though, is that the size of the home range increases as the number of location points increases 61 (Jennrich and Turner 1969); the number of location points per deer in this study ranged from 4 to 30. The HM and AK methods describe the intensity of use at a specific point. Additionally, the AK method provides a probability density function and a means of smoothing data to make more efficient use of the data (W orton 1989). No assumption is made about the form of the utilization distribution, and therefore, 30 to 100 observations per animal should be collected to obtain an accurate estimate (Worton 1987). Being able to detect use intensity provides more detailed information about habitat requirements of the animal being observed. For example, Naef-Daenzer (1993) showed density estimates from a modified kernel estimation for a male blue tit were highly correlated with caterpillar density on individual trees and the distance of the trees from the nest. HABITAT USE Habitat availability was based on a circle centered around a central trap location in the 2 trapping areas. Most deer movements from wintering areas to summer range appeared to be in a north-northeast direction during this study. Although this pattern existed, a circle instead of a wedge was used to estimate habitat availability because deer movement patterns may change (e. g., northwest-southwest) direction if habitat quality would change. The 2 vegetation types with the highest percentages of use for the 2 study areas were northern hardwoods (20.33%) and lowland conifers (16.67%) in the WRB and wet hardwood/conifer mix (19.49%) and northern hardwoods (18.41%) in the SP. Although 62 these types had high use, they also cover a large amount of land area and, except for northern hardwoods in the WRB, had use approximately equal to availability. The overall chi-square goodness-of-fit tests were significantly different (P<0.0001) for each study area for both years showing that deer were not using vegetation types in proportion to availability (Table 8). Vegetation types used more than expected during spring/summer were aspen/birch and mixed pine in the WRB and aspen/birch, mixed pine, and white-cedar in the SP. In Wisconsin, McCaffery and Creed (1969) found significantly more deer track crossing in aspen than in northern hardwood types. Kohn and Mooty (1971) also found deer preferred birch and aspen-birch-conifer cover types in Minnesota. The aspen/birch vegetation type provides forage during the summer period and has been shown to dominate the deer diet through early and late summer (McCaffery et a1. 1974). Forbs have also been found to be an important part of deer summer diet (McCullough 1985) and may be provided in the northern hardwood vegetation type. Mixed pine types being used more than expected may be due to an understory and herbaceous component encroaching from adjacent hardwood stands. Vegetation types used less than expected included northern hardwoods, agricultural-croplands, and the other category in the WRB and wetlands, agricultural- croplands, and the other category in the SP. In a study in Minnesota, deer were found to avoid red pine, aspen-conifer, and birch-conifer (Kohn and Mooty 1971 ); trees in the red pine stands were usually 3 to 6 m apart, approximately 15 m tall and 35 to 45-years-old. 63 The authors did find distance between trees and height influenced their use by deer, so avoidance or selection differences between studies may be due to stand characteristics. The agricultural-cropland vegetation type was not used in the WRB or SP in either year of the study, although availability ranged fiom 9.3 to 14.6% for the 2 study areas. Although the study area was not designated as one of the problem deer damage areas of the state, many farmers report deer damage in neighboring Menominee County and other regions of Michigan (Michigan State University 1989). No use by deer of the agricultural-cropland vegetation type during this study may appear unusual but could be due to a couple of factors. The designation of agricultural-cropland in the vegetation database used for this study is specifically for row crops and does not include open, grassy areas such as hayfields; these areas would be classified as herbaceous openland. Alfalfa and mixed hay fields occupied approximately 1.5% and 0.8% of the land in Delta and Alger Counties, respectively (Farm Service Agency pers. comm). Approximately 3,109 ha of corn, dry beans, and potatoes are grown in Delta County; none of these row crops are grown in Alger County (Michigan Dept. of Agriculture 1993). Secondly, a higher percentage of areas classified as agricultural-cropland occur in areas of the availability circles radio- collared deer did not use, e.g., the west side of Little Bay de Noc where approximately 20.2% of the land is agricultural-cropland compared to approximately 9.5% in the SP. Cedar vegetation type availability was low for both study areas (<1%). Spring/summer use of this vegetation type was 1.63% and 3.25% for the WRB and SP, respectively, which was as expected for the WRB and more than expected for the SP. 64 The low use may be due to other foods being available in other vegetation types (e.g., northern hardwoods, herbaceous openland) during the spring/summer. PRODUCTIVITY There was not a significant difference in the fawn:doe ratio between the WRB and SP. The total number of does and fawns observed during the 3 driving surveys was 34 and 8, respectively, for the SP and 37 and 6, respectively, for the WRB (T able 11). Initially, 3 driving surveys were conducted June 15th to 20th with only 5 fawns being observed between both areas. Although experienced does give birth in late May or early June (Ozoga ct al. 1994), the majority of fawns were probably still spending a high percentage of their time bedding during this first set of surveys. Testing the fawn:doe ratio did not show a significant difference between the WRB and SP. Although there is more open area on the SP and an expected greater opportunity to see deer, docs are secretive for the first 4 to 6 weeks after giving birth (Ozoga et a1. 1994), possibly equating the observation level for both study areas and resulting in a non- significant difference. Dense understories also reduces opportunity for viewing deer; this has also been shown to be a problem of night spotlighting (McCullough 1982). A low sample size may also have contributed to the non-significant difference. According to sample size requirement calculations (F reese 1978), 409 and 1066 surveys would be required in the SP and WRB, respectively, to obtain an accurate estimate of the fawn:doe ratio. 65 VEGETATION SAMPLING C .. IS [111 INT-[215 1 Long-term studies of deer browsing on various forest types and on timber products has not been well documented. A 20-year study (1942 to 1962) on the Allegheny National Forest in Pennsylvania reported declines in the understory of a virgin hemlock-hardwood forest (Hough 1965). A more recent study in Pennsylvania reported changes in species composition, growth impacts, and dramatic stocking differences between fenced and unfenced areas 9 to 22 years after clearcutting (Marquis 1981). The exclosure study initiated in this project will provide the opportunity to document long- term impacts of deer on the structure and composition of the cedar type. Although data recorded for baseline purposes were extensive, monitoring may focus on specific aspects of the community, e.g., cedar and rare herbaceous plant species. The herbaceous component of forest communities is an important part of a deer's diet, but few studies have investigated the impacts of deer foraging on herbaceous species (Miller et al. 1992). Herbaceous plants comprised 87% of the summer diet of white- tailed deer in Wisconsin (McCaffery et a1. 1974). This grazing impact is especially important regarding rare plants. Of the 95 species identified in the 4 study areas, several species of the Qrehidaeeae family were observed including showy and yellow lady's slipper which are favored by deer (Alverson et al. 198 8). The intensity of herbaceous plant use in the cedar stands was not documented during this project but could be incorporated into the long-tenn monitoring to be conducted at these sites. 66 Exclosure and Open Area Site Data Combined Significance tests among the study area gradients in which the exclosure and open area site data were combined revealed significant differences for all the vegetation components measured. Some of the differences were much more evident even ocularly (e.g., WRB-North vertical cover 0.5-2.0 m stratum compared to the other 3 study areas). With the densities of nomdominant woody species and the frequency of herbaceous species, many multiple comparison tests detected a difference among study areas, but the multiple comparison could not detect where the difference occurred. This may be due to the small sample sizes involved. Trends were evident when comparing the study area gradients. Mean percent vertical cover dropped dramatically in the 0.5-2.0 m stratum from WRB-North to other study areas; this may be due to the higher amount of water found in the WRB-North sites. Stem densities of northern white-cedar in the 0-0.5 m stratum were much higher in the WRB than the SP. This may be due to site variation among the study areas or lower deer densities. The greater stem densities in the 00.5 m stratum compared to the 0.5-2.0 m stratum in the WRB may indicate the problem with cedar is being recruited into higher height classes and not regeneration. Once cedar grows above snow levels, browsing pressure may impact growth. Stem densities of other woody species ofien appeared to be related to the vegetation types surrounding the cedar stands, e.g., black ash had 14,444 stems/ha in SP-North (where a black ash stand was adjacent to the cedar stand) versus no stems in WRB-North where the cedar stands were surrounded primarily by northern hardwoods. 