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S. degree in Wildlife Ecology Major professor 0-7639 MS U i: an Affirmative Action/Equal Opportunity Institution 2 ”Name? 5mm" Michigan 3’3” l University PLACE iN RETURN BOX to remove this checkout from your record. TO AVOID FINES return on or before date due. DATE DUE DATE DUE DATE DUE l ii? l MSU Is An Affirmative Action/Equal Opportunity institution chS-nt WILDLIFE RESPONSE TO CONVERSION OF HARDWOOD FOREST TO RED PINE by Suzanne Tomassi 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 1991 ABSTRACT WILDLIFE RESPONSE TO CONVERSION OF HARDWOOD FOREST TO RED PINE by Suzanne Tomassi Small mammal, bird, and deer populations were compared between 5 red.pine plantations converted from clearcuts and 5 naturally regenerating clearcuts. The conversion process included mechanical and chemical site preparation. Measured vegetation variables differed significantly between the converted sites and controls in all years. Total small mammal abundance increased with treatment and increases were negatively correlated with vegetative abundance and structural complexity. Bird density, species richness, and species diversity were lower on treated.plots than controls. Changes in bird populations were correlated with decreasing vegetative complexity. No significant changes in deer use were observed after treatment. Control sites provided potentially better summer deer habitat than converted sites. Results suggest that the conversion process may cause undesirable shifts in some wildlife populations, particularly bird species, and that the preservation of a hardwood. component in converted plantations might improve the suitability of these areas for wildlife. ACKNOWLEDGEMENTS This project was funded by Mead Paper Company, Escanaba, Michigan. I would like to thank.my major professor, Dr. Jonathan B. Haufler, for his advice and support throughout my graduate studies. Appreciation is also extended to my other committee members, Dr. Henry Campa III and Dr. Kurt Pregitzery for their input and suggestions. The expert statistical advice of Dr. Scott Winterstein was also greatly appreciated. My sincere thanks go to Shari Dann for providing employment and many learning opportunities over the past 2 years. Thanks are due to Tony Lazzari and John Johnson of Mead Woodlands Division for their help and the use of the Mead facilities during my field season. Appreciation also goes to Jane Brand and all the interns and volunteers who helped with fieldwork during the course of this study. My fellow graduate students contributed.more to this work than I can begin to list. Many thanks are extended to them for their advice, suggestions, and friendship. Finally, I would like to thank my parents for their unwavering support and confidence in me. ii TABLE OF CONTENTS Rage LIST OF TABLESOOO0.0000.0.0.0...O......OOOOOOOOOOOOOOOOOC iv LIST OF FIGURES..........................................Vii INTRODUCTION............................................. 1 OBJECTIVES............................................... 8 STUDY AREA DESCRIPTION................................... 9 METHODS.................................................. 16 VEGETATIVE SAMPLING................................. 16 WILDLIFE RESPONSES.................................. 19 Small Mammal Census............................ 19 Bird Census.................................... 20 Deer Census.................................... 21 DATA ANALYSIS....................................... 22 RESULTSOO......OCIOOOOOOCOCO......OOOOOOOOOOOOOOO00...... 24 VEGETATIVE RESPONSES................................ 24 Vegetative Frequencies......................... 24 Woody Densities................................ 47 Cover and PHD.................................. 56 WILDLIFE RESPONSES.................................. 56 Small Mammals.................................. S6 BirdSO0.0I.0.0...O...O.......OOOOOOOOOOOOOOOOOO 66 DeerOOOOOOOO.........OOOOCOOOOOC......OOOOO.... 72 DISCUSSION.................OOOOOOOOOOOOOOO.......OOOOOOOO 73 VEGETATIVE RESPONSES O O O O O I O O O O O O I O I O C O O O O O O O O O O O O O O O 73 WILDLIFE “SPONSES O O O O O O O O O O O O O O O O O O O O O O O O O O O I O I O O O O 77 small mammals. O O O O O O O O O O O O O O I O O O O I O O O O O O O O O O O O O 77 Birds.......OOOOOOOOOOOOOO......OOOOOOOOOOOOOOO 80 DeerOOOCOOO......OOOOOOOCOOO0.000.000.00000.... 87 DEER SUMMER HABITAT SUITABILITY..................... 89 SMARY AND RECOMNDATIONS O I O O O O O O O O O O O O O O O O O O O O O O O O O O O C 9 3 LITERATURE CITEDOOOO0.0.IO............OOOOOOIOOOOOOOO.... 97 APPENDIXOOOOOOOOO......OOOOOOOOOOCO......00.000.000.00000106 iii LIST OF TABLES Treatment histories of converted red pine plantations and naturally regenerating clearcuts in the Upper Peninsula, Michigan..................................... 17 Absolute (AF) and relative frequencies (RF) of common herbaceous species occurring on converted red pine plantations (treatments) and clearcuts (controls) in the Upper Peninsula, Michigan, in 1986....... 25 Absolute (AF) and relative frequencies (RF) of woody species <1m in height occurring on converted red pine plantations (treatments) and clearcuts (controls) in the Upper Peninsula, Michigan, in 1986....... 28 Absolute (AF) and relative frequencies (RF) of common herbaceous species occurring on converted red pine plantations (treatments) and clearcuts (controls) in the Upper Peninsula, Michigan, in 1987....... 30 Absolute (AF) and relative frequencies (RF) of woody species <1m in height occurring on converted red pine plantations (treatments) and clearcuts (controls) in the Upper Peninsula, Michigan, in 1987....... 32 Absolute (AF) and relative frequencies (RF) of common herbaceous species occurring on converted red pine plantations (treatments) and clearcuts (controls) in the Upper Peninsula, Michigan, in 1988....... 35 Absolute (AF) and relative frequencies (RF) of woody species <1m in height occurring on converted red pine plantations (treatments) and clearcuts (controls) in the Upper Peninsula, Michigan, in 1988....... 39 Absolute (AF) and relative frequencies (RF) of common herbaceous species occurring on converted red pine plantations (treatments) and clearcuts (controls) in the Upper Peninsula, Michigan, in 1990....... 43 iv 10 11 12 13 14 15 16 Absolute (AF) and relative frequencies (RF) of woody species <1m in height occurring on converted red pine plantations (treatments) and clearcuts (controls) in the Upper Peninsula, Michigan, in 1990....... Mean densities of woody species >lm in height occurring on converted red pine plantations (treatments) and clearcuts (controls) in the Upper Peninsula, Michigan, in 1986, 987, 1988, and 1990....... Absolute (AD) and relative densities (RD) of woody species >1m in height occurring on converted red pine plantations (treatments) and clearcuts (controls) in the Upper Peninsula, Michigan, in 1988....... Absolute (AD) and relative densities (RD) of woody species >lm in height occurring on converted red pine plantations (treatments) and clearcuts (controls) in the Upper Peninsula, Michigan, in 1990....... Percent cover of vegetation in 3 height strata on converted red pine plantations (treatments) and clearcuts (controls) in the Upper Peninsula, Michigan, in 1986, 1987, 1988, and 1990......................... Foliage height diversity (FHD) values for converted red pine plantations (treatments) and clearcuts (controls) in the Upper Peninsula, Michigan, in 1986, 1987, 1988, and 1990......................... Mean (SE) small mammal abundance (recaptures not included), species richness, and species diversity on converted red pine plantations (treatments) and clearcuts (controls) in the Upper Peninsula, Michigan, in July and August of 1987......... Mean (SE) small mammal abundance (recaptures not included), species richness, and species diversity on converted red pine plantations (treatments) and clearcuts (controls) in the Upper Peninsula, Michigan, in July and August of 1988......... 48 51 52 54 57 58 6O 61 17 18 19 Mean (SE) small mammal abundance (recaptures not included), species richness, and species diversity on converted red pine plantations (treatments) and clearcuts ‘(controls) in the Upper Peninsula, Michigan, in July and August of 1990......... 64 Bird density, species richness, and species diversity on converted red pine plantations (treatments) and clearcuts (controls) in the Upper Peninsula, Michigan, in 1987............................ 67 Bird density, species richness, and species diversity on converted red pine plantations (treatments) and clearcuts (controls) in the Upper Peninsula, Michigan, in 1990............................ 70 Vegetative species found on converted red plantations and clearcuts in the Upper Peninsula, Michigan, in 1986, 1987, 1988, and 1990...............................106 Correlation coefficients (r;) for statistically significant associations between small mammal population variables and vegetative characteristics on converted red pine plantation and clearcuts in the Upper Peninsula, Michigan, in 1987, 1988, and 1990...............................115 Bird species identified on converted red pine plantations and clearcuts in the Upper Peninsula, Michigan, in 1987 and 1990.........................................117 Correlation coefficients (r;) for statistically significant associations between bird population variables and vegetative characteristics on converted red pine plantations and clearcuts in the Upper Peninsula, Michigan, in 1990.......120 vi LIST OF FIGURES EQQE Location of the study area in Menominee County, MiChiganOO0.0..........OOOOOOOOOOOOOOOO0.0... 10 Mean monthly long term temperature and mean monthly temperature during the study period for Stephenson, Michigan.............. 13 Mean monthly long term precipitation and mean monthly precipitation for the study period for Stephenson, Michigan.............. 15 vii INTRODUCTION Red pine (Rings :esingga) plantations and natural stands comprise about 567,000 hectares of commercial forest in the Lake States (Dickmann et a1. 1987). The species historically made up about 1/3 of all pine forests in this area (Benzie 1977). Red pine is highly valued as a timber species because of its straight and rapid growth, adaptability to a range of soil types (Eyre and Zehngraff 1948), response to thinning (Dickmann et a1. 1987), and relatively high resistance to disease and insects (Benzie 1977). It transplants easily, is successful in commercial plantations, and is the most intensely managed conifer in the Lake States (Capen 1979). Red pine, in natural stands, is commonly associated with jack pine (2- We) . quakinq aSPen (Bangles mileage). paper birch (fistula W), and scrub oak (211915.25 iligifglig) on drier sites, and additionally with white pine (P. W), red maple (Age: W), northern red oak (gm mam), balsam fir (5219.5 balsam), white spruce (gig-g; glance), black spruce (B. W), black cherry (Ezugug ggzgtina), and chestnut oak (Q. prinug) on moister sites (Fowells 1965, Benzie 1977). On loamy soils, American basswood.(1111a3amezigana), yellow birch.(fi. lQLQQ). American beech (Legals grandifelia) . eastern hemlock (1151.13: W) . 1 2 white ash (frames antigens). red spruce (2- aliens). northern white-cedar (Tania occidentalig), and eastern hop hornbeam. (Q§;11g_ gigginiana) are also common associates (Fowells 1965). Red pine also grows naturally in relatively pure stands in the Lake States (Fowells 1965, Benzie 1977). Because the species is long-lived, it often remains dominant after other shade intolerant associates have died. It is in turn replaced by more tolerant associates as succession proceeds (Benzie 1977). Associated understory species typically include Canada blueberry (W W) , lowbush blueberry (y. W) , sweetfern (mi; peregrine), American hazel (Q. cornuta), bearberry (W era-41ml). prairie willow (Felix mute). striped maple (A. pgngylyaniggm), dwarf bush-honeysuckle (Digznilla lgnigera), Jerseytea ceanothus (Qeangtnns amgziggnus), sand cherry (Eznngg 93311;), and American fly honeysuckle (Lgniggrg canadensis) (Fowells 1965), as well as most common overstory associates except jack pine. Unthinned mature red pine stands are typically closed- canopy, and the amount of understory is inversely related to the density of the overstory (Dickmann et a1. 