67 A wide variety of herbaceous species (95) were identified in the 4 study areas, but species richness was similar for the areas ranging from 49 to 59. Twenty species were common to the 4 areas with several of these species also being the most common within the study area gradients, e.g., bunchberry (Cm-nus emdensis), Canada mayflower, naked miterwort, and starflower, indicating their ability to exist in variable habitats. Thirty- three species, though, were identified in only 1 study area gradient indicating more specific habitat requirements. Exclosure and Open Area Site Data Separated Paired t-tests comparing composition and structure data of exclosure to open area sites showed minimal significant differences (Tables A3-A14, Appendix), due to the fact that, even on sites where exclosures have been constructed, enough time has not elapsed for vegetation differences to be detected. Differences that were detected were probably due to natural variation and microhabitat differences between exclosure and open area sites. The comparisons among study areas resulted in 47 occurrences of significant differences primarily in the density of other woody species and frequency of herbaceous species. Multiple comparison tests for each of the species showing a significant difference were not always able to detect where the difference occurred probably due to the small sample size from each study area. Wanna Deer browsing affects forest vegetation in different ways. Regeneration may decrease as reported in several studies (Dahlberg and Guettinger 1956, Beals et a1. 1960, 68 Behrend et al. 1970, Anderson and Loucks 1979, Frelich and Lorimer 1985, Case and McCullough 1987). Vegetation may be able to regenerate and be recruited, but their growth may be detrimentally impacted by deer browsing (T ilghman 1989). Some species, though, are able to repair injuries received from deer browsing and recover to grow beyond the reach of ungulates (Graham 1958). In contrast, mountain maple (Aeer spieamm) has been found to slightly increase as browse pressure increases (Beals et a1. 1960) To compare the relative difference in browsing of selected tree species in the WRB and SP, browsing estimates of 3 tree species in stands adjacent to cedar stands were obtained in the 2 study areas. Sugar maple browsing (<10%) appeared to be much lower than that for balsam poplar and black ash (ranging from 31.3% to 45.3%) in each study area. The lower browsing levels for sugar maple may be due to a couple of reasons. First, the lower average height of sugar maple stems at the SP sampling site making the stems unavailable due to snow pack. Secondly, the extensive land area of northern hardwoods available where the sugar maple was sampled at the WRB sampling site (and in the WRB in general [27.31%]), which may spread the browsing intensity over a larger area reducing the percent browsed in the sampling area. CONCLUSIONS Overall use of vegetation types in the 2 areas (WRB and SP) was not in proportion to availability in either year. During spring/summer, vegetation types used more than expected were aspen/birch and mixed pine in the WRB and aspen/birch, mixed pine, and white-cedar in the SP. Deer use of the agricultural- cropland and “other” vegetation type categories was not observed in either study areas. Fall habitat use was similar to spring/summer use. Deer primarily used 5 vegetation types during the fall: northern hardwoods, aspen/birch, wet hardwood/conifer mix, mixed pine, and lowland conifers. Deer movement from wintering areas to spring/summer home ranges reached maximum distances of 54.4 and 52.9 km from the WRB and SP trapping areas, respectively. During the 2 years of the study, 27% of WRB summer range deer were SP-trapped deer. Spring/summer home ranges were significantly larger in the WRB than the SP for both years. Fall home ranges were significantly larger in the WRB than the SP in 1994. Female spring/summer home ranges were significantly larger in the WRB than the SP in 1993 and both seasons in 1994. Few significant differences were detected in vegetation characteristics between exclosure sites and open areas in all 4 study area gradients. Quantifying the vegetation characteristics over time, however, may provide natural resource 69 _.r. b 7O managers with a greater understanding of the long-term impacts of deer on the northern white-cedar type. Sugar maple had a lower browsing intensity than black ash and balsam poplar in the WRB and SP. Productivity (fawnszdoe) was not significantly different between the WRB and the SP. sho $19 abur type 10021 CODE 13311 “fine redLlc SUpp] MANAGEMENT RECOMMENDATIONS Long-term forest management practices in the Hiawatha National Forest (HNF) should focus on maintaining vegetation types throughout the landscape that meet wildlife, timber, and other multiple use objectives. Maintaining vegetation types for deer populations can provide habitat for other species, both game and non-game, and be a part of the overall management scheme for the HNF. Deer movements can be extensive from wintering areas to spring/summer ranges, so selected vegetation types should be provided to minimize the effects of locally abundant numbers on vegetation communities. A review of the distribution of vegetation types across the landscape may indicate the possibility of high concentrations of deer in local areas. Locations of future timber harvests should be planned to help reduce deer concentrations and alleviate browsing pressure on some forest species. Spring supply of vegetation types that provide foods used by deer, such as graminids and evergreen ground plants (McCaffery et a1. 1974) in close proximity to wintering areas will provide deer with the necessary food component to recover from reduced food intake during the winter months (Ozoga and Verrne 1970). Spring food supply is also important for pregnant does that are nearing the fawning period. Deer population goals for the state are set at 1.3 million; population size as of October 1994 was approximately 1.6 million (Ozoga et a1. 1994). Seasonally high concentrations of deer may impact the forest community through changes in species 71 72 composition (Marquis 1981) and impaired regeneration (Behrend et al. 1970, Anderson and Loucks 1979, Case and McCullough 1987). In areas where plant community composition and structure is a concern and deer are locally abundant, further herd reduction by removing more antlerless deer should be considered. The spatial relationships of seasonal habitat use patterns and movements of deer need to be considered when setting harvest regulations and delineating Deer Management Unit boundaries. A challenge in setting harvest regulations is determining where wintering deer that are impacting vegetation are located during hunting season (e. g., what Deer Management Unit). Education of the hunting public needs to be conducted about management objectives and how antlerless deer hunting helps maintain forest ecosystem composition and structure and why deer seasonal demographics need to be considered when setting harvest quotas. APPENDIX 73 Table A1. Seasonal home ranges (ha) of white-tailed deer in the Whitefish River Basin (WRB) and Stonington Peninsula (SP) using minimum convex polygon (MCP), harmonic mean (HM) with 95% contours, and adaptive kernel (AK) with 95% contours in the Hiawatha National Forest in Michigan's Upper Peninsula, 1993. Spring/Summer Home Ranges (ha) Study Area Sex Age Deer # MCP HM AK WRB Female Adult 1091 4.5 61.9 78.1 750 9.5 90.6 145.1 691 4.5 98.8 128.4 1340 30.0 393.7 415.1 591 16.0 326.5 273.6 1011 20.0 160.5 174.4 990 9.5 39.6 74.5 1290 48.5 771.4 624.2 1320 3.5 63.5 87.0 411 3.0 66.7 77.7 850 40.5 1462 1 18.4‘ 480 06.0 236.9 186.9 Female Adult Mean 24.6 204.7 198.6 (S.E.) 33.5) (60.9) (48.1) Yearling 540 217.0 319.7 362.1 390 151.0 209.4 249.3 550 77.5 78.7 129.9 499 127.5 175.9 166.1 420 8.0 6.7 34.9 400 54.0 70.8 99.0 370 576.0 3517.0 928.4 1300 27.0 31.3 38.7 Female Yearling Mean 154.8 551.2 251.1 (8.15.) (64.9) (425.3) (104.2) Female Mean 136.7 343.3 219.6 (S.E.) (32.0) (171.6) (49.4) Male Adult 791 193.5 266.8 234.2 710 247.5 287.3 382.8 1230 99.0 257.1 172.8 060 133.5 163.4 242.4 079 l 1.0 39.1 50.3 Male Adult Mean 136.9 202.7 216.5 ($13.) (40.5) (46.1) (53.9) Yearling 490 85.5 136.4 145.6 . 270 559.0 6313.7 13970.0 360 37.5 63.3 77.3 520 90.5 175.3 197.7 439 265.5 354.4 466.2 460 22.0 51.5 60.4 1370 165.5 222.5 257.0 Male Yearling Mean 175.1 1044.0 2167.7 (SE) (71.2) (879.1) (1967.7) Male Mean 159.2 693.5 1354.7 (S.E.) (43.5) (511.7) (1147.4) WRB Study Area Mean 145.1 474.6 645.3 (S.E.) (25.4) (216.9) (431.1) Table A1 (cont'd). 