1987). Total area of ground cover by woody vegetation less than 1.8m in height was only 4.2% in a mature red pine stand in Michigan (Gysel 1966). Tappeiner and Alm (1975) and Dickmann et a1. (1987) also noted a lack of understory vegetation in red pine stands. \ u -_/ Because of red pine's intolerance to shade, the species 3 responds best to even-aged management methods. Dense stocking (up to 2000 trees/ha) of red pine in plantations is recommended for maximum timber production (Eyre and Zehngraff 1948, Lundgren 1983), and intensive site preparation is often necessary to reduce hardwood competition. To increase softwood production, foresters have converted hardwood sites to red pine. Site preparation may involve the application of herbicides to reduce hardwood competition. Slay et al. (1987) compared 4 types of site preparation (chop and burn, windrow, fuelwood harvest, and fuelwood harvest plus herbicide) and reported higher seedling growth rates, greater groundline diameters, greater volume, and lower biomass of competing species on herbicided plots. Loblolly pines (2. taggg) on sites treated with hexazinone for 2 years following planting averaged 1.5 to 2.0 times greater in height than trees on untreated sites after 4 growing seasons. In addition, trees on treated plots were 2cm to 5cm greater in groundline diameter and up to 7 times greater in volume than those on untreated plots (Knowe et al. 1985). Creighton et al. (1987) observed similar height and diameter responses for loblolly, longleaf (B. augtzglis) and slash pine (P. elligttii) plots treated with hexazinone and other herbicides for 1 year, and they reported additional height and diameter increases on treated plots after a second year of herbicide application. They also observed a significant increase in pine survival after 1 year of treatment. Nelson et al. (1981) demonstrated a positive response in loblolly growth in plots treated with 4 hexazinone during the first 3 growing seasons and attributed the increase to a reduction in competition for water by weeds. McKee and Wilhite (1988) estimated a rotation length decrease of 1 year in loblolly pine as a result of herbicide treatment. Loblolly plantations treated with hexazinone in the first growing season showed a decrease in herbaceous vegetation after that growing season, but differences between treatment and.control plots disappeared.by the end of the second growing season (Blake et al. 1987). Michael (1985) compared herbicides and recommended the use of hexazinone on soils with high organic matter content. Results of studies investigating forage and cover changes associated with conversion practices suggest that the cultivation of pine monocultures with the sole objective of maximizing timber'production would adversely affect.wildlife. In addition to decreasing vegetative structural complexity and diversity, conversion to red pine alters natural succession and eliminates snags (Meslow 1978) . Planting essentially eliminates the grass-forb sere. It is widely accepted that certain wildlife species are characteristic of different stages of succession. Clearly, truncation of the successional process could negatively impact species dependent on those seres eliminated or shortened. Due to these vegetative changes and the resulting lack of wildlife food and cover species, red pine plantations are generally considered poor wildlife habitat (Benzie 1977, Johnson 1987). A red pine plantation in Michigan supported 5 fewer bird species and individuals than surrounding natural pin oak (W Wis) and jack pine communities (Gysel 1966). The plantation also supported significantly fewer white-tailed deer (M11932 11W) than surrounding communities, and small mammal numbers decreased with decreasing cover. Hurst and Warren (1982) reported reduction of deer forage, elimination of hard and soft mast, and a decrease in cover on a converted pine plantation. Forage reduction has been shown to be more extreme when herbicides are used, when compared to plantations prepared by mechanical methods alone (McComb and Hurst 1987). Some wildlife species may temporarily benefit from site preparation with herbicide. Herbicides improved mourning dove (291111.85 mm) feeding grounds on a Mississippi loblolly plantation by decreasing ground cover and increasing wooly croton (grater: W) and poke weed (211mm amalgam.) abundance (Blake et al. 1987), and vegetation changes following herbicide treatment in Oregon resulted in temporary increases in some seed eaters (Morrison and Meslow 1983) . Conversion without the use of herbicides has been shown to improve habitat for some species by increasing low growth (Felix et al. 1986). This method has also resulted in increased deer forage, when compared to conversion with herbicides (McComb and Hurst 1987). The benefits derived from populations of wildlife species emphasize the importance of considering wildlife when managing forested land for timber production. Wildlife in forest 6 ecosystems has both consumptive and nonconsumptive uses, as well as ecological value. Birds and small mammals are instrumental in seed dispersal and reduction of populations of some detrimental insects (Bruns 1959, Sloan and Coppel 1968, Shugart et al. 1975, Chew 1978, West 1968, Ream and Gruell 1980, Crawford et al. 1983). Bird species sensitive to habitat disturbance are early indicators of forest degradation. Small mammals are an important prey base (Chew 1978, Potter 1978), and they have been shown to aid aeration of soil and movement of organic matter into the soil (Chew 1978, Ream and Gruell 1980). Chew (1978) also suggested that the cutting of runways and feeding of small mammals increase fertility and rate of nutrient cycling in soil. Deer are the most popular game species in the United States (U.S. Fish and Wildlife Service 1987) and are the main prey of the timber wolf (gagis lupus) in the Great Lakes Region (Rogers et al. 1981). Wildlife species are valuable for nonconsumptive recreation as well. A total of 64 million people spent $14.3 billion on nonconsumptive wildlife related activities in 1985 (U.S. Fish and Wildlife Service 1987). Birds, small mammals, and deer, in addition to being of demonstrated importance in forest ecosystems, are generally present in relatively large numbers in Michigan forested land. Thus, these animals were selected as indicators of habitat quality in this study. An increasing demand is being placed on both timber and wildlife resources, but present silvicultural practices are often incompatible with the needs of wildlife. The effects of 7 converting hardwood sites to red pine need to be documented for wildlife to be given proper consideration in forest management practices. OBJECTIVES The overall objective of this study was to evaluate bird, small mammal, and white-tailed deer responses to conversion of hardwood sites to red pine. Specifically, the study evaluated: 1. Differences in. vegetative structure and composition between converted red pine plots and naturally regenerating clearcuts. 2. The effects of conversion to red pine on small mammal populations. 3. Differences in species composition and relative densities of bird populations between conwaied red pine plots and clearcuts. 4. The effects of conversion to red pine on deer use . 5. The relative suitability of converted red pine plantations as deer summer habitat. STUDY AREA DESCRIPTION The study was conducted in the northwestern part of Menominee County in the Menominee hill land region on Michigan's central Upper Peninsula (Fig. 1). The physiographic region is a rolling ground moraine consisting of ridges, drumlins, moderate slopes, and outwash plains (Schwenner 1989). Numerous wet areas and ridges with slopes of up to 25% occur in the region. The study area is drained by the Big Cedar River, which flows southeasterly and empties into Green Bay. Soil types on the study area include deep, well drained, moderately permeable Onaway and Nadeau fine sandy loans and soils of the Lupton, Tawas, Cathro, Deford, and Ensley series. The Lupton, Tawas, and Cathro series occur on moraines and outwash plains and consist of deep, very poorly to poorly drained, moderately permeable loamy to sandy soils. The Deford series consists of deep, poorly drained, rapidly permeable fine sandy soils. The Ensley series soils are similar to those of the Deford series, except that permeability may be somewhat slower. All soil types occur on 0% to 2% slopes with the exception of Onaway and Nadeau fine sandy loams, which may occur on slopes of up to 12% (Schwenner 1989). 10 Study Site Figure 1. Location of the study area in Menominee County, Michigan. 11 The climate of the region is semi-continental, with the prevailing wind from the southwest. The mean annual temperature of the area is 5.8°C, ranging from a high of 19.7°C in July to a low of -9.9°C in January. Annual precipitation averages 82.9cm, with 65.6% of the total occurring in April through September. Mean annual snowfall is 165.9cm. Mean monthly temperatures during the study period are shown in Figure 2 (NCAA 1986, 1987, 1988, 1989, 1990). Mean monthly precipitation during the study period is shown in Figure 3 (NCAA 1986, 1987, 1988, 1989, 1990). Due to the treatment effects, treated sites were sparsely vegetated. Planted red pine predominated, and other woody species included balsam fir, balsam poplar (W W) , paper birch, cherry (Plums spp.) , speckled alder (Linus zuggsa), quaking aspen, white spruce, and less commonly, bigtooth aspen (W W, dogwood (9.9mm app-i. red maple. sugar maple (Ace: W). serviceberry (W spp.), and willow (53111;) . Quaking aspen predominated on the control plots. Other tree species present on controls sites were most of those species found on treated sites. Understory vegetation included most overstory species plus brambles (BM spp.) , blueberry (mm spp.) , elderberry (W spp.) , currant (Bibs; spp.) , grape (11.11.: app-) . Virginia creeper (W Mansions) . and Viburnum (11m spp.) . All control plots and most treatment plots had thick ground cover composed of numerous herbaceous species. 12 Figure 2. Mean monthly long term temperature and mean monthly temperatures during the study period for Stephenson, Michigan. 13 Month 198" l l O O O N w-t 30 (3°) eJnieJeduie; Kluiuoui ueew 1989 1990 1988 Long term norm 1986 14 Figure 3. Mean monthly long term precipitation and mean monthly precipitation during the study period for Stephenson, Michigan. 15 0mm: mama mama bwmfi wwmfi 8.8: Show 984 £82 om (we) uoneudioeid Mmuow ueew METHODS Ten study plots of 8 ha each were established in newly clearcut hardwood stands on the study area. Plots were selected based on their similarity in vegetative composition and date of clearcutting. Plots were of irregular shape. Hexazinone, in the herbicide Velpar, was applied to all treatment sites. All sites had previous mechanical manipulations (Table l). VEGETATIVE SAMPLING Vertical cover was measured on all plots by the line intercept.method (Canfield 1941). Percent cover was measured in 3 height strata: <1m, 1-7m, and >7m. Vegetation covering a measuring tape was recorded in centimeters with gaps of <5cm ignored. Transects were located randomly and run in the same direction. In 1986, 1987, and 1988, numbers and lengths of transects were varied to meet efficiency and sample size requirements. Due to the wide variation in vegetative cover within sites in 1990, a maximum of 30 transects 30m in length was located on each study site. In 1986, cover was not measured in 3 unplanted treatment plots because of the nearly total lack of vegetation on these sites. In all years, a 5m- 16 Table 1. 17 Treatment histories of converted red pine plantations and naturally regenerating clearcuts in the Upper Peninsula, Michigan. SITE NAME (SITE TYPE) DATE PLANTED MECHANICAL AND CHEMICAL TREATMENT HISTORY Broken Pine (Treatment) Camp 7 (Treatment) Sand Road (Treatment) Cut Across (Treatment) Indian Head (Treatment) Burnt Stump (Control) Slessinger (Control) North Ridge (Control) 9/85 9/83 9/86 9/86 9/86 Roundwood harvest 1/84 Brushraked 12/84 Velpar treated 6/85 (5 qts/ac.) Disc trenched 6/85 Velpar treated 5/87 (4.2 qts/ac.) Velpar treated 5/88 (4.3 qts/ac.) Roundwood harvest 1/82 Brushraked, burned 1/82 Disc trenched 8/83 Velpar treated 8/83 (5-6 qts/ac.) Velpar treated 6/85 (3 qts/ac.) Velpar treated 5/86 (4 qts/ac.) Velpar treated 5/88 (4.4 qts/ac.) Roundwood harvest 1/85 Brushraked 5/86 Velpar treated 5/86 (5.2 qts/ac.) Velpar treated 6/86 (4.7 qts/ac.) Velpar treated 5/89 (4.2 qts/ac.) Whole tree harvest 1/85 Disc trenched 6/86 Velpar treated 6/86 (3.2 qts/ac.) Velpar treated 7/86 (5.1 qts/ac.) Velpar treated 5/89 (4 qts/ac.) Whole tree harvest 12/84 Velpar treated 5/86 (5.9 qts/ac.) Disc trenched 6/86 Velpar treated 6/86 (4 qts/ac.) Velpar treated 5/88 (4.2 qts/ac.) Roundwood harvest 1/83 Roundwood harvest 12/82 Roundwood harvest 12/82 Table 1 (cont'd). 18 MECHANICAL AND CHEMICAL TREATMENT HISTORY SITE TYPE DATE (SITE TYPE) PLANTED Cut Across (Control) Railroad Grade (Control) Whole tree harvest 1/85 Roundwood harvest 1/84 19 10m buffer at plot edges was left unsampled to minimize edge effects. Randomly located nested plots were used to determine density of woody species and frequency of woody and herbaceous species. Density of woody vegetation >1m in height was measured in randomly located 2m x 30m plots. The number of plots varied with sample size requirements. Frequency of woody species <1m in height was recorded in 2m x 10m plots, and frequency of herbaceous vegetation was recorded in 2m x 5m plots, except in 1987, when 1m x 1m plots were used. In 1986, 1987, and 1988, frequency was measured in 20 plots, and in 1990, 30 plots were located on each study plot. Grasses, brambles, and some tree species were recorded by genera, while forbs and most trees and shrubs were recorded by species. WILDLIFE RESPONSES Small Mammal Census Small mammals were censused by live trapping twice annually, in July and August of 1987, 1988, and 1990. A 6 x 6 trapping grid with 2 Sherman live-traps (H.B. Sherman Co., Tallahassee, Fla.) per station and trap spacing of 15m was established at the center of each study plot. Traps were placed next to logs and in other small mammal travel lanes and covered with vegetation. Bait was a mixture of oats, beef fat, and anise extract. Each treatment plot was randomly paired with a control, and pairs were trapped on the same 20 nights. Trapping was conducted in 2 groups (1 group of 6 plots and 1 group of 4 plots). Each group was trapped for 5 consecutive days in each month. Newly captured animals were ear tagged, and species, tag number, and grid location were recorded. Bird Census Bird populations were censused from mid-May to mid-June, 1987 and 1990, using the variable circular-plot method (Reynolds et al. 1980). Four stations were established on each plot and marked before censusing. Stations were placed so that there was at least a 10m buffer between circular plots, in order to avoid double counting. A 10m buffer was also left between the circular plots and study plot borders to minimize edge effects. Each treated site was randomly paired with a control, and each of 2 observers censused 1 pair each morning. Each observer censused all plots an equal number of times. The order in which the plots, and stations within plots, were censused was rotated to eliminate biases caused by changes in bird activity throughout the census period. Censuses began at sunrise, when breeding bird detectability is greatest (Kendeigh 1944, Hall 1964, Jarvinen et al. 1977, 1978, Connor and Dickson 1980). Censusing periods were 10 minutes in duration, and 1 minute was allowed before each census for birds to resume normal behavior. Birds were not censused in rain, wind, fog, and unseasonable cold, since these conditions may alter bird behavior and detectability 21 (Robbins 1981). In 1987, 3 treatment plots and their controls were censused 4 times each. The radius of each sampling station was 50m. In 1990, all 5 treatment plots and their controls were censused 8 times each and the sampling radius was reduced to 25m to increase sighting efficiency and identification accuracy. Different observers conducted the censuses in each year. Singing males were recorded separately from all other observations. The number of singing males was doubled and compared to the number of males plus all other observations and found to be less. Therefore, all observations were used in the final analysis. Bird density, species richness, and species diversity were calculated. Deer Census Deer response to conversion was assessed using track counts in 1986, 1987, and 1988 and pellet counts in 1990. Plots usable for track counts were limited by the presence of suitable road surfaces Sample sizes in 1986 and 1987 were too small for analysis of results. In July 1988, counts were conducted on 3 treatment and 3 control plots. A 0.15km-0.30km section of road bordering each sampled plot was dragged in the evening and checked in the morning for tracks. Five replications were conducted and results standardized to tracks/0.20km of road. In August 1990, pellet-group counts were conducted on all study plots. Four 10m x 10m plots were located randomly on 22 each plot and cleared of all pellets. After 5 days, plots were checked for pellet groups. Five or more pellets in a line or cluster were considered a group. All counts were made by the same observer to eliminate observer bias (Neff 1968). Results were expressed as pellet groups/ha and used for relative comparisons between treatments and controls. DATA ANALYSIS A completely randomized design was used for this study. The linear model for this design is: YU‘=”'+‘i+‘% mean of all observations 1 u t = variability due to treatments a, variability due to errors. Statistically adequate sample sizes for vegetative sampling were determined using Freese's (1978) sample size formula: n = ti’sz/E2 t = tabulated t value at the 90% confidence limit 52 = sample variance E = allowable error (mean x 0.10). 23 A maximum of 30 samples per plot were taken for any measurement when required sample sizes were extremely large, as they were for the upper stratum of vegetative cover. The nonparametric Mann-Whitney U test (Seigel 1956) was used to test density of woody vegetation, frequency of woody vegetation, frequency of herbaceous vegetation, percent cover, foliage height diversity, bird density, bird species richness, bird species diversity, small mammal abundance, small mammal species richness, small mammal species diversity, and deer use. The Spearman Rank Correlation Coefficient was used to describe associations between vegetative responses and bird, small mammal, and deer responses. Foliage height diversity, bird species diversity, and small mammal species diversity were determined using the Shannon-Weiner diversity index (Hair 1980): H. = -2 Pi log Pi Pi = decimal fraction of total individuals or total cover of the ith category. The numbers of individual small mammals of each species captured for the first time on each plot were used for comparison in all trapping periods. RESULTS VEGETATIVE RESPONSES Vegetative Frequencies All vegetative species found on the study plots are listed in the Appendix (Table A-1) . 0f the 67 herbaceous species identified in 1986, 32 occurred on treated sites and 63 on controls. Absolute and relative frequencies of 19 common (>5% absolute frequency) herbaceous species are listed in Table 2. Of these, 15 were found on both treatments and controls. Bedstraw (m spp.) , Canada mayflower (unionization caan—de se). moss (M. 98mm). and spreading dogbane (8292mm andresaemifolilam) were found on controls only, and all common species except dandelion (W sop-i and horseweed (8:192:98 99851998919) were more abundant on controls than on treated sites. Bedstraw, moss, round-lobed hepatica (m M) , spreading dogbane, and wild strawberry (W W) were significantly higher in both absolute and relative frequency on control siteS- Cinquefoil (Martina sop-i. grass (ma sop-i. hawmeed (W spp.) , and bunchberry (Corpus ganadensis) exhibited significantly higher absolute frequencies on controls, and dandelion and violet (1121B spp.) were higher in 24 25 Am.ov n.a Ae.ev o.nH An.av s.~ Am.nv o.m Haeummuom Ao.HV m.~ .e.~av o.on Am.o. c.H .Ae.mv o.o 8883338: Am.ov n.oa .e.mv o.em Am.av «.ma .AH.~AV o.m¢ amuse Am.av m.~ .o.eav o.- Ae.ov o.a .m.mv o.m eoucmoaoo 15.Hv n.» Am.nav o.me 15.83 ~.eH .~.~Hv o.ae cums Am.ov m.~ .o.mv o.e~ gzlm.ov s.m as.ev o.~n condenses .e.ov o.H A~.mv o.oa .m.ov m.” .AH.HV o.n aqoumsvceo 8.8 o4 8.3 92 8.8 m6 3.: ed 6883.35 A~.ov e.o 16.nv 8.6 Ao.ov o.o Ao.ov o.o umaoaosma eeocmo A~.Hv e.e Am.mv o.~e .m.a. m.n _:.m.~v o.ca muumnsocam Am.ov m.~ 1o.mv o.¢~ :mo.ov o.o :lo.ov o.c annumcmm .m.ov m.m in.mv o.om .¢.~V s.m Aa.mdv o.p~ umum< Ammo mm a lame m< « laws as » Anew 84 « mqomszoo mszmzsemme mmHommm newcomm nnooooouon :oeaoo mo Ammv nowososveuu m>aueamu one Amsoo so msfiuusooo .N UHQMB “Ho.o v as maouucoo soup pamumuuae saucmosuacuam:. “mo.o v my maouucou sous ucmumuuse saucmofiuacmsm: Aofl.o v m8 maouucoo aouu ucmumuuae saunaoauacmsm. 26 As.ov m.» 16.88 o.os .:ls.o8 m.~ 2:16.~V o.m suuohseuum can: A».o. e.e «p.58 o.ne 9:1m.o8 o.» .s.vv o.m~ uoHoH> Ao.av m.n .e.HH8 o.mn .H.nv o.m 6o.HHV o.en «Humane A~.HV s.a A~.mv o.es _:Ao.ov o.o :mo.oc o.o weenmoe ocaeemuam l¢.ov e.a Ao.ev ¢.ma :la.ov H.o s:lm.o. o.a «nausea: emboaiecsom .88 8o :48 oé 2.8.8 o.o :8.8 o.o mmo: Am.ov m.o .e.nv o.e .o.~v o.v in.mav o.e~ ommsmmuom Ammv mm » lame me a Ammo mm a lame we » mqomazoo mazmzeemms manommm .18.»:088 a means 27 relative frequency on treated sites. All 16 woody species <1m.in height occurring on the study area in 1986 were present on both treatment and control sites, with the exception of blueberry, which was not found on treated sites (Table 3). All species were more common on controls, though not all differences were statistically significant. Species which differed significantly in absolute frequency were aspen, balsam fir, balsam poplar, cherry, and dogwood, and balsam fir and balsam poplar also differed in relative frequency. Seventy-five herbaceous species were found on the study plots in 1987. Of these, 62 occurred on treated sites and 54 on controls. Absolute and relative frequencies of common species are listed in Table 4. Daisy fleabane (E. engage), fern (family Polypodiaceae) , and yellow avens (gem M) were significantly more common on controls than on treatments, and daisy fleabane, grass, and yellow avens were significantly higher in relative abundance on controls. Thirteen woody species <1m in height were identified on the study plots in 1987 (Table 5). Balsam poplar and dogwood were found only on controls, and vaccinium, hop hornbeam, and northern white-cedar were found only on treatments. Aspen and balsam poplar were significantly higher in both absolute and relative abundance on controls than on treatments, and pine (red and white) was significantly higher in absolute and relative frequency on treated sites. Birch had a higher relative frequency on treated sites. "J 28 Ae.~v m.m AH.NHV c.mn Am.av s.m A~.nv o.m~ wanna ems AH.H8 e.n 1H.ev o.na .s.ov N.H Am.Hv o.n aeohcuom AH.~V e.m AH.oHV o.e~ .e.~8 n.m A~.e. o.os usaamuem Ae.av ~.n AH.mV o.mH Am.ov m.o .Ao.av o.H ooosooo Ao.Hv o.¢ An.e. o.ma .m.ov w.n A».HV o.m unmuuso an.av H.oH An.s8 o.mo Ae.~v e.» ..~.HHV o.sn anyone Aa.n8 o.m .m.nav o.mm .~.nv a.m Ao.mav o.ee house Ao.av n.q Am.ev o.o~ .Ao.H. o.H :ge.~8 o.e season ammamm Am.ov n.n A>.ev o.- :lm.ov ¢.H :le.~v o.m use aemHmm Ae.~8 H.¢H Ae.av o.mm Ae.m. s.s :lm.s8 o.en comma AH.~. n.n A¢.m8 o.- .H.n8 s.~ A¢.ev o.~a awe .¢.o8 n.H 15.H8 o.e AH.H8 o.H Am.av o.n swede Ammo mm a lame me a lame mm a lame m< » mqomszoo mazmzesoo so mswuusooo voodoo a“ adv nmwooau >6003 no “hay nowocosuouu o>fiueaeu one Andy musaonna .n manna 29 “Ho.o v my Houucoo acne ucmumuuae sauceosuacmamzz lmo.o v my Houucoo sous ucououuse saucmoaoacmam: Aoa.o v my Houusoo aouu ucououuwo aauceowuwcmamc Am.HV o.¢ Ae.mv o.a~ Am.o8 e.H AH.H8 o.n sedans 3.8 m.~ 3.8 83 8.8 ma 3.: oé museum when: Ammv mm a lame ma » .mmc mm » .mmv me « mqomezoo mszmzsamme mmnommm .18.»:088 n wanes 3O A~.H8 «.6 16.68 n.5a 18.68 H.HH AH.6V n.n~ advance Ae.6. 6.6 An.H8 6.~ .6.H. 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Forty-one of these species were present in frequencies greater than 5% (Table 6). Aster (Aster spp.) , bedstraw, fringed polygala (W M) , grass and sedge (gang); spp.) , pearly everlasting (W W), round-lobed hepatica, tall cinquefoil (B. W), thimbleweed (W W), wild columbine (661.111.69.16 W): wild lettuce (m 261163366616). wild strawberry, wood anemone (A. W) , and yellow hawkweed (a. M) were significantly more abundant on control sites, while common mullein (mm mama), horseweed, and yellow goatsbeard (W M) were higher in both absolute and relative frequency on treatments. In addition to these species, common plantain (215193339 major), moss, thistle (m spp.), and yellow goatsbeard were higher in relative abundance on treated sites. Relative frequencies of fringed polygala, orange hawkweed (H. aurgntiaggm), pearly everlasting, round-lobed hepatica, wild columbine, wild lettuce, wood anemone, and yellow hawkweed were significantly higher on controls than treatments. Thirty-two woody species occurred on the study area in 1988, all of which except for elderberry (W W) , elm (31mg spp.) , red pine, and Virginia creeper were found on the controls (Table 7). 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H.o A>.cv n.o Ao.H. m.H Am.~v >.v swans spoouofim Aa.~v a.m Ae.mv ¢.em g:.~.ov ~.o 2:A>.ov >.o amazon ammflum A~.Hv m.¢ Ao.mv o.~n .o.nv ¢.m Ah.oav o.mH awn Banana an.nv m.¢ Am.eav n.m~ an.av m.a .a.nv n.m and Aa.ov H.o Au.ov h.o AH.HV H.H .>.~V >.~ comma cmoduma< Ammv mm w Ammv ac » Aumv mm w Ammv ma » mqomazoo mazmzacmma mmHommm .mwma ca .cwmfinoaz .mazmcwcmm human 0:» :« Amaouucoov musoummao 0cm Amucmaummuuv mcoauwucmam mafia vmu umuum>coo :0 ocfiuuaooo unwwm: ca adv mmwommm >UOO3 no Ammv wwwocmavmuu 0>fiumamu can “may muzaomnd .5 wanna 40 A~.ov e.o An.av o.~ .o.ov o.c Ao.ov o.o manna Human Ae.ov >.o .H.nv n.m A¢.ov e.o an.av n.a umcaa cmaxomam Au.ov H.n An.mv o.o~ 2:.o.ov o.o Ao.ov o.o munmnmofi>umm Ao.ov o.o Ao.o. o.o 2;Ao.vv ~.Hn 2:.¢.m. >.om «can omm Am.av n.m A~.HHV «.mn Ae.av o.n Ao.vv o.oH «Home omm An.av o.md Ae.~v n.nm 2:Ao.~v o.e Aa.mv 5.2” swans ocfixuso Am.ov o.o .o.¢. o.¢ Ao.ov o.o .o.ov o.o umou musumam An.ov m.o Am.av n.n .o.ov o.o .o.ov o.o uncmonmufins :umnuuoz .m.ov v.a .o.ev n.m A~.ov ~.o A>.ov >.o maxosmamcosncmsn :uonuuoz 3.8 ~.~ 31$ ...: 2.8.3 o.o 2.8.8 o.o 53:qu no: An.ov n.o Ao.mv o.~ Ao.ov o.o “o.ov o.o maxuammmcon >ufim: A~.ov ~.o An.HV n.” A¢.ov e.o an.av n.H macaw Ao.ov o.o .o.ov o.o .m.ov o.o .o.~v o.~ Sam Ammv mm « Ammv mm » Aumv mm » Awmv m¢ » mqomszoo mazmza .~.ov n.o An.av o.~ Ao.ov o.o .o.ov o.o assuanfi> Ao.~3 o.m Av.