74 Spring/Summer Home Ranges (ha) Study Area Sex Age Deer # MCP HM AK SP Female Adult 731 19.5 28.6 39.4 811 55.0 78.3 97.5 1270 44.5 76.2 94.9 771 104.5 106.8 178.4 1221 45.0 59.6 73.1 601 37.5 49.6 74.0 831 37.5 31.6 43.9 1110 133.5 205.7 305.3 Female Adult Mean 59.6 79.6 1 13.3 (8.8.) (13.7) (20.2) (31.4) Yearling 510 32.5 48.8 51.7 570 28.0 37.3 46.3 350 23.0 31.4 48.7 1280 26.0 34.4 38.0 Female Yearling Mean 27.4 38.0 46.2 (8.13.) (2.0) (3.8) (2.9) Female Mean 48.9 65.7 90.9 (S.E.) (10.1) (14.5) (22.6) Male Adult 1310 138.0 337.7 354.4 Male Adult Mean ____ ...... __ (S.E.) Yearling 530 46.0 75.3 97.1 1330 190.5 468.9 398.5 Male Yearling Mean 118.3 272.1 247.8 (S.E.) (72.3) (196.8) (150.7) Male Mean 124.8 294.0 283.3 (8.15.) (42.2) (115.7) (94.0) SP Study Area Mean 64.1 111.3 96.7 (5.8.) (13.4) (33.3) (31.6) 75 Table A1 (cont'd). Fall Home Ranges (ha) Study Area Sex Age Deer # MCP HM AK WRB Female Adult 1091 308.0 559.3 816.7 750 36.5 51.8 60.5 691 50.0 54.0 56.4 591 3742.0 1971.3 9411.0 1011 27.5 43.6 54.4 990 735.5 1842.5 3404.0 1290 19.5 25.2 59.1 1320 24.5 32.9 58.4 411 44.0 48.9 82.6 850 290.0 4702.0 5274.0 480 166.0 196.6 313.6 Female Adult Mean 494.9 866.2 1781.0 (5.8) (331.2) (442.1) (923.8) Yearling 390 92.0 104.0 157.3 550 46.0 61.0 100.0 499 2.5 7.2 6.3 400 66.5 81.6 119.4 370 25.0 32.0 61.4 Female Yearling Mean 46.4 57.2 88.9 (3.8) (15.6) (17.2) (25.8) Female Mean 354.7 613.4 1252.2 (8.15.) (230.6) (314.6) (657.4) Male Adult 791 343.5 472.3 909.2 710 34.0 56.3 163.9 1230 121.0 179.0 230.2 060 48.0 31.5 141.7 Male Adult Mean 136.6 184.8 361.3 (S.E.) (71.5) (101.1) (183.6) Yearling 270 99.5 148.6 139.8 360 31.0 25.5 84.7 520 4.0 8.8 8.5‘ 460 109.0 60.7 228.9 Male Yearling Mean 60.9 60.9 1 15.5 (5.15.) (25.7) (31.2) (46.4) Male Mean 98.8 122.8 238.4 (5.8.) (38.0) (54.3) (99.2) WRB Study Area Mean 269.4 449.9 914.3 (8.8.) (154.6) (213.7) (445.9) 76 Table A1 (cont'd). Fall Home Ranges (ha) Study Area Sex Age Deer # MCP HM AK SP Female Adult 81 1 70.0 70.9 98.2 1270 46.0 90.7 81.6 771 34.0 54.0 59.2 1221 39.5 50.8 86.6 601 35.5 36.8 67.3 831 24.5 29.9 28.3 1110 60.5 72.8 103.3 Female Adult Mean 44.3 58.0 74.9 (SE) (6.0) (8.1) (9.8) Yearling 510 150.0 257.1 322.8 570 72.5 131.5 98.1 350 111.0 74.0b 175.8 1280 42.0 91.8 79.5 Female Yearling Mean 93.9 138.6 169.1 (8.8.) (23.4) (41.3) (55.3) Female Mean 62.3 87.3 109.2 (S.E.) (11.4) (19.0) (24.0) Male Adult 1310 38.5 109.1 91.6 Male Adult Mean ...... ...... ...... (8.8) Male Mean --- --- --- (S.E.) SP Study Area Mean 60.3 89.1 107.7 (8.8.) (10.6) (17.5) (21.9) ‘80°/e contour; CALHOME program (Kie et a1. 1994) was not able to calculate 95% contour. t’80% contour; TELEM88 program (Coleman and Jones 1988) was not able to calculate 95% contour. 77 Table A2. Seasonal home ranges (ha) for white-tailed deer in the Whitefish River Basin (WRB) and Stonington Peninsula (SP) using minimum convex polygon (MCP), harmonic mean (HM) with 95% contours, and adaptive kernel (AK) with 95% contours in the Hiawatha National Forest in Michigan’s Upper Peninsula, 1994. Spring/Summer Home Ranges (ha) Study Area Sex Age Deer # MCP HM AK WRB Female Adult 1091 187.0 300.3 309.4 691 168.0 207.1 232.3 591 239.0 797.5 480.1 1011 161.0 195.6 191.9 390 51.0 94.4 77.8 990 161.5 177.9 181.1 499 152.5 1 10.0‘ 208.1 550 74.5 85.1 103.3 1320 56.5 51.5 69.8 400 38.5 61.0 57.9 411 110.0 24.7‘ 155.5 370 45.5 63.4 61.8 480 58.0 79.3 83.6 Female Adult Mean 115.6 192.1 170.2 (5.8) (18.3) (65.1) (33.6) Yearling 530 1302.0 4573.0 10640.0 1310 40.5 53.3 61.0 360 126.5 126.7 159.9 Female Yearling Mean 489.7 1584.3 3620.3 (8.15.) (406.9) (1494.5) (3510.0) Female Mean 185.8 490.4 817.1 (S.E.) (76.0) (318.3) (655.5) Male Adult 270 90.0 137.9 104.5 1230 150.5 185.5 169.9 460 328.5 483.4 693.0 Male Adult 189.7 268.9 322.5 Mean (71.6) (108.1) (186.2) (5.8) Yearling 640 40.5 60.6 62.8 520 235.0 179.3 226.0 079 115.5 185.6 169.0 710 90.0 201.8 140.0 Male Yearling Mean 120.3 156.8 149.5 (8.8) (41.3) (32.4) (34.0) Male Mean 150.0 204.9 223.6 (S.E.) (37.6) (49.8) (80.7) WRB Study Area Mean 174.9 366.7 636.5 (5.13.) (53.6) (194.4) (455.7) ‘ 78 Table A2 (cont'd). Spring/Summer Home Ranges (ha) Study Area Sex Age Deer # MCP HM AK SP Female Adult 1221 53.5 84.2 81.0 771 58.0 73.2 98.6 601 61.0 76.4 68.9 831 96.5 105.0 116.4 1 1 10 27.0 10.4 87.2 570 28.5 45.1 49.0 350 65.5 111.8 94.7 1280 31.0 17.9 44.1” Female Adult Mean 52.6 65.5 80.0 (5.8) (8.3) (13.3) (8.8) Yearling 1330 75.0 129.3 111.0 490 59.0 73.1 98.4 730 51.0 91.5 612 790 25.0 40.4 33.4 Female Yearling Mean 52.5 83.6 76.0 (8.15.) (10.4) (18.5) (17.7) Female Mean 52.6 71.5 78.7 (SE) (6.3) ( 10.6) (7.8) Male Yearling 620 27.5 51.8 51.3 1170 67.0 76.1 150.6 1370 76.0 167.5 96.1 Male Yearling Mean 56.8 98.5 76.0 (SE) (14.9) (35.2) (17.7) Male Mean 56.8 98.5 76.0 (8.1-I.) (14.9) (35.2) (17.7) SP Study Area Mean 53.4 76.9 82.8 (8.8.) (5.6) (10.7) (8.2) 79 Table A2 (cont'd). Fall Home Ranges (ha) Study Area Sex Age Deer it MCP HM AK WRB Female Adult 691 474.0 1527.1 517.0 591 1595.0 14952.0 22830.0 1011 28.5 64.5 32.9 990 917.0 1772.9 3736.0 499 38.0 204.2 50.2 550 22.5 32.9 52.7 1320 40.0 55.6 89.4 400 79.0 82.6 95.5 41 1 27.5 46.8 48.8 480 36.0 45.4 61.0 Female Adult Mean 325.8 1878.4 2751.4 (5.8) (168.7) (1467.5) (22602) Yearling 530 50.0 120.1 126.7 1310 1516.0 3544.0 259.7c 360 7.5 14.0 19.8 Female Yearling Mean 524.5 1226.0 135.4 (8.8) (495.9) (1 159.4) (69.4) Female Mean 371.6 1727.9 2147.7 (5.8.) (162.7) (1140.3) (1746.0) Male Adult 270 20.5 13.8 42.3 1230 69.0 149.0 164.8 Male Adult Mean 44.8 81.4 103.6 (S.E.) (24.3) (67.6) (61.3) Yearling 640 105.5 197.3 162.7 520 7.5 12.5 23.4 079 7.0 15.9 19.2 710 613.0 856.4 1373.0 Male Yearling Mean 183.3 270.5 394.6 (88.) (145.1) (200.0) (327.8) Male Mean 137.1 207.5 297.6 (5.8) (96.5) (133.8) (216.8) WRB Study Area Mean 297.6 1247.7 1563.4 (5.5) (1 16.4) (789.0) (1198.3) 8O Table A2 (cont'd). Fall Home Ranges (ha) Study Area Sex Age Deer # MCP HM AK SP Female Adult 1221 42.5 55.8 77.0 771 6.0 9.2 11.1 601 10.5 10.2 21.9 831 45.6 46.5 82.1 350 64.0 89.0 100.5 Female Adult Mean 33.7 42.1 58.5 (8.15.) (11.0) (15.0) (17.7) Yearling 1330 9.0 17.3 27.0 490 5.5 8.1 8.7 730 4.5 6.7 11.9 790 7.5 8.6 12.6 Female Yearling Mean 6.6 10.2 15.1 (8.8) (1.0) (2.4) (4.1) Female Mean 21.7 27.9 39.2 (8.8) (7.5) (9.8) (12.2) Male Yearling 620 16.0 23.9 22.7 1370 29.5 44.5 40.3 Male Yearling Mean 22.8 34.2 31.5 (SE) (6.8) (10.3) (8.8) Male Mean 22.8 34.2 31.5 (8.8) (6.8) (10.3) (8.8) SP Study Area Mean 21.9 29.1 37.8 (SE) (6.2) (8.0) (10.0) ‘80% contour; TELEM88 program (Coleman and Jones 1988) would not calculate 95% contour. t’80% contour; CALHOME program (Kie et a1. 1994) would not calculate 95% contour. c50% contour; CALHOME program (Kie et a1. 1994) would not calculate 95% contour. \ 81 Table A3. Mean percent vertical cover (and standard error) for height strata in exclosure and areas open to browsing sites in the Hiawatha National Forest in Michigan's Upper Peninsula, 1993-1994. Site Study Area Stratum (m) Exclosure Open Area Whitefish River <0.5 90.7A' (1.2) 8.4A (3.6) 3381111101111 0.5-2.0 60.3A (5.6) 77.9A (8.7) >2.0 80.7A (2.9) 84.2A (2.1) DWMb 5.3 (1.4) 5.8 (1.9) Whitefish River <0.5 74.4A (8.0) 83.3A (1.8) Basin-South 0.5-2.0 5.2AB(2.1) 5.1AB(1.3) >2.0 94.08 (0.3) 91.6A (3.2) DWM 2.2 (0.7) 2.0 (0.3) Stonington <0.5 73.2A (4.4) 73.7A (2.3) Peninsula-North 0.5-2.0 4.3AB(1.8) 10.2AB(7.7) >2.0 90.5AB(1.7) 86.0A (4.5) DWM 2.3 (0.7) 6.3 (1.7) Stonington <0.5 66.6A“ (7.2) 78.2A (6.9) Peninsula-South 0.5-2.0 078* (0.1) 2.48 (0.4) >2.0 91.7AB(2.2) 93.8A (0.6) DWM 4.0 (0.3) 2.8 (0.5) aMeans with different letters within a site and stratum were significantly different (P<0.10) among study areas with the Kruskal-Wallis one-way analysis of variance (Siegel and Castellan 1988) using SYSTAT (1992) and Kruskal-Wallis multiple comparison test (Siegel and Castellan 1988). I”Downed woody material; descriptive only, no tests conducted. *Significantly different (P<0.10) from open area with paired t-test (Steel and Torrie 1980). 82 Table A4. Mean percent horizontal cover (and standard error) for height strata in exclosure and areas open to browsing sites in the Hiawatha National Forest in Michigan's Upper Peninsula, 1993-1994. Site Study Area Stratum (m) Exclosure" Open Area Whitefish River Basin- <0.5 753' (5.2) 60.6A (12.0) North 0.5-1.0 58.3A" (7.1) 38.9A (8.9) 1.0-1.5 54.9' (5.5) 46.0'l (7.1) 1.5-2.0 39.5'I (6.1) 35.7A (6.1) 2.0-2.5 44.2A (6.3) 43.8A (6.7) Whitefish River Basin- <0.5 29.2 (3.4) 29.2A (5.8) South 0.5-1.0 13.8AB(5.9) 21.0A (6.9) 1.0-1.5 11.5 (4.2) 18.6 (4.7) 1.5-2.0 8.8 (2.2) 20.6AB (3.4) 2.0-2.5 7.98 (1.3) 21.6AB(4.3) Stonington Peninsula— <0.5 24.0 (4.2) 18.1A (3.2) North 0.5-1.0 14.0AB(2.7) 21.8A (8.6) 1.0-1.5 14.5 (3.9) 10.4 (4.1) 1.5-2.0 16.7 (4.6) 9.38 (3.8) 2.0-2.5 22.4AB(7.5) 13.0AB(5.2) Stonington Peninsula- 0.10) between exclosure and open area sites for all study areas and strata with paired t-test (Steel and Torrie 1980). llKruskal-Wallis multiple comparison test (Siegel and Castellan 1988) was unable to detect where the significant difference occurred within a site and stratum among the study areas as detected by the Kruskal-Wallis one-way analysis of variance (Siegel and Castellan 1988) using SYSTAT (1992). bMeans with different letters within a site and stratum were significantly different (P<0.10) among study areas by the Kruskal-Wallis one-way analysis of variance (Siegel and Castellan 1988) using SYSTAT (1992) and Kruskal-Wallis multiple comparison test (Siegel and Castellan 1988). 