~Hv 5.22 Azan.ov n.o A:AM.HV n.a soda“: A5.H3 m.n Ao.oav n.m~ Am.o3 «.5 An.n. o.oH magnum mugs: A5.ov 5.o A~.mv o.» Ao.ov o.o Ao.ov o.o mafia mugs: 3.3 1o 3.3 5.o 8.3 o.o 8.3 o.o finmaxmz Aa.ov ~.o Am.ov n.H Ao.o3 o.o .o.o3 o.o xomumaaa Ammv mm 5 “may m¢ » Ammv mm « Ammv ma « mqomezou mazmzaamme meommm ..c.uaouv 5 manna 42 (1.2222: 1222121212). waxberry (W new). white pine, and Viburnum occurred on controls only. Balsam poplar, brambles, currant, dogwood, hop hornbeam, quaking aspen, serviceberry, and willow were all significantly more common on control sites, and all of these species except brambles and currant also differed significantly in relative frequency. Birch had a significantly higher relative frequency on treated sites, and red pine was higher in both absolute and relative frequency on treatments. In 1990, a total of 108 herbaceous species were identified on the study plots, with 81 species occurring on treated sites and 83 species on controls. Common St. Johnswort (mm W) and lamb’ s quarters (ML: 112“!!!) were present only on treated sites, while ground cedar (mm W) , sweet coltsfoot (W palmgtus) , and trillium (m spp.) were found only on controls. Aster, bedstraw, fern, fringed polygala, fungus, grass and sedge, lichen, orange hawkweed, sweet coltsfoot, and wood anemone were significantly more common on controls than treatments, and all of these except fern, fungus, and grasses and sedges differed in relative frequency as well. Treated sites had significantly more chickweed (53931133; spp., m1}!!! spp.) , common mullein, horseweed, rough cinquefoil (B. W), and yellow goatsbeard, and all of these species, along with thistle, also had higher relative frequencies on treatments. Table 8 lists absolute and relative frequencies of common herbaceous species 43 6o.oe m.o .m.me n.m Aa.ov m.o an.ae o.e oamsoaan ommcaum .w.oe ~.5 .m.5e n.m5 .n.~v 5.m .Am.eav o.~e cums Ao.ov m.n Ae.5v 5.mn Am.ae ¢.o .m.eav n.nm coaamocmo an.ov ¢.o Am.~e 5.¢ A5.ov e.a .5.me 5.~H mamnmmau suave Ao.oe o.o .o.oe o.o :gn.ae 5.” .:AH.we n.m uuosmcnou .um cosaoo Aa.oe ~.o an.ae o.~ 2:.e.ae o.m 2:.o.5e n.mn :Huaass coaaoo Ad.oe ~.o .n.ae o.~ .A5.oe H.~ .Am.5. 5.oa oamzxofinu A~.oe o.o A¢.~e 5.e A5.oe n.H A¢.5v 5.~H umaoaumns noncmo an.av N." .m.aae n.m~ A~.oe o.o Am.He 5.» auumnsocsm Ae.oe m.o .m.me 5.5 Am.ov m.o Am.ne 5.o sesame sonam A5.oe m.~ “c.5e n.5m .zga.oe H.o .2km.oe n.H auuumomm Am.oe m.5 AH.mV o.~m .:An.av 5.n Azam.oav 5.on umum< “may mm » “may m4 » “may as a lame m4 » mqomszoo mazmzsamms mmHoumm .omma cw .smofinowz .maamcflsom Momma on» :« Amaouusoov musouooau can “magmauoouuv msowuouscaa mafia own omuum>coo so oswuusuoo mowooam msoooonuon sosaoo mo Ammv mowocmsvouu m>fiuoamu can Amu om>om32omumn Ao.oe o.o Ao.oe c.o .5.oe 5.5 Am.5e 5.5 mumuunsv u.namq A~.oc 5.5 .5.33 5.3 .¢.33 5.5 2:35.53 o.om cumsmmuom “5.53 5.o 35.53 o.5 .m.ov 3.3 .5.53 5.o3 3359mmuom A5.oe 5.o 35.53 5.5 Ao.ov o.o Ao.oe o.o unooo canouo Ao.33 5.5 Ao.oe o.oo3 A5.oe 5.o3 :;5.m. o.~5 mmmomm can mmmmmuo Am.oe 5.5 35.53 o.5~ A5.ov 3.3 .5.53 5.53 nonsmoaoo 3.3 5.3 3.3 33 3.3 5.o 2.3.3 5.5 mavens .o.oe 5.3 Ae.me o.~3 :33.oe 3.o .:A5.oe 5.o nHumaaoa ommc3um “may mm 5 “mm. mm a lame mm 5 game 54 5 mqomszoo mazmzs1m in height was significantly greater on controls than treatments in all years (P < 0.01) (Table 10). In 1988, balsam poplar and quaking aspen had significantly higher absolute and relative densities on controls than treatments (Table 11). In 1990, bigtooth aspen, cherry, balsam poplar, quaking aspen, speckled alder, white spruce, and willow had significantly higher absolute densities on controls than treatments (Table 12). In addition, quaking aspen, balsam poplar, bigtooth aspen, and white spruce had higher relative densities on controls. 48 A~.Hv o.¢ .n.aac >.~n :xm.ov >.o Azam.av n.~ noosmoa Am.oc ~.n Am.ac h.o~ .o.~c m.m Am.hc n.mH acmuuso A~.~c n.oa A».~H. >.oo A¢.Hc o.¢ 2:1H.mc >.ea xuuono A~.Hc m.ea Am.ac o.mm AH.nc n.H~ .:AH.>V n.ao moansmum Am.ov m.H Am.mc o.~a 1:1o.oc o.o Lzao.oc o.o muumnmsam .Ao.ac n.m Am.>. >.oo .Au.nc n.oa Am.oac c.cm conga A¢.ov m.c Aa.nc o.m A~.~c m.n Am.mv n.m comma nucouowm Ae.oc m.o A>.n. o.m Ao.oc c.o Ao.oc o.o ooosmmum .~.Hc m.n .m.>v o.w~ .H.Hv v.~ ..o.nc n.» nuance ammaam Am.ac ~.¢ Aw.mc n.h~ .s.~c m.¢ Am.mc «.ma gnu ammaom Ae.av n.e .m.oac s.on A~.~c n.n Am.o. >.oa baa A~.Hc o.H Am.mc >.~H Ao.oc o.o “o.oc o.o comma cmofiums< Ammo mm « Ammo o< » Ammo mm » Ammo ma « anomazoo mazmzacmma mmHowmm .omma a“ .smmwnowz .maamsfismm woman 0:» ca Amaouusoov musoummao can Amusoauomuuv mcoaumusmam mean can omuhm>soo so oswuusooo panama sfi BHv mmfiommm >600: no Ammv mmwososvouu m>wuoaou can Amdv musaomnd .m manna 49 A¢.cc n.o .H.nc n.m A>.Hc 5.” an.mc n.m «Home ummam Ao.ac m.H Am.mc s.oH A¢.oc m.o an.Hc >.~ umoflm omaxomdm A¢.oc e.~ A>.HV ".ma 21m.oc o.a .zao.av n.n auuonmoe>umm .o.oc o.o Ao.ov o.o 2:1».mc m.ma :zao.mac o.wm «and com Am.ac H.m Am.mac >.mn A«.Hc m.H .Am.nc n.m mamas omm .~.HV m.HH Am.vv o.ms Am.~c n.» Aa.uc n.5a amen» ocwxmso Ao.oc o.o «o.oc o.o A~.ov ~.o Au.oc >.o a>fi condom A~.oc ~.o .n.ac n.H Ao.oc o.o Ao.oc c.o mmou «gunman .n.ov s.o .m.ac o.¢ A~.oc ~.o .h.ov h.o maxosmmmcosunmsn cumzuuoz .n.ov >.o .n.mc h.v Ao.oc o.o Ao.oc o.o oooacouH A~.oc ~.o An.av n.H Am.ov m.o .>.~V n.~ mango A~.Hc m.~ Am.mv n.ma an.oc ¢.o .Am.oc n.a saw Am.ov o.H Am.~v p.c “o.oc m.o Am.ac n.~ auumnumoau Ammo mm » Ammo ma » Ammo mm » .mmc m4 « mqomezoo mezmzeamme mmHommm .Ao.ucooc m manna 50 Afio.o v my maouucoo scum ucmumuuflc aaucuufiuficofimza Amo.o v .3 maouusoo aouu usmumuuwo Nausmofiuasmfim: Aoa.o v my maouucou some ucmumuufio aaucuofiuwcmam. An.av o.¢ Am.mv >.o~ A~.Hc n.H 14.4. o.w seas“: Am.ac ~.n .m.~av s.¢~ Am.o. o.~ Am.~. >.o magnum mugs: An.oc m.o Am.ac n.n Ao.ov o.o Ao.ov o.o mafia opens An.ac o.n an.~ac >.- :g~.o. ~.o 1:1».oc s.o umemwuo aficwmufi> A¢.Hc m.H Am.>c n.m Am.oc H.H Am.~c n.n escusnfl> An.oc m.o Am.mv n.» Ao.oc o.o Ao.oc o.o xomumame Ammo mm a lame m4 » Ammo mm » .mmv he » mqomezoo mazmzemmme mmHommm .Ao.ucooc a means 51 Table 10. Mean densities of woody species >1m in height occurring on converted red pine plantations (treatments) and clearcuts (controls) in the Upper Peninsula, Michigan, in 1986, 1987, 1988, and 1990. YEAR TREATMENTS CONTROLS Stems/ha (SE) Stems/ha (SE) 1986 41.7 (24.3)* 8786.7 (2068.7) 1987 51.0 (39.4)* 2186.0 (641.0) 1988 326.7 (92.0)* 10556.9 (1893.8) 1990 983.0 (319.4)* 7838.2 (540.2) 'Significantly different from controls (P < 0.01) 52 Ao.oc o.o AH.HV H.H Ao.oc o.o .o.oc o.o mamas cum A¢.mc m.om A~.ocaav «.mmmm ..m.~v 5.4 .An.¢nv ~.~¢ cmomn asexuao .o.o. o.o .q.ac ~.~ Ao.oc o.o Ao.oc o.o sumacuos do: .H.oc H.o .n.nc m.mn mo.oc o.o .o.oc o.o uoosooo .H.oc H.o .m.mc m.m Am.~v m.~ Aq.mc 5.0 ucmuuao an.ac H.~ .H.H¢Hc >.ma~ .o.nc m.» .>.mc n.n~ muuoso Am.mv H.oH A~.mm¢c H.Ho> “H.5Hc H.- A~.nmmc o.mmm «manaoum AH.oc ~.o .m.sc «.ma Am.~c 0.4 .>.mc m.m bongo “6.6. «.H Au.mmc o.onH .e.nc m.n A~.6Hv >.m~ comma buoouofim .m.h. v.4a 15.nnmac w.mmH~ ..H.o. H.o .A~.~c ~.~ nuance anmaum Am.oc o.a .m.mmc m.mHH Am.mc H.m A«.oc 5.8” new anmaum Ao.ov o.o .o.oc o.o A».oc h.o AH.H. H.H comma coofiumam .mmv om » Ammo o< » Ammo om » Ammo o< » mqomazoo mazmzacmma mmHommm .mwma a“ .somw£0fiz .mHsmswsom none: 0:» ca Amaouusoov musouomao can .mucmaummuuv mafia own on Umuum>soo muuwm so onwuuzooo unvflms aw EHA moaommm wooos mo Aomv mmwuwmsmo o>wvmaou can Andy ousaomnc .HH manna 53 AHo.o v my «Houucoo scum usmumuufiu aflucmofiuacmam. “o.nmmac 6.0mmoH .Ao.~mv >.m~n qaaoa Am.oc ¢.H A~.amc n.nm Ao.ov o.o AH.HV H.H soda”: An.oc m.o .m.-. m.mo “4.6. o.oa .«.eac n.n~ 0050mm mugs: AH.oc H.o An.nac n.na Ao.oc o.o lo.oc 6.6 mean mugs: Ao.av a.” 15.00Hv ~.~>H Am.oac H.HH A¢.mac m.m~ “moan omaxummm Ao.oc o.o Ao.oc o.o Aa.ov H.o A¢.ev «.4 auumn00H>umm «o.oc o.o mo.oc o.o Am.mac H.o~ Aa.onac m.mna mafia cmm Ammo am » Ammo o4 » Ammo am a lame am « mqomazoo mazmzeamme mmHommm .Au.u:ooc Ha manna 54 Ao.~v o.mn Ao.>mmc m.~¢~m .:.o.~. e." .zgm.oac m.¢a among ocfixmso Ao.oc o.o Ao.~c ~.~ .o.o. 6.6 Ao.oc o.o saw A~.ov n.o A>.oac H.H~ .n.oc n.o 1o.ac H.H noosmoo Ao.oc o.o A~.Hc ~.~ Ao.oc o.o Ao.o. o.o unmuuso An.oc m.a .~.mmc m.>¢a .>.H. m.~ :;o.mc n.4H auuono AH.HV >.~ .o.e>c o.oma ..¢.o. n.0a .m.wnv m.eaa umansnum Ao.oc o.o Ao.ac H.H .o.o. o.o ho.oc o.o auumnusam A~.ov n.o .~.0Hc e.n~ A~.Hv m.~ Am.>~c m.mn souwm Ae.oc n.a Am.~nc H.ooa :ga.ov H.o :go.ac H.H cmomm sucouafim Aa.av h.m Am.mmac «.mmo :gm.ac >.H g:Am.¢. a.» usages Banana A¢.ov H.H .5.an m.em .H.H. n.n Am.mav n.6n 8““ sundae Ao.oc o.o .o.ac H.H Aa.oc o.H Ao.nc n.n sac Ammo om » Ammo ac » Ammo am » Ammo as » mqomazoo mazmzasoo so mswuusouo usmwoz cw BAA moaomam mooo3 no “any mofiuwmsmo 0>wumamu can «adv opsHomn< .ma manna 55 adooo V my WHOHHCOU EON“ UQQHTMHHU *HUCMUHNfiCDfimga “moo v .3 odouusoo Bouu usoumuuwo hausmowuwsvwm: Aoa.o v my «Houucoo some unmumuueu aducaofiuficmflm. A~.ocmc «.mmms 6:34.6an o.mmm nHc e.nm :mm.nc 6.5 .An.oac n.~v magnum «was: AH.oc H.o Am.nv w.m .o.ov o.o .o.ov o.o mead mugs: Ao.ov o.o “o.oc o.o Ao.oc H.o Ao.av H.H aacusnfi> Aa.oc H.o Am.¢v 5.6 Aa.oc H.o Ao.~c ~.~ 90600869 Ae.ov H.H Am.~nv m.em Am.o. 5.6 .Ao.oc 6.6 umoau coaxomam Ao.oc o.o Ao.~c ~.~ An.oc n.o Ao.ac H.H muumnmofi>umm Ao.oc o.o Ao.oc o.o :mn.nav 0.5m zlm.~m~c o.omo mafia cum Ao.ov o.o Ao.~c ~.~ A~.ov n.o .o.Hc H.H mamas 00m Ammo om » Ammo o4 w Ammo am » Ammo am » mqomezoo mazmzaamms mmHommm .Ac.ucoov we wanna 56 Relative densities of brambles and red pine were higher on treated sites, as was absolute density of red pine. Cover and FHD Percent cover in the lowest height stratum was significantly greater on controls than treatments in all years of the study, as was percent cover in the middle height stratum (Table 13). No statistically significant differences in percent. cover in. the highest. stratum 'were. observed. Foliage height diversity also differed between treatment and control sites in all years (Table 14). WILDLIFE RESPONSES Small Mammals A total of’9 small mammal species were trapped.during the study. Small mammal species composition varied somewhat from year to year. Only 3 species, deer mouse (W W) , least chipmunk (W minim) , and thirteen- lined ground squirrel (We LEW) . were trapped on both treatment and control sites in all trapping periods. Deer mouse was the most abundant species on treated sites. Thirteen-lined ground squirrel was the most commonly captured species on controls until 1990, when numbers were exceeded by jumping mouse (ngng spp.), least chipmunk, and masked shrew (gore); clams) . Shorttail shrews (glaring bzgyiggndg) were captured consistently, but in low numbers. 57 :uo.O V my WHOHHCOO BORN “COHGHHfiU %H9CflUfiNHGOHm: Amo.o v my maouucoo scum unmummufio aaucmofiuflcwam. A~.oc m.o A~.qc p.44 .e.ac H.5m oomazoo Ao.oc. o.o .zxm.oc m.~ :gm.mv n.~¢ azmze¢mms omma An.oc 5.6 .H.mc n.o~ Ao.ac >.mm aomezoo 8.3 o.o :34: o; .293 «.mn sausage $3 Ao.ov o.o Am.mc m.m~ An.mc «.mm nomazoo Ao.ov o.o :go.oc o.o .Ae.aflc e.me azmzacmme some Ao.oc o.o A~.ov e.~a Ao.oflc >.v> aomezoo Ao.oc o.o _:Ao.oc o.o .Ao.nac «.ma ezmzadmma mama Ammo 55A Ammo an-” Ammo adv zoecmam groan: mews maHm mew» .omma can .omma .hmma .omma an .cmmfinodz .masmswsom woman 0:» ca Amaouusoov usouooao can Amusoauoouuv msoHumusoHa mafia own omuum>soo so vacuum usofimn n ca sofiuoummo> no u0>oo usmoumm .nH manna 58 Table 14. Foliage height diversity (FHD) values for converted red pine plantations (treatments) and clearcuts (controls) in the Upper Peninsula, Michigan, in 1986, 1987, 1988, and 1990. YEAR SITE TYPE FHD (SE) 1986 TREATMENT 0.01 (0.01)* CONTROL 0.13 (0.04) 1987 TREATMENT 0.00 (0.00)“ CONTROL 0.22 (0.02) 1988 TREATMENT 0.05 (0.01)" CONTROL 0.20 (0.03) 1990 TREATMENT 0.09 (0.02)“ CONTROL 0.28 (0.01) 'Significantly different from controls (P < 0.05) “Significantly different from controls (P < 0.01) 59 Masked shrews and meadow voles (aim pgnnsylyanigus) were trapped consistently in 1988 and 1990, but were not present in 1987. Jumping mouse, eastern chipmunk (mm M) , and redback vole ( g1 gthrionomys ml) were trapped inconsistently on a limited number of sites. Total numbers of individual animals captured were consistently higher on treated sites, though not always significantly so. 0f the 6 species captured in 1987, only one, deer mouse, was common (Table 15). Shorttail shrews were found on controls only in July, and only 1 individual was captured. One jumping mouse and 1 redback vole were trapped in August on a control site. The total number of individual animals was significantly greater on treated sites than controls in August (P < 0.05). No other significant differences were observed, but a few associations were apparent. August abundance was negatively correlated with percent cover in the lowest height stratum, and August deer mouse abundance was negatively correlated with both percent cover <1m in height and density of woody stems >1m in height. Correlation coefficients for statistically significant associations between abundance of small mammal population variables and vegetative characteristics in all years are listed in Table A-2. A total of 8 species were trapped in 1988 (Table 16). Eastern chipmunks were captured on controls only, and 2 species, shorttail shrew and jumping mouse, were captured only during the August trapping period. The only significant difference found was in deer mice, which were more abundant on Table 15. 60 included), species richness, and species diversity on converted red pine plantations Mean (SE) small mammal abundance (recaptures not (treatments) and clearcuts (controls) in the Upper Peninsula, Michigan, in July and August of 1987. PERIOD SPECIES July August Deer mouse Least chipmunk Shorttail shrew Thirteen-lined ground squirrel TOTAL INDIVIDUALS SPECIES RICHNESS DIVERSITY Deer mouse Jumping mouse Least chipmunk Redback vole Shorttail shrew Thirteen-lined ground squirrel TOTAL INDIVIDUALS SPECIES RICHNESS DIVERSITY ABUNDANCE Treatments Controls 7.00 (2.67) 1.20 (0.72) 0.60 (0.36) 0.40 (0.22) 0.00 (0.00) 0.20 (0.18) 1.80 (1.61) 2.20 (1.21) 9.40 (2.96) 4.00 (0.75) 1.40 (0.36) 1.40 (0.22) 0.15 (0.06) 0.10 (0.06) 8.40 (4.28) 2.20 (0.87) 0.00 (0.00) 0.20 (0.18) 0.80 (0.44) 0.20 (0.18) 0.00 (0.00) 0.20 (0.18) 1.00 (0.89) 0.40 (0.36) 2.60 (2.33) 3.20 (1.95) 12.80 (3.92)‘ 6.40 (1.19) 1.60 (0.22) 1.80 (0.44) 0.16 (0.06) 0.16 (0.09) 'Significantly different from controls (P < 0.05) Table 16. 61 included), species richness, diversity on converted red pine plantations and species Mean (SE) small mammal abundance (recaptures not (treatments) and clearcuts (controls) in the Upper Peninsula, Michigan, in July and August of 1988. PERIOD SPECIES July August Deer mouse Eastern chipmunk Least chipmunk Masked shrew Meadow vole Thirteen-lined ground squirrel TOTAL INDIVIDUALS SPECIES RICHNESS DIVERSITY Deer mouse Eastern chipmunk Jumping mouse Least chipmunk Masked shrew Meadow vole Shorttail shrew Thirteen-lined ground squirrel TOTAL INDIVIDUALS SPECIES RICHNESS ABUNDANCE Treatments Controls 10.60 (2.93) 1.20 (0.87) 0.00 (0.00) 0.60 (0.54) 5.60 (3.21) 2.60 (1.28) 0.00 (0.00) 0.40 (0.36) 0.20 (0.18) 0.80 (0.52) 1.40 (1.25) 2.60 (0.22) 17.80 (5.10) 8.20 (0.87) 1.80 (0.44) 2.60 (0.22) 0.22 (0.05) 0.32 (0.05) 8.00 (1.81)‘ 1.60 (0.83) 0.00 (0.00) 0.60 (0.36) 0.20 (0.18) 1.20 (0.44) 4.20 (1.31) 1.60 (1.00) 0.40 (0.22) 1.00 (0.57) 1.80 (1.00) 3.40 (2.43) 0.40 (0.22) 0.20 (0.18) 3.00 (1.70) 4.00 (0.94) 18.00 (2.21) 13.60 (2.22) 4.00 (0.40) 4.00 (0.94) 62 Table 16 (cont'd). PERIOD SPECIES ABUNDANCE TREATMENTS CONTROLS DIVERSITY 0.48 (0.05) 0.39 (0.09) *Significantly different from controls (P < 0.05) 63 treated sites than controls in August (P < 0.05). As in 1987, total August abundance in 1988 was negatively correlated with percent cover <1m in height, and July abundance was negatively correlated with density of woody vegetation >1m in height. Small mammal species richness increased with percent cover in the 1-7m height stratum and PHD in July. Deer mouse abundance decreased with increasing percent cover in both the <1m and 1-7m height strata and with density of woody vegetation in August. All of the 8 species trapped in 1990 occurred on both treatment and control sites, although 4 species (jumping mouse, meadow vole, redback vole, and shorttail shrew) were captured in very small numbers (Table 17) . Redback voles were found only during the July trapping period, and only 1 individual was trapped on each site type. Deer mice were significantly more abundant on treated sites in both July and August (P < 0. 05) , and masked shrews were more abundant on controls in August (P < 0.10). The total number of individuals was greater on treatments in July (P < 0.05) , and species diversity was higher on controls in August (P < 0.05) . A number of associations between small mammal populations and vegetative characteristics were observed in 1990. Total abundance was negatively correlated with percent cover in both the <1m and 1-7m height strata in July, while species diversity was positively correlated with percent cover in the lowest stratum and density of woody vegetation in August. Deer mouse abundance decreased as PHD and percent cover in the Table 17. 64 included), species richness, and species diversity on converted red pine plantations Mean (SE) small mammal abundance (recaptures not (treatments) and clearcuts (controls) in the Upper Peninsula, Michigan, in July and August of 1990. PERIOD SPECIES ABUNDANCE Treatments Controls July August Deer mouse Jumping mouse Least chipmunk Masked shrew Meadow vole Redback vole Shorttail shrew Thirteen-lined ground squirrel TOTAL INDIVIDUALS SPECIES RICHNESS DIVERSITY Deer mouse Jumping mouse Least chipmunk Masked shrew Meadow vole Shorttail shrew Thirteen-lined ground squirrel TOTAL INDIVIDUALS 19.80 (4.69)" 0.00 8.40 0.00 1.00 0.20 0.00 0.37 19.00 2.00 0.50 0.00 0.17 1.83 27.33 (0.00) (2.84) (0.00) (0.57) (0.18) (0.00) (1.99) 2.40 4.40 0.40 2.80 1.20 0.20 0.40 0.40 (5.59)“ 12.20 (0.22) (0.05) (5.62)*' (0.93) (1.14) (0.34)‘ (0.00) (0.17) (0.95) (6.91) 4.00 0.44 5.00 1.67 5.83 3.83 0.67 0.67 0.83 18.50 (1.15) (1.66) (0.22) (1.31) (0.44) (0.18) (0.22) (0.22) (1.91) (0.69) (0.06) (2.16) (1.05) (2.44) (1.40) (0.49) (0.42) (0.54) (5.89) 65 Table 17 (cont'd). PERIOD SPECIES ABUNDANCE Treatments Controls SPECIES RICHNESS 3.33 (0.76) 3.50 (0.81) DIVERSITY 0.40 (0.05)“r 0.55 (0.03) 'Significantly different from controls (P < 0.10) “Significantly different from controls (P < 0.05) 66 lowest and middle height strata increased in both trapping periods. In addition, deer mouse abundance was negatively correlated with density of woody vegetation in August. Also in August, masked shrew abundance was positively associated with percent cover in the lowest height stratum and density of woody vegetation. Birds Forty-nine bird species were identified in censuses on the study plots in 1987 and 1990, 20 of which were common to both years (Table A-3) . The most abundant species were American goldfinch (N 1115115) , American robin (W 21132012211115). chipping sparrow (5.0119116 035591103) . eastern kingbird (W W) , song sparrow (MM W) , and white-throated sparrow (mm Wis) . Of the 33 species censused in 1987, white-throated sparrow was the most common, followed by red-winged blackbird (Mus phoenigggg) and song sparrow (Table 18). These species were relatively common on both treatment and control sites. A single observation was made for 9 species; American crow (gems W) . American woodcock (Bhilehela miner) . black-capped chickadee (Ram W) , northern raven (Q. m) , red-breasted nuthatch (film W) , red- tailed hawk (3.92.9 Wig), upland sandpiper (mi; W), yellow-bellied sapsucker (W yarius), and yellow warbler (mm W). Blue jays (511002911310 M) . brown threshers (W 111mm) . and 67 Table 18. Bird density, species richness, and species diversity on converted red pine plantations (treatments) and clearcuts (controls) in the Upper Peninsula, Michigan, in 1987. SPECIES TREATMENTS Birds/ha (SE) CONTROLS Birds/ha (SE) American crow 0.00 (0.00) 0.03 (0.02) American goldfinch 0.48 (0.17) 0.45 (0.21) American robin 0.43 (0.15) 0.40 (0.04) American woodcock 0.03 (0.02) 0.00 (0.00) Black-capped chickadee 0.00 (0.00) 0.03 (0.02) Blue jay 0.00 (0.00)* 0.21 (0.06) Brown-headed cowbird 0.13 (0.06) 0.00 (0.00) Brown thrasher 0.00 (0.00) 0.13 (0.11) Chipping sparrow 0.48 (0.31) 0.50 (0.09) Clay-colored sparrow 0.03 (0.02)‘ 0.21 (0.02) Common flicker 0.19 (0.03) 0.08 (0.07) Common grackle 0.03 (0.02) 0.05 (0.04) Common snipe 0.00 (0.00) 0.11 (0.09) Eastern bluebird 0.13 (0.06) 0.00 (0.00) Eastern kingbird 0.24 (0.11) 0.13 (0.11) European starling 0.11 (0.09) 0.00 (0.00) Fox sparrow 0.05 (0.04) 0.16 (0.08) Killdeer 0.13 (0.08) 0.11 (0.09) Nashville warbler 0.05 (0.04) 0.11 (0.04) Northern raven 0.03 (0.02) 0.00 (0.00) Ovenbird 0.24 (0.16) 0.24 (0.13) Table 18 (Cont'd). 68 SPECIES TREATMENTS CONTROLS Birds/ha (SE) Birds/ha (SE) Red-breasted nuthatch 0.00 (0.00) 0.03 (0.02) Red-tailed hawk 0.00 (0.00) 0.03 (0.02) Red-winged blackbird 1.06 (0.31) 0.19 (0.12) Rufous-sided towhee 0.08 (0.07) 0.00 (0.00) Song sparrow 0.90 (0.21) 1.35 (0.11) Tennessee warbler 0.03 (0.02) 0.08 (0.08) Tree swallow 0.03 (0.02) 0.08 (0.04) Upland sandpiper 0.03 (0.02) 0.00 (0.00) White-throated sparrow 1.25 (0.22) 1.94 (0.21) Yellow-bellied sapsucker 0.00 (0.00) 0.03 (0.02) Yellow-rumped warbler 0.05 (0.02) 0.24 (0.11) Yellow warbler 0.00 (0.00) 0.03 (0.02) TOTAL INDIVIDUALS 6.18 (0.80) 6.92 (0.45) SPECIES RICHNESS 14.00 (0.47) 16.00 (1.70) DIVERSITY 0.95 (0.02) 0.98 (0.03) 1'Significantly different from controls (P < 0.05) 69 common snipes (Capella galnnagg) were found on controls exclusively in multiple sightings. Brown-headed cowbirds (Mom agar) , eastern bluebirds (5.18.1.1; siglis) , European starlings (m 111193115) , and rufous-sided towhees (£121.19 W) were identified only on treated sites and in several sightings. Two species, blue jay and clay-colored sparrow (S. 211113;) , were found in significantly higher densities on controls than treatments (P < 0.05) . No correlations between bird population variables and vegetative characteristics were apparent in 1987. A total of 36 bird species were identified during the 1990 census period (Table 19) . Of these, 7 were sighted only once; American woodcock, Carolina wren (W W) , house wren (W W) , northern oriole (M Ma) , ruby-throated hummingbird (W M) , ruffed grouse (3.9.0.15: M1135) , and white-eyed vireo (11189 Mg.) . 0f the species that were counted more than once, black-capped chickadee, cedar waxwing (figmbygilla W) , chestnut-sided warbler (p_,_ W) , common flicker (921293;: 2111:2195) . hermit thrush (931mm 92111151115) . Nashville warbler (mm W), rufous-sided towhee, and whip-poor-will (W W) occurred on controls exclusively, and common yellowthroat (Wig W), field sparrow (5. P3121112). killdeer (gharggzius W), tree swallow (W Mon), and vesper sparrow (2mm wings) occurred only on treatments. All of the 4 species that exhibited significant differences in 70 Table 19. Bird density, species richness, and species diversity on converted red pine plantations (treatments) and clearcuts (controls) in the Upper Peninsula, Michigan, in 1990. SPECIES TREATMENTS CONTROLS Birds/ha (SE) Birds/ha (SE) American goldfinch 0.77 (0.26) 1.18 (0.39) American robin 0.67 (0.15) 0.57 (0.34) American woodcock 0.00 (0.00) 0.03 (0.03) Black and white warbler 0.03 (0.03) 0.13 (0.08) Black-capped chickadee 0.00 (0.00)’ 0.38 (0.12) Blue jay 0.06 (0.06)‘ 0.67 (0.14) Brown-headed cowbird 0.03 (0.03) 0.13 (0.08) Brown thrasher 0.06 (0.06) 0.19 (0.11) Carolina wren 0.03 (0.03) 0.00 (0.00) Cedar waxwing 0.00 (0.00) 0.06 (0.06) Chestnut-sided warbler 0.00 (0.00) 0.10 (0.06) Chipping sparrow 1.59 (0.58) 1.40 (0.15) Clay-colored sparrow 0.06 (0.04) 0.10 (0.04) Common flicker 0.00 (0.00) 0.10 (0.09) Common grackle 0.03 (0.03) 0.26 (0.15) Common yellowthroat 0.06 (0.04) 0.00 (0.00) Eastern bluebird 0.10 (0.06) 0.06 (0.06) Eastern kingbird 0.92 (0.32) 0.32 (0.16) Field sparrow 0.06 (0.06) 0.00 (0.00) Hermit thrush 0.00 (0.00) 0.22 (0.09) House wren 10.00 (0.00) 0.03 (0.03) Table 19 (Cont'd). 71 SPECIES TREATMENTS CONTROLS Birds/ha (SE) Birds/ha (SE) Indigo bunting 0.29 (0.11) 1.27 (0.37) Killdeer 0.10 (0.09) 0.00 (0.00) Nashville warbler 0.00 (0.00)“’ 2.04 (0.41) Northern oriole 0.00 (0.00) 0.03 (0.03) Red-eyed vireo 0.03 (0.03) 0.16 (0.08) Red-winged blackbird 0.16 (0.09) 0.10 (0.06) Ruby-throated hummingbird 0.00 (0.00) 0.03 (0.03) Ruffed grouse 0.00 (0.00) 0.03 (0.03) Rufous-sided towhee 0.00 (0.00) 0.10 (0.06) Song sparrow 1.85 (0.51) 1.18 (0.30) Tree swallow 0.16 (0.09) 0.00 (0.00) Vesper sparrow 0.10 (0.06) 0.00 (0.00) Whip-poor-will 0.00 (0.00) 0.13 (0.05) White-eyed vireo 0.00 (0.00) 0.03 (0.03) White-throated sparrow 0.06 (0.06)‘ 1.21 (0.22) TOTAL INDIVIDUALS 7.23 (0.92)‘ 12.23 (0.69) SPECIES RICHNESS 10.00 (0.85)‘ 16.20 (0.52) DIVERSITY 0.83 (0.05)“’ 1.04 (0.01) 'Significantly different from controls (P < 0.05) "Significantly different from controls (P < 0.01) 72 density were more common on controls. These were black-capped chickadee, blue jay, white-throated sparrow, and Nashville warbler. In addition, total density, species richness (P < 0.05) , and species diversity (P < 0.01) were higher on controls than treatments. A number of significant trends in bird population variables and vegetative characteristics were observed in 1990 (Table A-4) . Total bird density, species richness, and species diversity increased with percent cover in all height strata. Species richness and diversity were highly positively correlated with FHD and density of woody vegetation, and bird density also increased with FHD. Densities of 2 species were correlated with vegetative variables; American robin density tended to decrease as density of woody vegetation increased, and indigo bunting density (W 111m) exhibited positive correlations with FHD, density of woody vegetation, and percent cover in the middle stratum. Deer A mean of 19.93 12.32 tracks/0.2km was counted on treated sites in 1988, and 30.07 $3.73 tracks/0.2km were counted on controls. In 1990, 35.00 18.94 and 25.00 112.25 pellet groups/ha were found on treatments and controls, respectively. No statistically significant differences were found. DISCUSSION VEGETATIVE RESPONSES Herbaceous species richness differed to varying degrees between treated sites and controls in all years, with the greatest difference occurring in 1986, when controls supported nearly twice as many herbaceous species as treatments. This difference was most likely the result of heavy herbicide treatments on 4 of the 5 treated plots in this year. Although hexazinone is used primarily to reduce woody vegetation, it is also effective against herbaceous species (Nelson et al. 1981, Blake