83 Table A5. Mean stem densities per hectare (and standard error) of dominant tree species in exclosure and areas open to browsing sites in the 4 study areas (Whitefish River Basin-North and -South [WRB-North and -South] and the Stonington Peninsula-North and -South [SP-North and -South]) in the Hiawatha National Forest in Michigan's Upper Peninsula, 1993-1994. Site Study Area Species Stratum (m) Exclosure Open Area WRB- Northern white cedar 0-0.5 12179A' (360) 10789A (1875) North Ihttjanszciticntaiis 0.5-2.0 148" (64) 256b (227) >2.0,<12.67 cm dbh 20A (10) 10" (5) >2.0,>12.67 cm dbh 523A (120) 478A (61) Balsam fir 0-0.5 9472A" (2112) 7855A (1918) mm 0.5-2.0 10695b (1319) 7761” (2718) >2.0,<12.67cmdbh 1622A (240) 2056” (334) >2.0,>12.67 cm dbh 104A (89) 39A (26) Sugarmaple 0-0.5 296A (281) 1183A (571) Amsasshamm 0.5-2.0 1681 (153) 84" (13) >2.0,<12.67 cm dbh 10A (5) 10A (5) >2.0,>12.67 cm dbh 10A (10) 0A (0) WRB- Northern white cedar 0-0.5 12288A (6783) 12663A (7042) South 1111114038210th 0.5-2.0 0 (0) 0 (0) >2.0,<12.67 cm dbh 1736B (1098) 1977 (765) >2.0,>12.67 cm dbh 12038 (242) 1016A (317) Balsam hr 005 19305A 16090A (8730) Ahiesbalsamea 0.5-2.0 (13632) 138 (110) >2.0,<12.67 cm dbh 251 (223) 237 (101) >2.0,>12.67 cm dbh 148A (97) 54A (39) 20A (10) Sugarmaple 005 212A (212) 163A (155) mm 05-20 0 (0) 0 (0) >2.0,<12.67 cm dbh 0A (0) 0A (0) >2.0,>12.67 cm dbh 0A (0) 0A (0) SP-North Northern white cedar 0-0.5 878A (431) 2362A (1265) Ihlijaossitlentalis 0.5-2.0 o (0) 79 (79) >2.0,<12.67 cm dbh 325AB (260) 705 (580) >2.0,>12.67 cm dbh 720AB (35) 695A (78) Balsam fir 0-0.5 1893A (1154) 3136A (1616) Ahieshalsamea 0.5-2.0 20 (13) 168 (168) >2.0,<12.67 cm dbh 207A (171) 227 (219) >2.0,>12.67 cm dbh 84A (36) 35A (21) Sugar maple 0-0.5 25A (13) 49A (42) Ammhanim 05-20 0 (0) 0 (0) >2.0,<12.67 cm dbh 0A (0) 0A (0) >2.0,>12.67 cm dbh 0A (0) 0A (0) 84 Table A5 (cont'd). Site Study Area Species Stratum (m) Exclosure Open Area SP-South Northern white cedar 0-0.5 1065A (459) 910A (200) Ihlu'a mldmtalis 0.5-2.0 0 (0) 0 (0) >2.0, <12.67 cm dbh 207AB (74) 207 (74) >2.0, >12.67 cm dbh 888AB (133) 799A (0) Balsam fir 0-0.5 1938A (1938) 377A (126) Allies balm 0.5-2.0 7 (7) 15 (15) >2.0, <12.67 cm dbh 74A (74) 155 (141) >2.0, >12.67 cm dbh 52A (52) 44A (15) Sugar maple 0-0.5 0A (0) 0A (0) Ass: sacchanlm 05-20 0 (0) 0 (0) >20, < 12.67 cm dbh 0A (0) 0A (0) >20, >12.67 cm dbh 0A (0) 0A (0) 'Means with different letters within a site, species, and stratum were significantly different (P<0.10) among study areas with the Kruskal-Wallis one-way analysis of variance (Siegel and Castellan 1988) using SYSTAT (1992) and Kruskal-Wallis multiple comparison test (Siegel and Castellan 1988). eruskal-Wallis multiple comparison test (Siegel and Castellan 1988) was unable to detect where the significant difference occurred within a site, species, and stratum among study areas as detected by the Kruskal-Wallis one-way analysis of variance (Siegel and Castellan 1988) using SYSTAT (1992). ‘Significantly difi‘erent (P<0.10) from open area with paired t-test (Steel and Torrie 1980). 85 Table A6. Mean stem densities per hectare (and standard error) of non-dominant woody species in exclosure and areas open to browsing sites in the Whitefish River Basin-North study area in the Hiawatha National Forest in Michigan's Upper Peninsula, 1993-1994. Site Species Stratum (m) Exclosure" Open Area Alder-leaved buckthorn 0-0.5 1972 (1188) 2750 (2014) Shamans alniiolia 0.5-2.0 1194 (989) 1917 (1792) >2.0, <12.67 cm dbh 0(0) 0(0) >2.0, >12.67 cm dbh 0(0) 0 (0) Altemate-leaved 0-0.5 417 (417) 611 (611) dogwood 0.5-2.0 0 (0) 0 (0) Camus allemifolia >2.0, <12.67 cm dbh 0 (0) 0 (0) >20, >12.67 cm dbh 0 (0) 0(0) American elm 0-0.5 83 (48) 28 (28) Lllnuls mild 0.5-2.0 28 (28) 28 (28) >20, <12.67 cm dbh 0 (0) 0(0) >2.0, >12.67 cm dbh 0(0) 0(0) American mountain ash 0-0.5 56 (28) 83 (48) 59112115 amenisana 0.5-2.0 0 (0) 0 (0) >20, < 12.67 cm dbh 0 (0) 0 (0) >20, >12.67 cm dbh 0 (0) 0(0) American red raspberry 0-0.S 0 (0) 28 (28) 81112115 idaells 05-2-0 0 (0) 0 (0) >20, <12.67 cm dbh 0 (0) 0(0) >2.0, >12.67 cm dbh 0 (0) 0(0) Balsam poplar 0-0.5 0(0) 0 (0) 1192111115 halaamifera 05-20 0 (0) 0 (0) >20, <12.67 cm dbh 28 (28) 111 (111) >2.0, >12.67 cm dbh 0 (0) 0 (0) Beaked hazelnut 0-0.5 111(111) 111(111) Conan: gamma 05-2-0 0 (0) 0 (0) >20, <12.67 cm dbh 0(0) 0(0) >2.0, >12.67 cm dbh 0 (0) 0 (0) Black ash 0-0.5 0 (0) 0 (0) Eminns nista 0-5-2-0 0 (0) 0 (0) >20, <12.67 cm dbh 28 (28) 0(0) >2.0, >12.67 cm dbh 0 (0) 0 (0) Black currant 0-0.5 833 (567) 2028 (916) 3112:: 1mm 0.5-2.0 83 (83) 194 (100) >2.0, <12.67 cm dbh 0(0) 0 (0) >20, >12.67 cm dbh 0(0) 0 (0) 86 Table A6 (cont'd). Site Species Stratum (m) Exclosure Open Area Black spruce 0-0.5 2639 (709) 1083 (614) Piece mariana 0.5-2.0 2000 (1380) 750 (289) >2.0, <12.67 cm dbh 1167 (l 167) 389 (389) >2.0, >12.67 cm dbh 0(0) 0(0) Flowering dogwood 0-0.5 1028 (628) 56 (56) Camus florida 0.5-2.0 83 (48) 0 (0) >20, <12.67 cm dbh 0(0) 0(0) >2.0, >12.67 cm dbh 0(0) 0(0) Honeysuckle 00.5 1361 (320) 972 (724) mm spp. 0.5-2.0 417 (293) 306 (227) >2.0, <12.67 cm dbh 0(0) 0(0) >2.0, >12.67 cm dbh 0 (0) 0(0) Hop hombeam 0-0.5 0 (0) 56 (56) 95mm minim 0.5-2.0 0(0) 0(0) >20, <12.67 cm dbh 0(0) 28 (28) >20, >12.67 cm dbh 0(0) 0(0) Ironwood 0-0.5 194 (100) 278 (278) 93112111113 muninna 0.5-2.0 139 (139) 83 (83) >20, <12.67 cm dbh 0(0) 0 (0) >20, >12.67 cm dbh 0 (0) 0 (0) Labrador tea 0-0.5 20139 (18740) 18639 (15541) Ledllm. amenlandicllm 0.5-2.0 889 (889) 3000 (1732) >2.0, <12.67 cm dbh 0(0) 0(0) >2.0, >12.67 cm dbh 0(0) 0(0) Mountain maple 0-0.5 56 (56) 0 (0) Ammicamm 0.5-2.0 0(0) 0(0) >20, <12.67 cm dbh 0(0) 0(0) >2.0, >12.67 cm dbh 0(0) 0(0) Paper birch 0-0.5 83 (83) 194 (194) Bemlananxrifsta 0.5-2.0 0(0) 83 (83) >20, <12.67 cm dbh 222 (222) 111 (111) >2.0, >12.67 cm dbh 0 (0) 0 (0) Red maple 0-0.5 1583 (756) 3833 (2821) Ace: mbnlm 0.5-2.0 139 (56) 472 (431) >2.0, <12.67 cm dbh 0(0) 28 (28) >20, >12.67 cm dbh 0(0) 0(0) Red-osier dogwood 0-0.5 278 (139) 972 (890) Camus stalenifera 0.5-2.0 139 (100) 0(0) >20, <12.67 cm dbh 0(0) 0(0) >2.0, >12.67 cm dbh 0(0) 0(0) 87 Table A6 (cont'd). Site Species Stratum (m) Exclosure Open Area Rug spp. 0-0.5 28 (28) 222 (222) 0.5-2.0 0 (0) 0 (0) >20, <12.67 cm dbh 0(0) 0(0) >2.0, >12.67 cm dbh 0(0) 0(0) Smooth gooseberry 0-0.5 0(0) 28 (28) Elbe: hinella 0.5-2.0 0 (0) 0 (0) >20, <12.67 cm dbh 0(0) 0(0) >2.0, >12.67 cm dbh 0 (0) 0(0) Speckled alder 0-0.5 1139 (901) 2667 (747) Alma m 0.5-2.0 1389 (578) 2222 (320) >2.0, <12.67 cm dbh 56 (56) 361 (147) >2.0, >12.67 cm dbh 0 (0) 0(0) Swamp red currant 0-0.5 389 (309) 333 (173) mm: 1:151: 0.5-2.0 56 (56) 0 (0) >20, <12.67 cm dbh 0(0) 0(0) >2.0, >12.67 cm dbh 0 (0) 0 (0) Tamarack 0-0.5 0 (0) 56 (56) Larix 1312121113 0.5-2.0 56 (56) 28 (28) >20, <12.67 cm dbh 83 (83) 0(0) >2.0, >12.67 cm dbh 0 (0) 0(0) Xaccinillm Spp. 0-0-5 83 (48) 0 (0) 0.5-2.0 28 (28) 0(0) >2.0, <12.67 cm dbh 0 (0) 0 (0) >20, >12.67 cm dbh 0(0) 0 (0) Virgin's bower 0-0.5 28 (28) 0 (0) clematis minim 0.5-2.0 o (0) 0(0) >20, <12.67 cm dbh 0 (0) 0(0) >2.0, >12.67 cm dbh 0 (0) 0 (0) White spruce 0-0.5 0 (0) 0 (0) 1219:; 81811.98 0.5-2.0 111 (73) 0 (0) >20, <12.67 cm dbh 111 (73) 0 (0) >20, >12.67 cm dbh 0 (0) 0(0) Willow 0-0.5 28 (28) 56 (56) $3118 spp. 0.5-2.0 0 (0) 139 (139) >2.0, <12.67 cm dbh 28 (28) 139 (139) >2.0, >12.67 cm dbh 0(0) 0 (0) Winterberry holly 0-0.5 0 (0) 28 (28) 11:21 miballata 0.5-2.0 0 (0) 0 (0) >20, <12.67 cm dbh 0 (0) 0 (0) >20, >12.67 cm dbh 0 (0) 0(0) 88 Table A6 (cont‘d). Site Species Stratum (m) Exclosure Open Area Yellow birch 0-0.5 56 (56) 167 (96) 3511113 1111:: 0.5-2.0 56 (56) 28 (28) >20, <12.67 cm dbh 0(0) 0(0) >2.0, >12.67 cm dbh 0 (0) 0 (0) Other 0-0.5 28 (28) 28 (28) 0.5-2.0 0 (0) 0 (0) >20, <12.67 cm dbh 0(0) 0(0) >2.0, >12.67 cm dbh 0(0) 0(0) 'No significant differences (P>0. 10) between exclosure and open area sites for any species and strata with paired t-test (Steel and Torrie 1980). 89 Table A7. Mean stem densities per hectare (and standard error) of non-dominant woody species in exclosure and areas open to browsing sites in the Whitefish River Basin-South study area in the Hiawatha National Forest in Michigan's Upper Peninsula, 1993-1994. Site Species Stratum (m) Exclosure Open Area Alder-leaved buckthorn 0-0.5 56 (56) 361 (147) Bhamnu: alnifblia 0.5-2.0 0 (0) 0 (0) >20, <12.67 cm dbh 0(0) 0(0) >2.0, >12.67 cm dbh 0(0) 0 (0) Altemate—leaved 0-0.5 28 (28) 0 (0) dogwood 0.5-2.0 0(0) 0 (0) Quinn: 81mm >2.0, <12.67 cm dbh 0(0) 0(0) >2.0, >12.67 cm dbh 0(0) 0(0) American elm 0-0.5 83 (83) 111 (111) 11111111: W 0.5-2.0 0 (0) 0 (0) >20, <12.67 cm dbh 0(0) 0(0) >2.0, >12.67 cm dbh 0(0) 0 (0) American mountain ash 0-0.5 250 ( 173) 56 (56) Sotbu: amalgam 05-20 0 (0) 0 (0) >20, <12.67 cm dbh 0(0) 0 (0) >20, >12.67 cm dbh 0 (0) 0 (0) Balsam poplar 0-0.5 306 (306) 806 (806) E99111“: balmifem 0.5-2.0 56 (56) 167 (167) >2.0, < 12.67 cm dbh 0(0) 0 (0) >20, >12.67 cm dbh 0 (0) 0 (0) Beaked hazelnut 0-0.5 417 (376) 28 (28) 9.911111: 99mm 05-2-0 56 (56) 0 (0) >20, <12.67 cm dbh 0(0) 0(0) >2.0, >12.67 cm dbh 0 (0) 0(0) Beech 0-0.5 56 (28) 0 (0) Ban: cranditolia 0.5-2.0 o (0) 0 (0) >20, <12.67 cm dbh 0(0) 0(0) >2.0, >12.67 cm dbh 0(0) 0 (0) Black ash 0-0.5 5889 (5847) 8889 (8639) Eminu: nigta 0.5-2.0 333 (333) 444 (403) >2.0, <12.67 cm dbh 333 (333) 56 (56) >20, >12.67 cm dbh 0(0) 0 (0) Black currant 0-0.5 28 (28) 28 (28) Ribs: hellfire 05-2-0 0 (0) 0 (0) >20, <12.67 cm dbh 0(0) 0(0) >2.0, >12.67 cm dbh 0(0) 0(0) 90 Table A7 (cont'd). Site