et al. 1987). The slight increasing trend in species richness over the course of the study is characteristic of newly-cut stands but may also have been the result of a decrease in herbicide applications after 1986. Although the number of herbaceous species was not significantly different between treated sites and controls, total plant frequencies were higher on control sites in all years. Occurrence of herbaceous species should be of interest to land managers managing red pine because many of the species found on the study sites are important wildlife forages. Effects of hexazinone on herbaceous species have been short- lived, with measurable differences in herb response 73 74 disappearing by the end of the second growing season (Blake et al. 1987). In this study, however, the effect of the herbicide on the frequency of occurrence of herbaceous plants is apparent for up to 3 years. Although soil profiles reveal the sites to be similar in overall soil quality, the scarcity of deciduous leaves in plantations may have retarded invasion of new species and reproduction of established species. Deciduous leaf litter decomposes rapidly and increases nutrient availability. It is thus assumed that residual and secondary effects of heavy herbicide treatments early in the study, along with the effects of scattered additional treatments in 1987, 1988, 1989, and 1990, were sufficient to suppress herbaceous growth throughout the study period. It is interesting to note that a number of the species which were more common on treatment plots were species which tend to do well in highly disturbed areas. This group of species included thistle, horseweed, lamb's quarters, common St. Johnswort, common mullein, and yellow goatsbeard, which are generally poor wildlife forages, except for certain granivorous species such as the American goldfinch. A number of the woody species occurring on the study area are important wildlife cover and forage species, and analysis of woody vegetative structure and composition reveals extreme differences between the treatment and control sites. In addition, the presence of hardwood species may affect site suitability by providing high quality leaf litter. The most valuable of these species to wildlife are aspen, maple, 75 cherry, willow, birch, dogwood, serviceberry, blueberry, raspberry, and honeysuckle. Frequencies of woody species <1m in height generally showed the same trends as those of herbaceous species. Only rarely was a woody species more common on treated plots than controls. Brambles, birch, cherry, red maple, white spruce, and balsam fir on treatments were comparable in frequency to controls in some years, but only birch and brambles regularly reached heights of >1m in significant numbers on treatments. Density of woody vegetation >1m in height increased temporally on controls as would be expected in natural regeneration, and the drastic differences in density between treatment and control sites illustrates clearly the effect of repeated herbicide treatments The result is a loss of biomass of woody vegetation. Blake et a1. (1987) reported a nearly 50% decrease in total plant biomass after'the first growing season on a loblolly plantation treated with hexazinone. Similar results.have been.reported.by Hurst and.Warren (1982), Slay et a1. (1987), and Hurst (1989), although effects were short- lived. Brambles and.honeysuckles have exhibited tolerance to hexazinone in the past (Michael 1985, Hurst 1989), and brambles appeared to be at least partially resistant in this study. Biomass was not measured directly in this study, but analysis of percent cover, frequency, and density data clearly show a reduction in plant material on treated plots. The similarity in species richness between treated sites and controls suggests that site to site variation was not a major 76 factor; the increased densities are the result of greater occurrence of species already present on both site types, and not of invasion by new species. Woody plants reached heights of >1m relatively infrequently on treated sites in the 2 years following any herbicide application. Those trees which were able to keep a tenuous but consistent foothold on plantations were fast-growing species, such as balsam fir, birch, and cherry, all of which are capable of reaching 1m in height in 1 to 2 years. These species occurred most frequently on sites that had not been treated in the 2 years before the sampling period. Other evidence suggests that repeated applications are necessary to prevent establishment of woody species (Carter et al. 1975, Hurst 1989). The extreme sparseness of trees on sites that had been treated up to 1 year before sampling in this study is further evidence of hexazinone's initial efficiency. Analysis of available cover and FHD revealed a greater structural complexity and better developed under- and midstories on control plots. The importance of cover and vegetative complexity to wildlife is well documented. The near absence of cover >1m in height on treated plots illustrates one of the most obvious and extreme differences between treated sites and controls, and results in significantly lower FHD values on these plots. This consequence of herbicide treatment, along with a significant decrease in cover <1m in height, has serious implications concerning the suitability of pine plantations for wildlife. 77 However, this reduction in competing vegetation is the primary reason for the increase in softwood production that has been demonstrated on plantations. WILDLIFE RESPONSES Small Mammals Small mammal abundance, species richness, and species diversity were low throughout the study, but tended to increase temporally. Several species were represented by only a few individuals, making accurate comparisons difficult, but some general trends were apparent. Invariably, abundance was higher on treated sites than controls, largely due to the high deer mouse populations on the plantations. Deer mice inhabit a variety of vegetation types and were relatively abundant in recent clearcuts (Gashwiler 1970, Kirkland 1977), as well as plantations prepared with herbicides (Borrecco et al. 1979, Santillo 1987). Deer mouse abundance in this study consistently increased in abundance with herbicide treatment, and the correlations between abundance and some vegetative variables suggest that the increase was a response to decreases in vegetative cover and stem density. A similar abundance-cover relationship was reported by Eaton (1986) for this species, while M'Closkey and Lajoie (1975) observed a positive association between white-footed mice (2. M) and cover. The diet of deer mice contains insects, seeds, and vegetation (Hamilton 1941) . Vegetative cover was clearly less 78 abundant on treated sites, and thus it is apparent that cover was not a limiting factor for deer mice. Although insect response was not measured in this study, herbicide treatment has increased insect numbers (Santillo 1987), and some invertebrates, particularly ants, appeared to be more abundant on plantations than controls“ Possibly’ an increase in invertebrates allowed for a greater population of deer mice on the treated plots. A number of species appeared to prefer controls to treatments, but sample sizes were rarely large enough to show a significant difference. Masked shrews were consistently more abundant on control sites, although significantly so only in August 1990, when coinciding positive correlations with cover <1m in height and density of woody vegetation were observed. Decreases in vagrant shrew (nggx yagzang) populations have been reported after herbicide treatment in Oregon clearcuts (Borrecco et al. 1979), when grass cover was reduced. Masked shrews have been associated with moist habitats in Michigan (Getz 1961) , due to their high water requirements (Chew 1951). The moist microclimate maintained by thick grass cover was probably the primary factor affecting use of control sites by masked shrews. Decreases in abundance with herbicide treatment have been observed in field voles (Migxgtugflagrestis) (Teivainen et al. 1986), Oregon voles (M. grgggni) (Borrecco et al. 1979), and redback voles (Santillo 1987). Meadow‘voles were captured on a limited number of plots, making sample sizes too small to 79 accurately show differences. However, the species was present solely on moist sites. The preference of meadow voles for moist habitats with dense grass cover (Eadie 1953, Yahner 1983) and their largely vegetarian diet (Burt 1946) were the most likely factors to affect abundance on the study area. Least chipmunk abundance was greater on treated plots in all years and trapping periods.except August 1990, despite the species' tendency to prefer forest habitat. No associations between abundance and vegetative characteristics were observed, and given the wide range of food and cover requirements of chipmunks (Hamilton and Whitaker 1979) , no conclusions can be drawn. Different capture probabilities may exist between treatment and control sites for this species, and may be the result of animals spending less time on the ground when trees are available; least chipmunks use trees not only for escape (Caras 1967), but for sunning, resting, and occasionally nesting (Hamilton and Whitaker 1979). Abundance, species richness, and species diversity increased somewhat from July to August in 1987 and 1988, as well as increasing from 1987 to 1990. Differences were due mainly to the additions of or increased numbers of shorttail shrews, redback voles, jumping mice, masked shrews, and meadow voles. Shorttail shrews have a breeding season in late summer (Burt 1957) , and the presence of young may have increased captures of this species. Rainfall may also have been a significant factor influencing occurrence of some species. Small mammal populations are sensitive to precipitation (Getz 80 1968) and may have fluctuated in response to spring and summer rainfall levels. Jumping mice, redback voles, meadow voles, shorttail shrews, and masked shrews all prefer moist habitats (Burt 1946, Caras 1967, Getz 1968, Hamilton and Whitaker 1979, Yahner 1986) , and the study area experienced below average rainfall in spring of 1987 and 1988. The slight increase in rainfall from June through August 1987 was accompanied by slight increases in small mammal abundance, species richness, and species diversity. Similarly, near-drought conditions in May and June of 1988 were followed by average rainfall in July and above average rainfall in August, and again, small mammal population variables increased from July to August. In 1990, rainfall was unusually high from May through July, dropping to just below the norm in August, and no trends in abundance or species richness were apparent in this year. Further evidence that rainfall may have affected small mammal populations is the overall increase in small mammal abundance from 1987 to 1990, corresponding with an increase in total spring and summer rainfall over these years. Small mammal populations fluctuate to relatively large degrees from year to year (Haufler, pers. commun.), however, and the temporal changes observed in this study may simply have been the result of normal population variations. Birds Bird density, species richness, and species diversity were consistently higher on controls than treatments, although 81 statistically significant differences were detected only in 1990, by which time controls had developed substantial structural diversity. A shift in species composition occurred from 1987 to 1990, with the loss of 13 species from the study area and the addition of 16 species. Much of the variation between years can be explained by observer bias, since different observers were used for the 2 censuses. Many of the species present in 1 year and absent in the other, such as American crow, northern raven, common snipe, European starling, red-breasted nuthatch, red-tailed hawk, Tennessee warbler (W W), upland sandpiper, yellow- bellied sapsucker, yellow warbler, Carolina wren, house wren, northern oriole, ruby-throated hummingbird, ruffed grouse, field sparrow, white-eyed vireo, and cedar waxwing, were identified in very low numbers and were most likely just passing through the study area rather than breeding there. The variable Circular-plot method involves counting all birds using an area, including individuals feeding, hunting, and resting. The record of 1 species, ovenbird (aims ggzggapillng), in 1987 was likely due to observers including birds identified by call from surrounding forest. Several species present in 1990, including black and white warbler (W 2111;) , chestnut-sided warbler, red-eyed vireo (11339 W) , hermit thrush, and whip-poor-will, are characteristic of young or early successional forest, and the study area habitat in 1987 may not have been mature enough to support them. Common yellowthroats were observed only on 82 plots bordered by low, flooded areas after a wet spring in 1990, and these areas may have been.too dry for the species in 1987. One species, indigo bunting, appeared late in the census period in 1990 and may have been absent during the 1987 census period, which was shorter than that in 1990. Additionally, the 1987 census was conducted on three plots of each site type while the 1990 census utilized all 10 plots. This change may have resulted in some of the less ubiquitous species appearing only in the 1990 census. Of the 5 species exhibiting a statistically significant preference for control sites, only 1, blue jay, was significant in both years. Blue jay diet consists of nuts, fruits, and berries (Wilmore 1977), and the relative sparseness of fruit-producing vegetation on plantations may have limited the species on these sites. Clay-colored sparrows inhabit grasslands, thick brush, and forest edges (Bent 1968), and the 1987 plantations were probably too barren for even some grassland species. Treated sites in 1990 lacked the semi-open mixed stands and thick brush preferred by white-throated sparrows (Bent 1968) . Eaton (1986) reported a positive correlation between this species and amount of slash, and treated sites in 1987 could have contained enough slash to accommodate some individuals. It is also possible that the record of this species in 1987 was the result of miscalculation of birds' distances from census stations; like the ovenbird, this sparrow has a very loud call. 83 The abundance of black-capped chickadees on control plots was probably related to their feeding habits; the Species feeds on insects in trees of conifer and deciduous forests, and such habits would have been severely limiting on young pine plantations, which lacked the density of trees available on controls. In 1990, Nashville warblers were much more abundant on controls, due to the deficiency of cover in pine plantations in the 1-7m height stratum. Nashville warblers inhabit young mixed stands of aspen, birch, and balsam poplar and require a developed midstory (Griscom and Sprunt 1957) , and controls clearly provided better habitat for the species than treated sites. In general, Species commonly observed on the control sites inhabit young forest or brushy areas, and species found on the plantations usually occur in more open vegetation types, such as grasslands and fields. In this study, treatments and controls can be categorized into open field and second growth fOrest vegetation types, respectively. Some factors contributing to the bird population differences between treatments and controls are apparent from the associations observed between bird population variables and vegetative measures. The consistent, positive correlations between number and variety of birds and habitat variables suggest that vegetative density and structural complexity enabled the control sites to support a denser and more diverse avifauna than the treated sites. Such associations have been 84 examined and documented previously. In general, bird species diversity has increased with successional development (Balda 1975, Weins 1975, Edgerton and Thomas 1978, Shugart et al. 1978, Weins 1978, Capen 1979) . Positive correlations between bird diversity and vegetative structural complexity have been demonstrated by Balda (1975), Thomas et al. (1975), Weins (1975), Meslow (1978), Shugart et al. (1978), and Capen (1979). Increases in overall bird densities on the naturally regenerating plots were likely due to greater resource availability, especially for species limited by cover or food on plantations. A red pine plantation in Michigan supported fewer bird species and individuals than surrounding natural pin oak and jack pine communities (Gysel 1966). Slagsvold (1977) reported a 30% reduction in breeding bird abundance after spraying of deciduous scrub with herbicide, with numbers remaining low for 5 years post-treatment. Bird abundance also decreased on a Jeffrey pine (P. M) plantation prepared with herbicide, and all avian species dependent on snowbush (W M) were eliminated from the treated plot (Savidge 1978) . The controls sites in this study provided more cover for both ground and tree nesters; this cover not only increases potential nest sites, but may also provide greater protection from predators, possibly increasing survival rates. The greater availability of seed- and fruit- producing plants on control sites also provides food for granivores and frugivores, as evidenced by the presence of 85 cedar waxwings, blue jays, clay-colored sparrows, chipping sparrows, fox sparrows (mug 111m) , white-throated sparrows, indigo buntings, and American goldfinches. These species also prefer open woods or brush vegetation types to grasslands. The effects of herbicides on insect populations are not well documented, but Santillo (1987) observed an increase in invertebrate numbers after herbicide treatment on conifer plantations, and Johansen (1981) reported an abundance of insects on young loblolly pine plantations. It has also been suggested that insect hawkers have difficulty maneuvering in dense stands (Willson 1974). These factors suggest that treated sites would support greater numbers of insectivores than controls. Three insectivorous species were found in greater numbers on treated sites, though differences were not statistically significant. Eastern bluebirds, eastern kingbirds, and tree swallows capture their prey in flight and may have been limited on control sites by the density of trees >1m in height. Eastern bluebirds and tree swallows are cavity nesters, and occurrence of these species was very likely related to the availability of snags in areas surrounding the study plots. Other insectivores were able to inhabit control sites in relative abundance, indicating that insect food was not severely limited on controls. These species included brown thrashers, hermit thrushes, chestnut-sided warblers, red-eyed vireos, and whip-poor-wills, as well as Nashville warblers and blackecapped chickadees, which showed a 86 significant preference for controls over treatments. Clearly, treated sites were more suitable for species limited to grassland areas, such as killdeer, vesper sparrow, and field sparrow. Species with broad food habitats, such as song sparrow, red-winged blackbird, common. grackle (Quisgalgs (guiggula), and. brown-headed. cowbird. also seemed. to fare particularly well on the plantations. Despite the affinity of some insectivores and grassland species to the treated plots, the naturally regenerating controls in this study possess a number of features that enable them to support more avian species than plantations. Pine forests generally' support fewer' species than :mixed forests (Tramer 1969, Thomas et al. 1975, weins 1975, Driscoll 1977, Dickson 1978, Meyers and Johnson 1978); however, this is largely‘ due to the early canopy closure and sparse understory typical of mature pine stands. Johnson (1987) reported canopy closure to occur earlier in slash pine (P. elliggtii) plantations than in natural stands, and noted a corresponding decrease in summer bird populations on plantations. The use of herbicides reduced low vegetative cover, thereby decreasing bird species richness even on young plantations. Controls, on the other hand, had significantly more cover in all height strata. Finally, species with wide distributions have been found in pure pine stands, while these species are joined by others with more specific habitat requirements in mixed and broadleaf forest (Eiberle and von Hirschheydt 1983). 87 The differences in bird density and species richness resulted in higher species diversity on controls than treatments. Thus, the relationship of bird species diversity to vegetative characteristics can be explained in terms of the variation in resource availability and vegetation structure between treated plantations and naturally regenerating stands. Deer This study yielded no statistically significant results pertaining to deer use of herbicided pine plantations. Results do indicate that deer made use of both treated sites and controls. Assumptions and biases associated with both the track count and pellet-count techniques must be considered before it can be concluded that conversion did not affect deer use of the study area. In 1988, track counts were conducted on roads bordering 3 treated and 3 control plots. Road surfaces varied from sand and mud, which provided good substrate, to gravel, which did not show tracks as clearly. Roads surrounding plantations were generally better maintained and had less vegetation growing on them, due to their frequent use for herbicide application and growth monitoring. Thus, tracks would tend to be more visible on these roads. Van Dyke et al. (1986) found the track method to be less sensitive to changes in mountain lion (29.1.1.5 W) density when tracking conditions were poor, suggesting that track counts on poor road surfaces may not be a reliable indicator of density. A difference in 88 accuracy between site types would invalidate comparisons of relative densities. It is also likely that the roads in close proximity to plantations were more frequently used by vehicles, since they were generally in better condition. Any difference in the amount of such disturbance between site types could affect the tendency for deer to cross roads. The locations of the roads used for track counts may have biased results; tracks on the roads could have been an indication of deer use of surrounding vegetation types, rather than the plots themselves. Roads may have been used by deer as travel lanes to and from areas adjacent to study plots, resulting in the inclusion of tracks made by deer that may never have entered the study area. Finally, deer use of plantations can A be expected to differ with distance to an edge, and deer could have avoided most of a plantation while still leaving tracks on boundary roads. In an attempt to eliminate these potential sources of error, pellet-group counts were employed to assess deer use in 1990. Estimates were used only to rank relative use of the study plots. This method has received criticism because of its associated assumptions (Fuller 1991). It must be assumed that no groups are lost during the census period and there is no observer bias in locating, aging, and identifying groups. It is further assumed that defecation rates are constant over time spent in an area. Collins (1981) recorded different defecation rates during various activities in mule deer (Qggggilggg hemigngg), and Neff (1968) summarized data 89 indicating that defecation rates varied with range condition and forage quality. It is therefore possible that differences in vegetative characteristics between site types could affect defecation rates. Thus, the assumption that defecation rates are linearly related to time spent in an area is of questionable validity, but was accepted in this study. However, because of the small total number of pellet-groups counted on the study plots, intraspecific variations in defecation rates could have affected results. Allowing a longer time for pellet deposition might improve accuracy of results by increasing the total number groups counted. DEER SUMMER HABITAT SUITABILITY Food quality and abundance are usually limiting for white-tailed deer in the southern United States (Hurst and Warren 1982), and availability of preferred forage appears to be the primary factor influencing deer summer habitat use in the Upper Great Lakes Region (Kohn and Mooty 1971). Summer diet of deer is varied, consisting of new growth of browse, herbaceouS‘vegetation, shrubs,‘and tree foliage (Blouch 1984), especially aspen leaves (McCaffery et al. 1974). Fruit, berries, and mushrooms are also readily eaten (Rogers et al. 1981, Blouch 1984). Pine is eaten only under stress conditions (Halls and Boyd 1982), and spruce, balsam fir, tamarack, and speckled alder are also poor foods (Rogers et al. 1981, Blouch 1984). A.hardwood.or'herbaceous component is 90 necessary to support a healthy deer population (Johnson 1987) . A number of studies evaluating deer forage in pine plantations have been conducted. Gysel (1966) attributed a decline in white-tailed deer numbers in a Michigan red pine plantation due to a lack of low-growing foods. Deer forage was reduced more drastically on pine plantations prepared with.hexazinone when compared to plantations prepared by mechanical methods (McComb and. Hurst 1987), and Savidge (1978) reported. a significant difference in mule deer use between a Jeffrey pine plantation treated with herbiCide and an untreated plantation. Favorable browse species were virtually eliminated by the herbicide treatment, and deer were not observed on treated areas. Conversion of mature pine-hardwood forests to loblolly pine through the use of herbicides not only reduced deer forage, but also removed hard and soft mast and mushrooms (Hurst and Warren 1982). Hurst and Warren also noted a lack of cover on intensely prepared sites. Blake et al. (1987) reported a decrease in forage biomass on a loblolly plantation following treatment with hexazinone. They noted, however, resistance of 3 key forages; blackberry, honeysuckle, and greenbrier (Smilax spp.). Rogers et al. (1981) listed results from several deer forage utilization studies conducted in the northern Great Lakes Region. Analysis of these data indicated approximately 50% to 60% of white-tailed deer diets in July and August to be woody