Species Stratum (m) Exclosure Open Area Black spruce 0-0.5 194 (73) 500 (459) Elma matiana 05-2-0 0 (0) 0 (0) >20, <12.67 cm dbh 0(0) 28 (28) >20, >12.67 cm dbh 28 (28) S6 (56) Choke cherry 0-0.5 111 (111) 139 (139) anu: mainline 0.5-2-0 28 (28) 0 (0) >20, <12.67 cm dbh 0(0) 0(0) >2.0, >12.67 cm dbh 0(0) 0 (0) Honeysuckle 0-0.5 1528 (434) 1472 (420) mm spp. 0.5-2.0 83 (48) 0 (0) >20, < 12.67 cm dbh 0 (0) 0 (0) >20, >12.67 cm dbh 0(0) 0(0) Hop hombeam 0-0.5 167 (167) 389 (227) more xirginiana 05-20 0 (0) 0 (0) >20, <12.67 cm dbh 0(0) 0(0) >2.0, >12.67 cm dbh 0 (0) 0(0) Ironwood 0-0.5 250‘(48) 28 (28) Caminu: carbliniana 05-2-0 0 (0) 0 (0) >20, <12.67 cm dbh 0(0) 0(0) >2.0, >12.67 cm dbh 0 (0) 0 (0) Labrador tea 0-0.5 136] (1361) 1472 (420) Mm amulandisum 0.5-2.0 0 (0) 0 (0) >20, <12.67 cm dbh 0(0) 0(0) >2.0, >12.67 cm dbh 0 (0) 0 (0) Low sweet 0-0.5 361 (217) 2111 (1788) blueberry 0.5-2.0 0 (0) 0 (0) Maximum angusn'fglium >2.0, <12.67 cm dbh 0(0) 0(0) >2.0, >12.67 cm dbh 0 (0) 0 (0) Mountain maple 0-0.5 694 (313) 250 (250) Am snibanlm 0.5-2.0 0 (0) 0 (0) >20, < 12.67 cm dbh 0 (0) 0(0) >2.0, >12.67 cm dbh 0 (0) 0 (0) Paper birch 0-0.5 3000 (928) 3778 (2749) W W 0.5-2.0 56 (28) 83 (83) >20, <12.67 cm dbh 28 (28) 111 (111) >2.0, >12.67 cm dbh 28 (28) 0(0) Red maple 0-0.5 30611 (21759) 12028 (6731) Ace: mbrum 0.5-2.0 28 (28) 28 (28) >20, <12.67 cm dbh 0 (0) 0(0) >2.0, >12.67 cm dbh 28 (28) 0(0) 91 Table A7 (cont'd). Site Species Stratum (m) Exclosure Open Area Red-osier dogwood 0-0.5 139 (28) 389 (100) Camus :tblbnifeza 0.5-2.0 0 (0) 0 (0) . >2.0, < 12.67 cm dbh 0 (0) 0 (0) >20, >12.67 cm dbh 0 (0) 0 (0) Red oak 0-0.5 0 (0) 28 (28) Qumu: rubta 0.5-2.0 0 (0) 0 (0) >20, <12.67 cm dbh 0 (0) 0 (0) >20, >12.67 cm dbh 0 (0) 0 (0) Ruby; spp. 0-0.5 28 (28) 0 (0) 0.5-2.0 0 (0) 0 (0) >20, < 12.67 cm dbh 0 (0) 0 (0) >20, >12.67 cm dbh 0 (0) 0 (0) Speckled alder 0-0.5 444 (444) 639 (598) Ahms mggsa 0.5-2.0 28 (28) 83 (48) >20, <12.67 cm dbh 0 (0) 111 (73) >20, >12.67 cm dbh 0 (0) 0 (0) Swamp red currant 0-0.5 56 (28) 56 (28) Ribs: 1:151: 0.5-2.0 0 (0) 0 (0) >20, <12.67 cm dbh 0 (0) 0 (0) >20, >12.67 cm dbh 0 (0) 0 (0) Tamarack 0-0.5 0 (0) 0 (0) Latizi mm 0.5-2-0 0 (0) 0 (0) >20, <12.67 cm dbh 0 (0) 56 (56) >20, >12.67 cm dbh 0 (0) 28 (28) 31399111111111 spp. 0-0.5 139 (139) 0 (O) 0.5-2.0 o (0) 0 (0) >20, <12.67 cm dbh 0 (0) 0 (0) >20, >12.67 cm dbh 0 (0) 0 (0) Velvet-[caved 0-0.5 111(111) 1611 (1611) blueberry 0.5-2.0 0 (0) 0 (0) WW >2.0,<12.67 cm dbh 111(111) 0(0) >2.0, >12.67 cm dbh 0 (0) 0 (0) Virginia creeper 0-0.5 56 (56) 500 (459) Earthenbciml: 0.5-2-0 0 (0) 0 (0) W13 >2.0, <12.67 cm dbh 0 (0) 28 (28) >20, >12.67 cm dbh 0 (0) 56 (56) White pine 0-0.5 0 (0) 0 (0) Rinu: :tmbu: 0.5-2.0 0 (0) 0 (0) >20, <12.67 cm dbh 0 (0) 0 (0) >20, >12.67 cm dbh 0 (0) 56 (56) 92 Table A7 (cont'd). Species Stratum (m) Exclosure Open Area White spruce 0-0.5 56 (56) 0 (0) Bicea glans: 0.5-2.0 0 (0) 0 (0) >20, <12.67 cm dbh 0 (0) 0 (0) >20, >12.67 cm dbh 0 (0) 0 (0) Willow 0-0.5 56 (56) 0 (0) Sam spp. 0.5-2.0 28 (28) 0 (0) >20, <12.67 cm dbh 0 (0) 0 (0) >20, >12.67 cm dbh 0 (0) 0 (0) Winterberry holly 0-0.5 1500 (542) 944 (823) 11:21 Human 0.5-2.0 11] (73) 167 (167) >2.0, <12.67 cm dbh 28 (28) 56 (56) >20, >12.67 cm dbh 0 (0) 0 (0) Yellow birch 0-0.5 1833 (1792) 1222 (864) Betula lute: 0.5-2.0 0 (0) 0 (0) >20, <12.67 cm dbh 28 (28) 0 (0) >20, >12.67 cm dbh 0 (0) 0 (0) Other 0-0.5 0 (0) 139 (73) 0.5-2.0 0 (0) 0 (0) >20, <12.67 cm dbh 0 (0) 0 (0) >20, >12.67 cm dbh 0 (0) 0 (0) ‘Significantly different (P<0. 10) than open area using paired t-test (Steel and Torrie 1980). 93 Table A8. Mean stem densities per hectare (and standard error) of non-dominant woody species in exclosure and areas open to browsing sites in the Stonington Peninsula-North study area in the Hiawatha National Forest in Michigan's Upper Peninsula, 1993-1994. Sites Species Stratum (m) Exclosure Open Area Alder-leaved 0-0.5 0"(0) 1 1 l (28) buckthorn 0.5-2.0 0 (0) 0 (0) 311mm); alumni; >2.0, < 12.67 cm dbh 0 (0) 0 (0) >20, >12.67 cm dbh 0 (0) 0 (0) Alternate-leaved 0-0.5 0 (0) 56 (56) dogwood 0.5-2.0 0 (0) 0 (0) Com: >2.0, < 12.67 cm dbh 0 (0) 0 (0) ahemifblia >2.0, >12.67 cm dbh 0 (0) 0 (0) American black 0-0.5 28 (28) 0 (0) currant 0.5-2.0 0 (0) 0 (0) Ribs-.5 mm >2.0, <12.67 cm dbh 0 (0) 0 (0) >20, >12.67 cm dbh 0 (0) 0 (0) American mountain ash 0.0.5 0 (0) 28 (28) $51th ameticana 05-20 0 (0) 0 (0) >20, <12.67 cm dbh 0 (0) 0 (0) >20, >12.67 cm dbh 0 (0) 0 (0) American red 0-0.5 0 (0) 0 (0) raspberry 0.5-2.0 0 (0) 139 (139) Rubu: idaeu: >2.0, <12.67 cm dbh 0 (0) 0 (0) >20, >12.67 cm dbh 0 (0) 0 (0) Balsam poplar 0-0.5 5778‘(609) 1417 (625) Ebnulu: balsamifena 0.5-2.0 472 (100) 361 (282) >2.0, <12.67 cm dbh 28 (28) 0 (0) >20, >12.67 cm dbh 139 (56) 111 (73) Black ash 0-0.5 14444 (10729) 43694 (36469) Emmy: mm 0.5-2.0 0 (0) 28 (28) >20, <12.67 cm dbh 0 (0) 28 (28) >20, >12.67 cm dbh 0 (0) 0 (0) Black currant 0-0.5 111] (1070) 2472 (2472) Kiln: lambs 05-20 0 (0) 0 (0) >20, <12.67 cm dbh 0 (0) 0 (0) >20, >12.67 cm dbh 0 (0) 0 (0) Black spruce 0-0.5 28 (28) 28 (28) Rim mam 0.5-2.0 28 (28) 0 (0) >20, <12.67 cm dbh 28 (28) 0 (0) >20, >12.67 cm dbh 56 (56) 0 (0) 94 Table A8 (cont'd). Site Species Stratum (m) Exclosure Open Area Eastern hemlock 0-0.5 28 (28) 0 (0) Inna madmsi: 05-2-0 0 (0) 0 (0) >20, <12.67 cm dbh 0 (0) 0 (0) >20, >12.67 cm dbh 0 (0) 0 (0) Honeysuckle 0-0.5 1167 (1125) 722 (556) Lgniggm spp. 0.5-2.0 0 (0) 139 (139) >2.0, < 12.67 cm dbh 0 (0) 0 (0) >20, >12.67 cm dbh 0 (0) 0 (0) Hop hombeam 0-0.5 83 (83) 139 (139) 9:111: minim 0.5-2.0 0 (0) 0 (0) >20, <12.67 cm dbh 0 (0) 0 (0) >20, >12.67 cm dbh 0 (0) 0 (0) Ironwood 0-0.5 139 (73) 167 (83) Caminu: bamliniana 0.5-2.0 0 (0) 0 (0) >20, <12.67 cm dbh 56 (56) 0 (0) >20, >12.67 cm dbh 0 (0) 0 (0) Mountain maple 0-0.5 2806 (2806) 6111 (6111) Ace: W 0.5-2-0 0 (0) 0 (0) >20, <12.67 cm dbh 0 (0) 0 (0) >20, >12.67 cm dbh 0 (0) 0 (0) Paper birch 0-0.5 694 (437) 56 (56) Bmla nanxntm 0.5-2.0 0 (0) 0 (0) >20, <12.67 cm dbh 28 (28) 28 (28) >20, >12.67 cm dbh 0 (0) 0 (0) My: spp. 0-0.5 250 (250) 139 (139) 0.5-2.0 0 (0) 0 (0) >20, <12.67 cm dbh 0 (0) 0 (0) >20, >12.67 cm dbh 28(0) 83 (83) Prickly gooseben'y 0-0.5 0 (0) 83 (83) macadamia 0.5-2.0 0(0) 0(0) >20, <12.67 cm dbh 0 (0) 0 (0) >20, >12.67 cm dbh 0 (0) 0 (0) Red maple 0-0.5 5000 (1849) 3611 (2183) Ace: nlbnnn 0.5-2.0 0 (0) 0 (0) >20, <12.67 cm dbh 0 (0) 0 (0) >20, >12.67 cm dbh 0 (0) 0 (0) Red-osier dogwood 0-0.5 222 (111) 83 (83) Corny: :tblbnifem 05-2-0 0 (0) 0 (0) >20, <12.67 cm dbh 0 (0) 0 (0) >20, >12.67 cm dbh 0 (0) 0 (0) 95 Table A8 (cont'd). Site Species Stratum (m) Exclosure Open Area Rosa spp. 0-0.5 28 (28) 0 (0) 0.5-2.0 0 (0) 0 (0) >20, <12.67 cm dbh 0 (0) 0 (0) >20, >12.67 cm dbh 0 (0) 0 (0) Speckled alder 0-0.5 28 (28) 28 (28) Alnu: mam 0.5-2.0 0 (0) 0 (0) >20, <12.67 cm dbh 0 (0) 28 (28) >20, >12.67 cm dbh 0 (0) 0 (0) Swamp red currant 0-0.5 83 (83) 1222 (1222) 3118:: 1:151: 0.5-2.0 0 (0) 0 (0) >20, <12.67 cm dbh 0 (0) 0 (0) >20, >12.67 cm dbh 0 (0) 0 (0) Trembling aspen 0-0.5 28 (28) 28 (28) Barium: tremulbide: 05-2-0 0 (0) 0 (0) >20, <12.67 cm dbh 0 (0) 0 (0) >20, >12.67 cm dbh 0 (0) 0 (0) White spruce 0-0.5 28 (28) 0 (0) Rim glans: 05-20 0 (0) 0 (0) >20, < 12.67 cm dbh 0 (0) 0 (0) >20, >12.67 cm dbh 0 (0) 0 (0) Winterberry holly 0-0.5 56 (28) 56 (56) 11:8 minimum 05-2-0 0 (0) 0 (0) >20, <12.67 cm dbh 0 (0) 0 (0) >20, >12.67 cm dbh 0 (0) 0 (0) Yellow birch 0-0.5 56 (28) 83 (48) 8.811113 lum 0.5-2.0 0 (0) 0 (0) >20, < 12.67 cm dbh 0 (0) 0 (0) >20, >12.67 cm dbh 0 (0) 28 (28) Other 0-0.5 56 (28) 0 (0) 0.5-2.0 0 (0) 0 (0) >20, <12.67 cm dbh 0 (0) 0 (0) >20, >12.67 cm dbh 0 (0) 0 (0) ‘Significantly different (P<0.10) from open area using paired t-test (Steel and Torrie 1980). 96 Table A9. Mean stern densities per hectare (and standard error) of non-dominant woody species in exclosure and areas open to browsing sites in the Stonington Peninsula-South study area in the Hiawatha National Forest in Michigan's Upper Peninsula, 1993-1994. Site Species Stratum (m) Exclosure‘ Open Area Alder-leaved 0-0.5 42 (42) 0 (0) buckthom 0.5-2.0 0 (0) 0 (0) Bharnnu: alnifnlia >2.0, <12.67 cm dbh 0 (0) 0 (0) >20, >12.67 cm dbh 0 (0) 0 (0) Altemate—leaved 0-0.5 42 (42) 167 (0) dogwood 0.5-2.0 0 (0) 0 (0) Camus >2.0, < 12.67 cm dbh 0 (0) 0 (0) altemifblia >2.0, >12.67 cm dbh 0 (0) 0 (0) American mountain ash 0-0.5 42 (42) 83 (0) Sbtbu: amerisana 05-20 0 (0) 0 (0) >20, <12.67 cm dbh 0 (0) 0 (0) >20, >12.67 cm dbh 0 (0) 0 (0) Balsam poplar 0-0.5 42 (42) 167 (167) mm: balsamifena 0.5-2.0 o (0) 83 (83) >20, <12.67 cm dbh 0 (0) 42 (42) >20, >12.67 cm dbh 0 (0) 0 (0) Black ash 0-0.5 12167 (10667) 39917 (38333) Eminu: nigta 0-5-2-0 0 (0) 0 (0) >20, <12.67 cm dbh 42 (42) 42 (42) >20, >12.67 cm dbh 42 (42) 0 (0) Black currant 0-0.5 750 (583) 1208 (375) mm 13211an 0.5-2.0 0 (0) 83 (83) >20, <12.67 cm dbh 0 (0) 0 (0) >20, >12.67 cm dbh 0 (0) 0 (0) Black spruce 0-0.5 83 (83) 0 (0) Risen manana 0.5-2.0 0 (0) 0 (0) >20, <12.67 cm dbh 0 (0) 0 (0) >20, >12.67 cm dbh 0 (0) 0 (0) Honeysuckle 0-0.5 583 (250) 2125 (2042) Lgnjggm spp. 0.5-2.0 42 (42) 0 (0) >20, <12.67 cm dbh 0 (0) 0 (0) >20, >12.67 cm dbh 0 (0) 0 (0) Hop hombeam 0-0.5 42 (42) 208 (208) Qsmminiana 0.5-2.0 0(0) 0(0) >20, <12.67 cm dbh 0 (0) 0 (0) >20, >12.67 cm dbh 0 (O) 0 (0) 97 Table A9 (cont'd). Site Species Stratum (m) Exclosure Open Area Ironwood 0-0.5 42 (42) 125 (125) Caminu: catnliniana 05-20 0 (0) 0 (0) >20, <12.67 cm dbh 0 (0) 0 (0) >20, >12.67 cm dbh 0 (0) 0 (0) Mountain maple 0-0.5 1292 (1125) 9833 (9667) Am salsanlm 0.5-2.0 0 (0) 0 (0) >20, <12.67 cm dbh 0 (0) 0 (0) >20, >12.67 cm dbh 0 (0) 0 (0) Paper birch 0-0.5 125 (125) 208 (42) Retina naturism 0.5-2.0 0 (0) 0 (0) >20, <12.67 cm dbh 42 (42) 0 (0) >20, >12.67 cm dbh 0 (0) 42 (42) Prickly gooseberry 0-0.5 0 (0) 42 (42) Ribs: mbbmi 0.5-2.0 0 (0) 0 (0) >20, <12.67 cm dbh 0 (0) 0 (0) >20, >12.67 cm dbh 0 (0) 0 (0) Red maple 0.0.5 15125 (13625) 18583 (17917) Abe: nlbnlm 0.5-2-0 0 (0) 0 (0) >20, <12.67 cm dbh 0 (0) 0 (0) >20, >12.67 cm dbh 0 (0) 0 (0) Red-osier dogwood 0-0.5 83 (83) 333 (333) Coma: siblbnifera 05-2-0 0 (0) 0 (0) >20, <12.67 cm dbh 0 (0) 0 (0) >20, >12.67 cm dbh 0 (0) 0 (0) Rings spp. 0-0.5 0 (0) 125 (125) 0.5-2.0 0 (0) 0 (0) >20, <12.67 cm dbh 0 (0) 0 (0) >20, >12.67 cm dbh 0 (O) 0 (0) Rubll: Spp. 0-0.5 42 (42) 0 (0) 0.5-2.0 0 (0) 0 (0) >20, <12.67 cm dbh 0 (0) 0 (0) >20, >12.67 cm dbh 0 (0) 0 (0) Smooth gooseberry 0-0.5 0 (0) 83 (83) allies hinella 0.5-2.0 o (0) o (0) >20, < 12.67 cm dbh 0 (0) 0 (0) >20, >12.67 cm dbh 0 (0) 0 (0) Speckled alder 0-0.5 0 (0) 42 (42) Alnu: mama 05-20 0 (0) 0 (0) >20, <12.67 cm dbh 0 (0) 0 (0) >20, >12.67 cm dbh 0 (0) 0 (0) 98 Table A9 (cont'd). Site Species Stratum (m) Exclosure Open Area Swamp red currant 0-0.5 0 (0) 208 (125) Ribs: 1:151: 0.5-2.0 0 (0) 0 (0) >20, <12.67 cm dbh 0 (0) 0 (0) >20, >12.67 cm dbh 0 (0) 0 (0) White spruce 0-0.5 0 (0) 83 (83) Piece glam 0.5-2.0 0 (0) 0 (0) >20, <12.67 cm dbh 0 (0) 0 (0) >20, >12.67 cm dbh 0 (0) 0 (0) Winterberry holly 0-0.5 83 (83) 208 (208) 11:21 leniballata 05-20 0 (0) 0 (0) >20, <12.67 cm dbh 0 (0) 0 (0) >20, >12.67 cm dbh 0 (0) 0 (0) Yellow birch 0-0.5 9208 (375) 5667 (5417) m 1mm 0.5-2.0 0 (0) 0 (0) >20, <12.67 cm dbh 42 (42) 0 (0) >20, >12.67 cm dbh 0 (0) 0 (0) Other 0-0.5 0 (0) 125 (125) 0.5-2.0 0 (0) 0 (0) >20, < 12.67 cm dbh 0 (0) 0 (0) >20, >12.67 cm dbh 0 (0) 0 (0) ‘No significant differences (P>0.10) between exclosure and open area sites for any species and strata with paired t-test (Steel and Torrie 1980). 99 Table A10. Mean stem densities per hectare (and standard error) of non-dominant woody species that were significantly different (P<0.10) within site and stratum among study areas (Whitefish River Basin-North and -South [WRB-North and -South] and Stonington Peninsula-North and -South [SP-North and -South]) in the Hiawatha National Forest in Michigan's Upper Peninsula, 1993-1994. Site Study Area Species Stratum (m) Exclosure Open Area WRB-North Alder-leaved buckthorn 0-0.5 1972A' (1188) 2750A (2014) Rhamnu: minim 0.5-2.0 1194” (989) 1917" (1792) Balsam poplar 0-0.5 0A (0) ----° Panulu:bal:ami£era 0.5-2.0 0" (0) .....- >20, >12.67 cm dbh 28" (28) ...... Black ash 00.5 0" (0) 0" (0) Eminu: nizta Black spruce 0-0.5 2639A (709) ----- Risen mariana 0.5-2.0 2000b (1380) 750" (289) Paper birch 0-0.5 83A (83) 194" (194) Benin napalm Red maple 0.5-2.0 139" (56) ..---- Abe: :acshanlm Speckled alder 0.5-2.0 1389" (578) 2222A (320) Ainu: mass Winterberry holly 005 o A (o) ..---- 11:8 xeniballata WRB-South Alder-leaved buckthorn 0-0.5 56AB (56) 361AB (147) Bhamnu: alnifllll'a 0.5-2.0 0 (0) 0 (0) Balsam poplar 0-0.5 306AB (306) ----- Ennulllsbalsamifcta 0-5-2.0 56 (56) mm >20, >12.67 cm dbh 0 (0) Black ash 0-0.5 5889 (5847) 8889 (8639) Eminu: aim Black spruce 0-0.5 194AB (73) mm Eicea mariana 0.5-2-0 0 (0) 0 (0) Paper birch 0-0.5 3000B (928) 3778 (2749) Bemla maxim Red maple 0.5-2.0 28 (28) ----- Abetnlbnlm Table A10 (cont’d). 100 Site Study Area Species Stratum (m) Exclosure Open Area WRB-South Speckled alder 0.5-2.0 28 (28) 83AB(48) (cont’d-) Alnusnlzbsa Winterberry holly 0-0.5 1500B (542) --- 11:8 misallata SP-North Alder-leaved buckthom 0-0.5 OB (0) 1 11AB(28) Bhamnu: alumni: 0.5-2.0 o (0) 0 (0) Balsam poplar 0-0.5 5778B (609) ----- Ronalu: balsamifm 0.5-2.0 472 (100) ...... >2.0, >12.67 cm dbh 28 (28) Black ash 0-0.5 14444 (10729) 43694 (36469) Eminu: aim Black spruce 0-0.5 288 (28) mm Rice: mariana 0.5-2.0 28 (28) 0 (0) Paper birch 0-0.5 694AB (437) 56 (56) 8:11.113 1239mm Speckled alder 0.5-2.0 0 (0) 0A (0) Alnu: mans: Red maple 0.5-2.0 0 (0) mm Abe: nlbnlm Winterberry holly 0-0.5 56AB (28) --..-- 11:21 Manuals SP-South Alder-leaved buckthorn 0-0.5 42AB (34) 0B (0) Rhamnll: alnifnlia 05-2-0 0 (0) 0 (0) Balsam poplar 0-0.5 42AB (34) um- anulusbalsamiim 0.5-2.0 0 (0) ..---- >2.0, >12.67 cm dbh 0 (0) ---- Black ash 0-0.5 12167 (8709) 39917 (31299) Eminu: nim Black spruce 0-0.5 83AB (68) ----- Pineal mariana 05-2-0 0 (0) 0 (0) Paper birch 0-0.5 125A (102) 208 (34) Bamlananxfifm 101 Table A10 (cont’d). Site Study Area Species Stratum (m) Exclosure Open Area SP-South Red maple 0.5-2.0 0 (0) ...... (cont’d.) Abernlbnlm Speckled alder 0.5-2.0 0 (0) DAB (0) Alnu: {Hausa Winterberry holly 0-0.5 83AB (68) ...... Ilsa miballata 'Means with different letters within a site, species, and stratum were significantly different (P<0.10) among study areas with the Kruskal-Wallis one-way analysis of variance (Siegel and Castellan 1988) using SYSTAT (1992) and the Kruskal-Wallis multiple comparison test (Siegel and Castellan 1988). eruskal-Wallis multiple comparison test (Siegel and Castellan 1988) was unable to detect where the significant difference occurred within a site, species, and stratum among the study areas as detected by the Kruskal-Wallis one-way analysis of variance (Siegel and Castellan 1988) using SYSTAT (1992). °Species not identified in any study area for this site type. 102 Table A1 1. Mean absolute (AF) and relative frequencies (RF) (and standard errors [SE]) of herbaceous species in exclosure and areas open to browsing sites in the Whitefish River Basin-North study area in the Hiawatha National Forest in Michigan's Upper Peninsula, 1993-1994. Site Exclosure Open Area AF‘ RF AF RF Species (SE) (SE) (SE) (SE) Am: spp. 20.8 1.4 37.5 2.3 (11.0) (0.8) (19.1) (1.1) Bunchberry 79.2 5.4 87.5 5.8 Cnmu: canadensi: (4.2) (0.4) (12.5) (1-1) Canada mayflower 66.7 4.5 70.8 4.7 Maianthemum W (4.2) (0-4) (8.3) (0-6) Cinnamon fern 0 0 4.2 0.3 Qsmunda cinnamnmea (0) (0) (4.2) (0.3) Common wood sorrel 0 0 4.2 0.3 Quail: mnntana (0) (0) (4-2) (0-3) Crested wood fern 4.2 0.3 8.3 0.5 Mariam (4.2) (0.3) (8.3) (0.5) Dewberry 62.5 4.3 62.5 4.4 Rubu: hisbidu: (31 -5) (2-2) (25-0) (1 .9) Dwarf enchanter‘s nightshade 4.2 0.3 0 0 Clram albina (4.2) (0-3) (0) (0) Ebulsenlm Spp. 4.2 0.3 4.2 0.2 (4.2) (0.3) (4.2) (0.2) Fragile fern 29.2 2.0 25.0 1.5 mm (8.3) (0.6) (14.4) (0.8) Fringed brome 4.2 0.3 0 O Emu: 911121111: (42) (0.3) (0) (0) Goldenrod 41.7 2.7 50.0 3.5 Solidago spp. (22.0) (1.4) (19.1) (1.6) Goldthread 70.8 4.7 91.7 6.1 901211: mnhndiaa (23-2) (1 -S) (4-2) (0-4) Grass spp. 8.3 0.5 62.5 4.2 (8.3) (0.5) (21.7) (1.5) 103 Table A11 (cont'd). Site Exclosure Open Area AF RF AF RF Species (SE) (SE) (SE) (SE) Hawkweed 8.3 0.6 8.3 0.6 Hiemiunl Spp. (8.3) (0.6) (4.2) (0.3) Intermediate wood fern 0 0 4.2 0.3 Damien: lntmneslla (0) (0) (4.2) (0-3) Jewelweed 8.3 0.6 0 0 Impatien: Spp. (8-3) (0-6) (0) (0) Joe-pye weed 20.8 1.4 16.7 1.2 Eunatbrium spp. (1 1.0) (0.8) (1 1.0) (0.9) Large-leaved aster 12.5 0.8 4.2 0.3 mammalian: (7.2) (0.5) (4.2) (0.3) Manna grass 0 0 20.8 1.2 Gilliam 8139- (0) (0) (20-8) (1-2) Marsh bedstraw 25.0 1.7 25.0 1.8 caliumaalustre (14.4) (1.0) (12.5) (0.9) Marsh marigold 0 0 12.5 1.0 Caitlin balumi: (0) (0) (12-5) (1 -0) Moss spp. 100 6.8 100 6.6 (0) (0-2) (0) (0-6) Naked miterwort 95.8 6.5 62.5 4.4 Milena nude (42) (0-5) (26-0) (29) Oak fern 12.5 0.9 20.8 1.4 omnocamiumspp. (7.2) (0.9) (11.0) (0.7) W spp. 4.2 0.3 8.3 0.5 (4.2) (0.3) (4.2) (0.3) Pamela spp. 12.5 0.9 8.3 0.5 (7.2) (0.5) (8.3) (0.5) Rattlesnake fern 8.3 0.6‘ 33.3 2.1 mm (4.2) (0.3) (11.0) (0.6) Rattlesnake plantain 0 0 8.3 0.5 Gaadxeta SPP- (0) (0) (4.2) (0.3) Table All (cont'd). 104 Site Exclosure Open Area AF RF AF RF Species (SE) (SE) (SE) (SE) Royal fern 4.2 0.3 0 0 Qamunda mall: (42) (0-3) (0) (0) Sedge 91.7 6.2‘ 87.5 5.7 Cam spp. (4.2) (0.1) (7.2) (0.1) Self-heal 8.3 0.6 8.3 0.5 Pruneila mm (8-3) (0-6) (4-2) (0-3) Showy lady's slipper 0 0 8.3 0.6 ann'neuiumncainae (0) (0) (4.2) (0.3) Small-flowered cranberry 4.2 0.3 8.3 0.5 labelnlum 9mm: (42) (0.3) (8-3) (0-5) Snowberry 75.0 5.0‘ 25.0 1.5 Qauithetia hisnlduia (12.5) (0-7) (14-4) (0-9) Starflower 37.5 2.5 62.5 3.8 Irlemaii: bnmaii: (12-5) (0-7) (31-5) (1 .9) Strawberry 83.3 5.6 66.7 4.3 mm spp. (1 1.0) (0.6) (16.7) (0.8) Sundew 4.2 0.3 0 0 12mm 5w (4.2) (0.3) (0) (0) Sweet coltsfoot 8.3 0.6 8.3 0.5 Parasite: naimanl: (4.2) (0.3) (8-3) (0.5) Sweet-scented bedstraw 62.5 4.2 20.8 1.2 Qaiillm ttlflbmm (0) (0- 1) (20-8) (1 -2) Thistle 45.8 3.1 54.2 3.6 mum spp. (4.2) (0.2) (20.8) (1 .5) Three-leaved Solomon's seal 75.0 5.0 66.7 4.2 Smliablnattifbiia (19.1) (1.2) (22-0) (1-1) Trailing arbutus 0 0 4.2 0.2 51118323 man: (0) (0) (4-2) (0.2) Twinflower 91.7 6.2 75.0 4.9 Linnaea Email: (83) (0-5) (7.2) (0.1) 105 Table A11 (cont'd). Site Exclosure Open Area AF RF AF RF Species (SE) (SE) (SE) (SE) Violet 75.0 5.0 83.3 5.5 M19]; spp. (12.5) (0.7) (8.3) (0.3) Wild sarsaparillo 37.5 2.6‘ 62.5 4.0 Aralia nudlsauil: (12-5) (0-9) (14.4) (0-8) Wintergreen 16.7 1.1 12.5 0.7 fiauithatia mumben: (8.3) (0-6) (7-2) (04) Wood anemone 12.5 0.8 4.2 0.2 Ansunbne quinaubfbiia (12-5) (0-8) (4.2) (02) Wood Sorrel 16.7 1.2 0 0 Quail: 5913- (16-7) (1 .2) (0) (0) Other 29.2 2.0 29.2 1.7 (4.2) (0.2) (23.2) (1.3) 'Significant differences between exclosure and open area sites were not determined for absolute frequencies, only relative frequencies. ‘Significantly different (P<0.10) from open area with paired t-test (Steel and Torrie 1980). 106 Table A12. Mean absolute (AF) and relative frequencies (RF) (and standard errors [SE]) of herbaceous species in exclosure and areas open to browsing sites in the Whitefish River Basin-South study area in the Hiawatha National Forest in Michigan's Upper Peninsula, 1993-1994. Site Exclosure Open Area AF‘ RF AF RF Species (SE) (SE) (SE) (SE) Aster spp. 12.5 0.9 25.0 1.8 (0) (0) (14.4) (1.1) Bedstraw 0 0 16.7 1.1 Gaillun SW (0) (0) (16-7) (1 . 1) Bracken fern 4.2 0.3 4.2 0.3 Biennium aaulilnum (4-2) (0.3) (4-2) (0-3) Bugleweed 4.2 0.3 8.3 0.6 mam SW (4.2) (0-3) (4-2) (0-3) Bunchberry 37.5 2.9 62.5 4.5 Commmadansi: (19.1) (1.6) (31-5) (2.2) Canada mayflower 70.8 5.1‘ 91.7 6.3 Maianthemum canadenm (1 1.0) (0.6) (4-2) (0-2) Cinnamon fern 8.3 0.6 4.2 0.3 mm cinnamnmea (8.3) (0-6) (4.2) (0.3) Columbine 0 0 4.2 0.3 Aguiiegia Spp. (0) (0) (4-2) (0.3) Club-spur orchid 4.2 0.3 0 0 Habenatia biaxeiiata (4-2) (0-3) (0) (0) Crested wood fern 16.7 1.2 0 O Damian: ctlstata (8.3) (0-6) (0) (0) Dewberry 54.2 4.0 66.7 4.5 Rubushisnislu: (11-0) (0.8) (16-7) (0.9) mm: spp. 4.2 0.3 0 0 (4.2) (0-3) (0) (0) Emiismm spp. 16.7 1.1 4.2 0.3 (11.0) (0.8) (4.2) (0.3) Fragile fern 4.2 0.3 0 0 W fragiii: (4-2) (0-3) (0) (0) 107 Table A12 (cont'd). Site Exclosure Open Area AF RF AF RF Species (SE) (SE) (SE) (SE) Fringed polygala 75.0 5. 6 79.2 5.5 Eelxzalaoaucitolia (12.5) (12) (4.2) (0.5) Golden ragwort 4.2 0.3 12.5 0.8 Smeaib entail: (4-2) (0-3) (12.5) (0-8) Goldenrod 29.2 2.1 33.3 2. 2 Saildaga SPP- (16.7) (1 2) (15-0) (0 9) Goldthread 95.8 7.0 91 .7 6.4 Camlsmniandiba (4.2) (0.6) (8-3) (0-8) Grass spp. 62.5 4.4 54.2 3. 6 (31.5) (2.2) (25.3) (1 6.) Interrupted fern 4.2 0.3 0 0 Qstmlnda Wine (42) (0.3) (0) (0) Jack-in-the-pulpit 0 0 16.7 1 .1 Ariaaalna Spp. (0) (0) (16-7) (1 - 1) Jewelweed 0 0 20.8 1. 3 Impatisn: SW (0) (0) (20-8) (1 3) Joe-pye weed 12.5 0. 9 8. 3 0.6 Eunalbnum Spp. (7.2) (0 5) (4 2) (0.3) Large-leaved aster 0 0 12.5 0. 8 Am Win: (0) (0) (12-5) (0 8) Lesser pyrola 0 0 12.5 0. 9 Pamela minor (0) (0) (12-5) (0 9) Long beech fern 0 0 4. 2 0.3 Ihbbauatl: nhsaomstl: (0) (0) (4 2) (0-3) Marsh bedstraw 4.2 0. 3 0 0 fiallum 2111qu (4.2) (0 3) (0) (0) Marsh fern 8.3 0.6 0 0 111mm (8.3) (0.6) (0) (0) Marsh marigold 0 O 4. 2 O. 3 Caitlin naiustrl: (0) (0) (4- 2) (0 3) 108 Table A12 (cont'd). Site Exclosure Open Area AF RF AF RF Species (SE) (SE) (SE) (SE) Moss spp. 100 7.3 100 6.9 (0) (0-4) (0) (03) Naked miterwort 83.3 6.2 91.7 6.3 Mimi]: and: (11.0) (1.1) (8.3) (04) Oak fern 8.3 0.6 0 0 fixmnacamium 5917- (8.3) (0-6) (0) (0) One-flowered pyrola 4.2 0.3 0 0 m5 unlflbta (4-2) (0-3) (0) (0) Wm: spp. 0 0 4.2 0.3 (0) (0) (4-2) (0-3) finals spp. 16.7 1.2 25.0 1.8 (8.3) (0.6) (14.4) (1.0) Rattlesnake fern 16.7 1.2 37.5 2.5 mm (4.2) (0.3) (26.0) (1.8) Rough bedstraw 54.2 3.9 37.5 2.6 Qailumasbtalium (15.0) (0.9) (0) (0.1) Royal fern 16.7 1.2 12.5 0.9 asmunda mall: (11.0) (0-7) (12-5) (0-9) Sedge 83.3 6.2 79.2 5.6 Cm); spp. (11.0) (1.1) (20.8) (1.6) Self-heal 8.3 0.6 4.2 0.3 Emnalla 1011831215 (33) (0-6) (4.2) (03) Sensitive fern 8.3 0.6 4.2 0.3 anclba sensibiii: (4.2) (0-3) (4-2) (0-3) Small-flowered cranberry 0 0 16.7 1.1 Masainlurn 98mm (0) (0) (16-7) (1-1) Snowberry 75.0 5.6 62.5 4.4 Qauithetiahisnidula (14-4) (1-3) (19-1) (1-4) Starflower 91.7 6.7 83.3 5.8 Inanlails bbmail: (8.3) (0.8) (4-2) (0.4) 109 Table A12 (cont'd). Site Exclosure Open Area AF RF AF RF Species (SE) (SE) (SE) (SE) Strawberry 20.8 1.5 33.3 2.1 Eragada spp. (15.0) (1.1) (33.3) (2.1) Sweet coltsfoot 0 0 4.2 0.3 Remus: naimatu: (0) (0) (4-2) (0-3) Sweet-scented bedstraw 4.2 0.3 0 0 Galium trifoium (4-2) (0.3) (0) (0) Tall meadow rue 4.2 0.3 0 0 lilaiibmlm mbmmum (4.2) (0.3) (0) (0) Thistle 33.3 2.4 20.8 1.5 Cit:ium Spp. (1 1.0) (0.7) (11.0) (0.8) Three-leaved Solomon's seal 37.5 2.8 20.8 1.4 Smliabina nifblia (7-2) (0-5) (15-0) (1 -0) Trailing arbutus 4.2 0.3 8.3 0.6 Estimation: (4.2) (0.3) (8.3) (0.6) Twinflower 50.0 3.8 45.8 3.3 Linnaea Email: (289) (2-3) (23-2) (1 -6) Twisted stalk 0 0 12.5 0.8 Stratum amnlaxlfbiiu: (0) (0) (12-5) (0-8) Violet 45.8 3.4 50.0 3.4 Mg spp. (8.3) (0.8) (25.0) (1.7) Wild ginger 0 0 4.2 0.3 Asanlm madame (0) (0) (4-2) (03) Wild sarsaparillo 29.2 2.0 33.3 2.1 Atalla nudibauii: (18.2) (1-3) (33-3) (2-1) Wintergreen 8.3 0.7 20.8 I .5 Clauithan'a mumbsn: (8.3) (0.7) (11-0) (0.8) Wood Sorrel 25.0 1.7 0 0 Ozailssrm- (19-1) (1-3) (0) (0) Table A12 (cont’d). 110 Site Exclosure Open Area AF RF AF RF Species (SE) (SE) (SE) (SE) Yellow lady’s slipper 4.2 0.3 0 0 Wm calbebiu: (4-2) (0-3) (0) (0) Other 4.2 0.3 4.2 0.3 (4.2) (0.3) (4.2) (0.3) 'Significant differences between exclosure and open area sites were not determined for absolute frequencies, only relative frequencies. ‘Significantly different (P<0.10) from open area with paired t-test (Steel and Torrie 1980). 111 Table A13. Mean absolute (AF) and relative frequencies (RF) (and standard errors [SE]) of herbaceous species in exclosure and areas open to browsing sites in the Stonington Peninsula-North study area in the Hiawatha National Forest in Michigan's Upper Peninsula, 1993-1994. Site Exclosure Open Area AF“ RF AF RF Species (SE) (SE) (SE) (SE) Anemone spp. 0 0 6.1 0.5 (0) (0) (3-0) (0-3) Arrow arum 0 0 3.0 0.4 Beltranda xitginlsza (0) (0) (3.0) (04) A519,: spp. 37.5 2.8 12.1 1.4 (12.5) (1.0) (6.1) (0.7) Bedstraw 4.2 0.4 O 0 aaiium Spp. (4-2) (0.4) (0) (0) Blunt-lobed woodsia 0 0 3.0 0.2 mm bbtusa (0) (0) (3-0) (0-2) Boot's wood fern 41.7 3.1 21.2 1.7 Damian: bacilli (15.0) (1 .2) (10-9) (0-9) Bracken fern 0 0 6.1 0.6 Biennium anuliinum (0) (0) (6- 1) (0-6) Bugleweed 8.3 0.5 15.2 1.1 Lyman: spp. (8.3) (0.5) (10.9) (0.7) Bulbet fern 4.2 0.3 0 0 93151991211: buibifena (4.2) (0.3) (0) (0) Bunchberry 29.2 1.9 21 .2 1.3 Camuscanadensi: (18-2) (1-1) (21-2) (1-3) Buttercup 0 0 42.4 4.9 Eanunaulll: SW (0) (0) (23-7) (29) Canada mayflower 91.7 7.1 72.7 6.9 Maianthsmumcanadanss (4.2) (1.0) (15-7) (1.9) Cinquefoil 4.2 0.3 0 0 Palemlila SW (42) (0-3) (0) (0) Clover 4.2 0.3 9.1 1.0 Irifoilum Spp. (4.2) (0.3) (5.2) (0.6) 112 Table A13 (cont’d). Site Exclosure Open Area AF RF AF RF Species (SE) (SE) (SE) (SE) Club moss 4.2 0.4 0 0 Lxmnadlum SW (4.2) (0-4) (0) (0) Crested wood fern 4.2 0.3 0 0 Duablanssnstala (4.2) (0.3) (0) (0) Dandelion 4.2 0.3 9.1 1.0 Iataxaeum Spp. (4-2) (0.3) (5.2) (0.6) Dewberry 33.3 2.2 33.3 2.1 Rubu: hisnidu: (22-0) (1 -3) (33-3) (2-1) Damned; spp. 25.0 2.0 6.1 0.4 (12.5) (1.1) (6.1) (0.4) Dwarf enchanter's nightshade 29.2 1.8 21.2 1.3 Cirsaea 81121113 (292) (1 -8) (21-2) (1 -3) Equisanlm spp. 50.0 3.6 36.4 2.7 (26.0) (1.8) (27.8) (1 .7) False Solomon's seal 4.2 0.3 0 0 Smiiaalna W (4.2) (0.3) (0) (0) Golden ragwort 4.2 0.3 0 0 Ssnesib aurau: (4-2) (0-3) (0) (0) Goldenrod 41.7 3.3 33.3 3.1 5121111389 SPP- (8-3) (0.9) (12- 1) (1 -3) Grass spp. 37.5 2.9 60.6 5.4 (0) (0.3) (21.2) (1.9) Grass/sedge spp. 75.0 5.6 42.4 4.1 (7.2) (0.4) (21.9) (2.6) Hawkweed 16.7 1 .4 60.6 6.7 W spp. (I 1.0) (1.0) (23.7) (3.2) Hooked crowfoot 0 0 6.1 0.4 Ranunbulu: mam: (0) (0) (6.1) (04) Intermediate wood fern 12.5 0.8 6.1 0.5 mm (7.2) (0.4) (3.0) (0.3) 113 Table A13 (cont'd). Site Exclosure Open Area AF RF AF RF Species (SE) (SE) (SE) (SE) Marsh bedstraw 8.3 0.5 3.0 0.2 cannmaaluatre (8.3) (0.5) (3.0) (0.2) Milkweed 0 0 3.0 0.2 Assknia: Spp. (0) (0) (3.0) (02) Mint 4.2 0.3 3.0 0.2 Mentha spp. (4.2) (0.3) (3.0) (0.2) Moss spp. 91.7 7.0 93.9 9.2 (8.3) (0.9) (3.0) (1 .8) Naked miterwort 54.2 3.8 39.4 2.8 Mitena nnda (23.2) (1.2) (30.8) (1.9) Oak fern 54.2 4.0 12.1 0.8 Gmnbcamium Spp. (15.0) (1 .2) (12.1) (0.8) Ostrich fern 4.2 0.3 0 0 Matiaubala minim (4-2) (0-3) (0) (0) Mala spp. 29.2 2.1 3.0 0.2 (15.0) (1 .0) (3.0) (0.2) Rattlesnake fern 58.3 4.3‘ 24.2 2.2 3.121135111111111 xltginianllm (1 1-0) (0-7) (8-0) (0.7) Rough bedstraw 33.3 2.2‘ 42.4 3.4 aaiiumasnmlillm (22.0) (1.3) (21.2) (1.2) Sedge 45.8 3.9 33.3 3.8 gm spp. (23.2) (1.9) (16.9) (2.0) Self-heal 25.0 1 .8 15.2 1 .4 Pnlneila inland: (7-2) (0-4) (3-0) (04) Sensitive fern 4.2 0.3 0 0 finale: sensibiii: (4-2) (0-3) (0) (0) Silvery glade fern 25.0 1.8 3.0 0.2 Athxriumthalxtuatlbida: (12.5) (0.9) (3.0) (0.2) Skullcap 0 0 6.1 0.4 Sautaliaria 8139- (0) (0) (6- 1) (0-4) 114 Table A13 (cont'd). Site Exclosure Open Area AF RF AF RF Species (SE) (SE) (SE) (SE) Spinulose wood fern 8.3 0.7 3.0 0.3 Damien: mm (8-3) (0-7) (3-0) (0-3) Spurred gentian 8.3 0.7 18.2 1.9 Halsnla deflm (8.3 (0.7) (182) (1 .9) Starflower 62.5 4.7 51.5 4.2 Inmtail: bateaii: (14.4) (1-2) (26-9) (2- 1) Strawberry 20.8 1 .4 15.2 1 .0 Engarja spp. (11.0) (0.7) (15.2) (1.0) Sweet coltsfoot 45.8 3.8 21.2 2.6 Emails: nairnatu: (22-0) (2-0) (21-2) (2.6) Tall buttercup 12.5 1 .1 0 0 Ranunbulllsaslzi: (12-5) (1.1) (0) (0) Thistle 33.3 2.4 33.3 3.2 Qirsjmn spp. (1 1.0) (0.7) ( 10.9) (1.3) Twinflower 4.2 0.3 6.1 0.4 Linnaea banaaii: (4-2) (0.3) (6- 1) (0-4) Twisted stalk 4.2 0.4 0 0 W ambieziifaiiu: (4.2) (0-4) (0) (0) Violet 75.0 5.5 72.7 6.8 £1913 spp. (12.5) (0.3) (10.5) (1.0) Wild sarsaparillo 66.7 5.1 36.4 2.6 Ataiia nudlballii: (4.2) (0.7) (27-8) (1 -7) Wood anemone 20.8 1.7 27.3 2.7 Anemoaeauinouerolia (15.0) (1.2) (5.2) (0.9) Other 12.5 1.0 21.2 1.9 (7.2) (0.6) (10.9) (1.1) ‘Significant differences between exclosure and open area sites were not determined for absolute fi'equencies, only relative fi'equencies. ‘Significantly different (P<0.10) from open area with paired t-test (Steel and Torrie 1980). 115 Table A 14. Mean absolute (AF) and relative frequencies (RF) (and standard errors [SE]) of herbaceous species in exclosure and areas open to browsing sites in the Stonington Peninsula-South study area in the Hiawatha National Forest in Michigan's Upper Peninsula, 1993-1994. Site Exclosure Open Area AF‘ RF” AF RF Species (SE) (SE) (SE) (SE) Bedstraw 0 0 6.3 0.4 Qaiium 5913- (0) (0) (6-3) (0-4) Boot's wood fern 12.5 1.2 6.3 0.5 W bcumii (12-5) (1-2) (6-3) (0-5) Bugleweed 31.3 2.3 43.8 3.1 Lymrms spp. (31.3) (2.3) (6.3) (0.3) Bunchberry 43.8 3.9 31.3 2.2 Camus madmsi: (18-8) (2- 1) (6-3) (0.4) Canada mayflower 81.3 6.8 68.8 4.9 Maianthemum canadanss (6.3) (0-4) (6.3) (0.3) Coralroot 6.3 0.6 6.3 0.4 Catalinrhiza 5139- (6.3) (0-6) (6.3) (0-4) Crested wood fern 6.3 0.5 25.0 1.8 mm (6.3) (0.5) (12.5) (1.0) Dewberry 50.0 4.4 87.5 6.3 Rubll: hlsnldus (12-5) (1 -6) (0) (02) Men's spp. 0 0 12.5 0.9 (0) (0) (12.5) (0.9) Dwarf enchanter's nightshade 37.5 3.0 62.5 4.5 Citbaaaainlna (12.5) (0.6) (12-5) (1.1) Eaulsanlm spp. O 0 18.8 1.3 (O) (0) (18.8) (1.3) Fragile fern 18.8 1.4 6.3 0.4 stiamaris Mill: (18.8) (1 -4) (6-3) (0-4) Fringed polygala 0 0 6.3 0.4 Pawn pausifblia (0) (0) (6.3) (04) Golden ragwort 6.3 0.5 0 0 Smash am (63) (0-5) (0) (0) 116 Table A14 (cont'd). Site Exclosure Open Area AF RF AF RF Species (SE) (SE) (SE) (SE) Goldenrod 6.3 0.5 12.5 0.9 flamingo spp. (6.3) (0.5) (12.5) (0.9) Goldthread 43.8 3.5 62.5 4.4 Calms emaniandiba (18-8) (1 - 1) (12.5) (0-7) Grass spp. 50.0 3.7 68.8 4.8 (50.0) (3.7) (31.3) (2.1) Grass/sedge spp. 18.8 1.8 12.5 0.9 (18.8) (1.8) (12.5) (0.9) Intermediate wood fern 31.3 3.0 37.5 2.8 Domini: lntannadia (31.3) (3-0) (37-5) (2-8) Interrupted fern 6.3 0.6 0 0 Qsmunda clanbnlana (6.3) (0-6) (0) (0) Jack-in-the—pulpit 37.5 2.8 31.3 2.3 m spp. (37.5) (2.8) (18.8) (1.4) Jewelweed 12.5 0.9 12.5 0.9 Impatien: 8139- (12-5) (0-9) (0) (0) Joe-pye weed 6.3 0.5 0 0 Eunalbtium 8913- (6.3) (0-5) (0) (0) Long beech fern 18.8 1.4 37.5 2.7 mm (18.8) (1.4) (12.5) (1.0) Marsh fern 12.5 0.9 12.5 0.9 Iliclxntetisaalmia (12.5) (0.9) (12.5) (0.9) Marsh skullcap 0 0 6.3 0.4 Ssutaiiatia eniibbifbiia (0) (0) (6-3) (0-4) Mint 6.3 0.5 6.3 0.4 Mgmha spp. (6.3) (0.5) (6.3) (0.4) Moss spp. 93.8 7.8 93.8 6.7 (6.3) (0.5) (6.3) (0.2) Naked miterwort 93.8 7.8 100 7.2 Mltaiia nuda (6-3) (0.5) (0) (0-3) 117 Table A14 (cont'd). Site Exclosure Open Area AF RF AF RF Species (SE) (SE) (SE) (SE) Oak fern 43.8 3.3 18.8 1.4 fixmnasamium Spp. (31.3) (2.2) (18-8) (1 .4) finals spp. 0 0 6.3 0.4 (0) (0) (6.3) (0.4) Rattlesnake fern 31.3 2.8 31.3 2.3 Banahlumxltglnlanllm (18-8) (1.9) (18.8) (1-4) Rattlesnake plantain 0 0 18.8 1.3 Gambian SPP- (0) (0) (6-3) (0-4) Rough bedstraw 68.8 5.7 68.8 4.9 Qallunl asmilum (6.3) (0-2) (18-8) (12) Royal fern 0 0 31.3 2.2 Qsmunda mail: (0) (0) (31-3) (2-2) Sedge 12.5 0.9 31.3 2.2 Cm spp. (12.5) (0.9) (18.8) (1.3) Self-heal O 0 18.8 1 .3 Enlnaiia mind: (0) (0) (18-8) (1 -3) Sensitive fern 0 0 25.0 1.7 Qnmiaa :snsibiil: (0) (0) (25.0) (1 -7) Silvery glade fern 0 0 18.8 1.4 Aihxmlm thsbmtan'aida: (0) (0) (18-8) (1-4) Starflower 81.3 7.1 68.8 5.0 Idsntaii: bnraaii: (18-8) (2-5) (18-8) (1-5) Strawberry 12.5 1.1 6.3 0.5 W spp. (0) (0.1) (6.3) (0.5) Thistle 6.3 0.5 0 0 Quinn) spp. (6-3) (0.5) (0) (0) Wm spp. 6.3 0.5 0 0 (6-3) (0-5) (0) (0) Turtlehead 6.3 0.5 0 0 (Ensign: SW (6.3) (0-5) (0) (0) 118 Table A14 (cont'd). Site Exclosure Open Area AF RF AF RF Species (SE) (SE) (SE) (SE) Twinflower 0 0 6.3 0.4 Linnaea bbteaii: (0) (0) (6-3) (0-4) Twisted stalk 6.3 0.6 0 0 W amblaxifailu: (6.3) (0-6) (0) (0) Violet 75.0 6.2 75.0 5.3 Viola spp. (12.5) (0.2) (25.0) (1.6) White adders mouth 0 0 6.3 0.4 Maiaxi: bmhmda (0) (0) (6-3) (04) Wild sarsaparillo 56.3 4.7 31.3 2.3 Atalia nudisauii: (6-3) (0-1) (18-8) (1 -4) Wood anemone 12.5 1.2 0 0 Anemone aulnausfillia (12-5) (1-2) (0) (0) Wood sorrel 43.8 4.3 50.0 3.7 933115 spp. (43.8) (4.2) (50.0) (3.7) Yellow lady's slipper 0 0 6.3 0.4 Winn baibcbiu: (0) (0) (6-3) (0-4) Other 12.5 0.9 6.3 0.4 (12.5) (0.9) (6.3) (0.4) 'Signlficant differences between exclosure and open area sites were not determined for absolute frequencies, only relative frequencies. bNo significant differences (P>0.10) of relative frequencies between exclosure and open area sites for any species with the paired t-test (Steel and Tom'e 1980). Table A15. Relative frequency (and standard error) of herbaceous species that were significantly different (P<0.10) among study areas (Whitefish River Basin-North and -South [WRB-North and -South] and Stonington Peninsula-North and -South [SP-North and -South]) in the Hiawatha National Forest in Michigan's Upper Peninsula, 1993-1994. Study Area Species Exclosure Open Area WRB-North Aster spp. 1.4AB' m..." (0.8) Boot’s wood fern 0° --- W 12921111 (0) Canada mayflower 4.5° ...... Maianthsmum canadenss (0-4) Crested wood fern ------ 0.5° Donated: enslata (0.5) autumn: 599- 0c mm (0) Dwarf enchanter’s nightshade ----- 0° Clmaaa albina (0) Fringed polygala 0° 0° Palxgaia nauaiibiia (0) (0) Goldthread 2.7AB 6.1AB Camisgmsnianslisa (1.4) (0.4) Grass-sedge 0c -..-.. (0) Hawkweed ------ 0.6AB Biennium 599- (0-3) Jack-in-the-pulpit -....- 0° Adm SPP- (0) Long beech fern ----- 0° maimed: nalusuls (0) Rattlesnake fern 0.6A .....- mum xlrginianum (0-3) Rough bedstraw 0A 0° aalium asnteiium (0) (0) 120 Table A15 (cont’d). Site Study Area Species Exclosure Open Area WRB-North Snowberry 50° 1 .5° (cont’d) fiaultheria hlsnidula (0-7) (09) Sweet coltsfoot 0.6° Emails: naimatu: (0.3) ...... Sweet-scented bedstraw 4.2° .....- Gailumttlflbnlm (0.1) Three-leaved Solomon’s seal 50° 42° Smiiaalnatrifnlia (1.2) (1.1) Wood anemone ----- 0.2° Amman: qulnauefilila (0.2) WRB-South Aster SPP- 0.9AB ...--- (0) Boot’s wood fern 0 ...... WM (0) Canada mayflower 5.1 -..--- Maianthamumranadanss (0.6) Crested wood fern mm 0 WW (0) Domitian: SW 0.3 ------ (0.3) Dwarf enchanter’s nightshade ------ 0 Cltcaea alpine (0) Fringed polygala 5.6 5.5 Ebixaaianauaimlia (1.2) (0.5) Goldthread 7.0A 6.4A finalisamsniandiba (0.6) (0.8) Grass-sedge 0 ------ (0) Hawkweed ----- 0A Biennium spp. (0) Table A15 (cont’d). 121 Site Study Area Species Exclosure Open Area WRB-South Jack-in-the-pulpit --- l . 1 (cont’d) Arisaema Spp. (1.1) Long beech fern mm 0.3 Ihahmtensnaiustris (0.3) Rattlesnake fern 1.2AB -....- Banshiummlnlanum (0.3) Rough bedstraw 3.9AB 2.6AB fiailum aspneiium (0.9) (0.1) Snowberry 5.6 4.4 aauitheriahisniduia (1.3) (1.4) Sweet coltsfoot 0 ..---- Bataaitesnaimanl: (0) Sweet-scented bedstraw 0.3 -..--- fiailurntn'flnnlm (0.3) Three-leaved Solomon’s seal 2.8 1.4 Smilablna nlfbila (0.5) (1.0) Wood anemone mm 0 Anemoneauinauafaila (0) SP-North Aster Spp. 2.8A mm (1.0) Boot’s wood fern 3.1 ------ Wadsworth (1.2) Canada mayflower 7.1 ------ Maianthelnumcanadense (1.0) Crested wood fern mm 0 Manama (0) Wanted: spp. 2 0 ----- (1:1) Table A15 (cont’d). 122 Site Study Area Species Exclosure Open Area SP-North Dwarf enchanter’s nightshade --- 1.3 (cont’d.) Cimeaalnina (1.3) Fringed polygala 0 0 WW (0) (0) Goldthread OB OB Contismnlandiea (0) (0) Grass-sedge 5.6 mm (0.4) Hawkweed mm 6.73 Mum spp. (3 .2) Jack-in-the-pulpit ---- 0 Adam SPP- (0) Long beech fern mm 0 Med: 231mm: (0) Rattlesnake fem 4.38 mm mm (0-7) Rough bedstraw 2.2AB 3.4AB fialium asmellum (1 .3) (1.2) Snowberry 0 0 Qaulthnia hisnidula (0) (0) Sweet coltsfoot 3.8 mm Eetasitesnalmam (2.0) Sweet-scented bedstraw 0 mm Qaliumttiflnmm (0) Three-leaved Solomon’s seal 0 0 Smilacinamfelia (0) (0) Wood anemone mm 2.7 Anemoneauinnuemlia (0.9) Table A15 (cont’d). 123 Site Study Area Species Exclosure Open Area SP-South Aster spp. OB --- (0) Boot’s wood fern 1.2 ...... W5 Milli (1.0) Canada mayflower 6.8 ....-- Maianthmnumm (0-3) Crested wood fern ---- 1.8 W Slim (0.8) W spp. 0 ...-.. (0) Dwarf enchanter’s nightshade mm 4.5 denina (0.9) Fringed polygala 0 0.4 Bulimia musiflzlia (0) (0-4) Goldthread 3.5AB 4.4AB 9.91315 mnlandiea (0.9) (0.6) Grass-sedge 1 ,8 ...--- (1.5) Hawkweed ----- OAB Himdum Spp. (0) Jack-in-the-pulpit mm 2.3 Arisaema Spp. (12) Long beech fern mm 2.7 111mm nalustnis (0-8) Rattlesnake fern 4.33 mm B l . . . . (0.7) Rough bedstraw 2.2AB 4.9AB fialium amellum (1 .3) (1 .0) Snowberry 0 0 Gaultherin hisnidula (0) (0) 124 Table A15 (cont’d). Site Study Area Species Exclosure Open Area SP-South Sweet coltsfoot 3.8 ---- (cont’d) Eetasitesnaimams (2.0) Sweet-scented bedstraw 0 ---- Galiumtn'flnmm (0) Three-leaved Solomon’s seal 0 0 Smilasina Ritalin (0) (0) Wood anemone mm 0 Amen: WW (0) 'Means with different letters within species and site were significantly different (P<0. 10) among study areas with the Kruskal-Wallis one-way analysis of variance (Siegel and Castellan 1988) using SYSTAT (1992) and the Kruskal-Wallis multiple comparison test (Siegel and Castellan 1988). "Species not identified in any of the study areas for this site type. cKruskal-Wallis multiple comparison test (Siegel and Castellan 1988) was unable to detect where the significant difference occurred within species and site among the study areas as detected by the Kruskal- Wallis one-way analysis of variance (Siegel and Castellan 1988) using SYSTAT (1992). LITERATURE CITED LITERATURE CITED Albert, D.A., S.R. Denton, and B.V. Barnes. 1986. 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