browse, and 30% to 40% mushrooms and forbs“ Grasses and sedges made up 6% or less of the summer diet. Woody species 91 eaten in the greatest quantities were aspen, hazelnut (m spp.), bush-honeysuckle (Digzgilla spp.), red maple, cherry, brambles, birch, and ‘willowu Of ‘these species, birch, brambles, cherry, and aspen were present in significant amounts on plantations in this study. By comparison, naturally regenerating plots contained all of these species, and all except birch were higher in frequency and density on controls. Additional common woody species on treated sites were generally poor forage species, including balsam fir, red pine, and white spruce. While adequate browse for small numbers of individuals may exist on plantations in the form of birch, cherry, and brambles, the browse component of controls was superior in both species composition and biomass. Herbaceous species constitute a significant portion of summer white-tailed deer food. Considerable amounts of wild strawberry, goldenrod (Soligggg spp.), aster, bracken fern (M19111!) M), rough cinquefoil, clover (W spp.) , violet, sarsaparilla (Anglia spp.) , vetch (112;; spp.) , spreading dogbane, and hawkweed were consumed by deer in the Upper Great Lakes Region (Rogers et al. 1981). Most of these species, in addition to several other desirable herbaceous species, were abundant on both controls and treatments. Availability of herbaceous forage would probably not limit deer on the study plots if an adequate woody browse component was available. However, the relative lack of woody forage on plantations might necessitate an increase in consumption of herbaceous species, in which case overall food availability 92 may become limiting to deer on treated sites. ‘When.both woody and herbaceous foods are considered, it is evident that controls were :more suitable for deer in forage species composition and availability. Hiding cover is also an important summer deer habitat component. Thomas et al. (1979) defined hiding cover as vegetation capable of concealing 90% or more of an adult animal from human vision at a distance of 61m or less. Heaviest deer use occurs within 183m of the edge between cover and forage areas (Thomas et al. 1979). Deer are capable of running through dense vegetation, and therefore cover can be used for both hiding and escape. Suitable cover for deer occurs at a density of 3000 stems/ha >1m in height (Bender and Haufler, unpubl.). By this standard, adequate cover was not available on treated plots in any year. Density was sufficient on control Sites in all years except 1987 (see Table 10). The low density estimate in this year resulted from extremely low density values obtained on 2 plots. Both of these plots yielded higher estimates in the other years. Different field personnel collected data in each year, and variations may have been due to sampling biases. SUMMARY AND RECOMMENDATIONS Results of this study illustrate the extensive habitat alteration caused by the convertion of hardwood forest to red pine. Structural changes in vegetation following herbicide treatment were evident in percent cover, stem density, and FHD estimates. Woody and herbaceous species composition was similar between site types, but total absolute frequencies were consistently greater on controls. These vegetative differences have implications for wildlife, particularly avian species. Small mammal numbers were low throughout the study period but increased slightly over time. .Abundance was consistently higher on plantations than on naturally regenerating plots, Significantly so in August 1987 and July 1990, due largely to the affinity of deer mice to the plantations. Treated sites and control sites supported similar numbers of species, but species diversity was significantly higher on controls in August 1990. Analysis of correlations suggests that percent of low'cover is one'variable‘affecting’small.mammal population dynamics, but moisture and food availability may also have affected populations. Bird density, species richness, and species diversity were higher on controls than treatments, and differences 93 94 between site types were significant in 1990. Between-year differences were largely explained by observer differences and small sample size. Site type differences were mainly the result of vegetative composition and structure alteration by herbicide treatment, although indirect effects, such as changes in insect abundance, may also have influenced habitat selection by birds. Treatments and controls belonged to different vegetation types. Treatments were basically open grasslands while controls more closely resembled second growth forest and thickets. Predictable shifts in avian species composition occurred as vegetative features of treatment and control plots diverged. Deer track counts and pellet-group counts yielded no significant results. Inherent biases, assumptions, and sources of error, along with small sample sizes, made deer use estimation extremely difficult. Deer did make use of both treated sites and controls. The effects of conversion on deer were more easily measured indirectly by qualifying the sites as deer habitat. Assessment of available forage species, forage abundance, and security cover indicated that converted sites provided sub-optimum deer habitat, while controls were more suitable due to their mixture of woody browse, herbaceous forage, and cover. The vegetative changes after herbicide treatment are severe enough to alter wildlife use of plantations. Considering the amount of land in the Upper Great Lakes Region presently or prospectively being used for softwood production, 95 it is imperative that wildlife be considered in management practices. The results of this study, as well as those in the literature, suggest that the preservation of a hardwood component in red pine stands would allow for a more diverse avifauna, better deer habitat, and possibly higher diversity of small mammal species. This might be achieved by encouraging regeneration of red pine on sites where the species occurs naturally. However, good seed crops occur relatively infrequently, and planting may be necessary to meet minimum timber stocking requirements. Because red pine grows most successfully on low quality sites, planting pine in such areas would reduce the need for competition control. Limiting site preparation to burning or a mechanical treatment would allow the survival of other woody vegetation. In addition, low stocking densities (1000 trees/ha) and thinning beginning at age 25 would improve understory for wildlife, as well as increase red pine growth rate. Important variables not considered in this study are plantation size and spacing. Plantations may be used by deer as forage areas provided cover is accessible. Recommended maximum width of forage areas is generally 220m, and deer forage areas should be no more than 183m from cover at any point. This would allow for plantations to be approximately 13 ha in size if square. At least 40% of a section should be left in cover. Consideration should also be given to the possibility of forest fragmentation. Complete interspersion of‘a management 96 area with plantations could allow edge species and predators to invade forest blocks, and may result in declines in interior species, particularly some nongame birds. Large areas in plantations of'varying'ages*will produce an unnatural and fragmented forest landscape. Therefore, some blocks of mature forest should be preserved, although additional research is needed to determine the exact stand size necessary to maintain populations of interior species. The effects of conversion to red pine are not well documented for all plantation ages. Vegetation and wildlife populations need to be monitored on plantations throughout the entire rotation to determine the success of 'management procedures on both timber and wildlife. 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Avian community organization and habitat structure. Ecology 55:1017-1029. Wilmore, S.B. 1977. Crows, jays, ravens, and their relatives. David and Charles, BC. 208pp. Yahner, R.H. 1983. Small mammals in farmstead shelterbelts. J. Wildl. Manage. 47:74-104. Yahner, R.H. 1986. Microhabitat use by small mammals in even-aged forest stands. Am. Midl. Nat. 115:174-180. APPENDIX 106 Table A-1. Vegetative species found on converted red pine plantations and clearcuts in the Upper Peninsula, Michigan, in 1986, 1987, 1988, and 1990. COMMON NAME SCIENTIFIC NAME HERBACEOUS Agrimony Agrimeaia 9122952231; American pennyroyal flggggmg pglggigiggg Aster Aster spp. Bluebead lily glintgnig bgzgglgg Bedstraw Gallup spp. Bicknell's cranesbill Geranigm pigkpglli Black-eyed Susan Bugbggkig hirta Black bindweed Eglyggnnm ggnyglynlgg Black medick Mgdigagg lupulina Black mustard Braggiga nigza Bladder campion .fiilgng Qgggpalgg Blue cohosh 922122021138 thalistreides Bristly crowfoot Bangngglgg pgngylyaniggg Bristly sarsaparilla 5:311; higpigg BunChberrY QQIDQ§.QensQen§i§ Butter-and-eggs Linaria 22192215 Canada mayflower Maiantngmum gaggggpgg chickweed Stellaria Spp-. Cerastium Spp- Common burdock Arctigm pings Common buttercup 8309022125 eerie Table A-1 (cont’d). 107 COMMON NAME SCIENTIFIC NAME Common cattail Common milkweed Common mullein Common nightshade Common plantain Common St. Johnswort Coneflower Cow vetch Curled dock Creeping snowberry Daisy fleabane Dandelion Downy wood mint Dwarf enchanter’s nightshade Early meadow rue Enchanter's nightshade Erect bindweed Eyebane Fern Field bindweed Field pussytoes Fireweed Fringed bindweed malatifelia mm mm mm 21811189911812: WW 81111932313891:- 21213210925 mm mammals Erigerenannnns IAIQXQQQE SPP- 21221111132111.8128 91132553812111: maligtmmdigim mm mm mm mapp- WW Mariam WW 29119201111911.1020: Table A-1 (cont’d). 108 COMMON NAME SCIENTIFIC NAME Fringed loosestrife Fringed polygala Fungus Goldenrod Goldthread Grasses and sedges Great lobelia Great St. Johnswort Ground cedar Hairy Solomon’s seal Hare figwort Hemp nettle Hooked crowfoot Hop clover Horsetail Horseweed Hound’s tongue Indian paintbrush Indian tobacco Lamb's quarters Large blue flag iris Large-leaved avens Least hop clover Welling WW Various 51211113912899- MW Beasppugarexsw- Lebeliasinhilitisa mm mm mm mm Table A-1 (cont’d). 109 COMMON NAME SCIENTIFIC NAME Lichen Mad-dog skullcap Maple-leaved goosefoot Marsh marigold Marsh skullcap Moss Narrow-leaved willowherb Night flowering catchfly Northern bugleweed Northern willowherb Orange hawkweed Oxeye daisy Oxeye sunflower Oyster plant Pale corydalis Pale smartweed Pale touch-me-not Partridgeberry Pearly everlasting Peppermint Plantain-leaved pussytoes Poison ivy Poke milkweed Various mm mm mm Various WW Silensneetiflera mm mm mm mm WW mm mm mm Imtisnenallida 8.1125282113392115 WW Meatbapiperita WWW 80118128122115 MM Table A-1 (cont'd). 110 COMMON NAME SCIENTIFIC NAME Prickly lettuce Purple-leaved willowherb Pyrola Queen Anne's lace Rattlesnake roots Red baneberry Red clover Rose twisted stalk Rough cinquefoil Rough-fruited cinquefoil Round-lobed hepatica Selfheal Sheep sorrel Shinleaf Shrubby St. Johnswort Spotted joe-pye weed Spotted knapweed Spotted touch-me-not Spreading dogbane Starflower Stinging nettle Strawberry blight Swamp milkweed 1.093112052011210 mm 81112108»- DQEQB§.£§IQ§§ Brenantheaepp- Agtaee IBDIQ Wm WM WW Waste 88000112000201.9000 81110911020190115 BBBQX.§£§§Q§§11§ Table A-1 (cont'd). 111 COMMON NAME SCIENTIFIC NAME Sweet Cicely Sweet coltsfoot Sweet everlasting Tall cinquefoil Tansy Thimbleweed Thistle Trillium Turtlehead Twisted stalk Violet Water horehound Water parsnip White campion White clover White lettuce White sweet clover White vervain Wild basil Wild bergamot Wild columbine Wild lettuce Wild mint mm Wm mm Wm WM Ansmenexirginiana Table A-1 (cont’d). 112 COMMON NAME SCIENTIFIC NAME Wild sarsaparilla Wild strawberry Wild thyme Wintergreen Wood anemone Yarrow Yellow avens Yellow corydalis Yellow goatsbeard Yellow hawkweed Yellow stargrass Yellow sweet clover Yellow wood sorrel WOODY American basswood American Beech Ash Balsam fir Balsam poplar Bigtooth aspen Birch Blueberry Brambles 000110 0001000110 80000010.2100101000 100000.000001100 5001000010 0000000000 0000000 001000010110 A0011100.01110f01100 0000 010001000 000200110.fl00010 1000000000 000100 810000100 00000000 8000010 0100000 001110000 00110100110 000110 00000000 $1110 000010000 80000 00000100110 80001000 800- 00100 00100000 800010100 00100010000 8000100.0000010000000 800010 800- 200010100 800- 80000 spp. Table A-1 (cont’d). 113 COMMON NAME SCIENTIFIC NAME Cherry Currant Dogwood Elderberry Elm Grape Hairy honeysuckle Hazelnut Hornbeam Ironwood Jack pine Northern bush-honeysuckle Northern white-cedar Pasture rose Poison ivy Quaking aspen Red maple Red pine Serviceberry Speckled alder Sugar maple Tamarack Viburnum £00000 sPP- 81000 800- 900000 800- 00000000 0000000010 11100.0 SPP- 11010 800. 10010000 01001100 90011100 800- 00.001011100101000 0000100000001101000 81000000001000 0.1001111101001030 10010000100000110 80000101011000 800000010000 20001000000010.1000 1100001000110 81011000810000 01010000100800- 01000210000 00000000001101 1001010010100 21000000800- 114 Table A-1 (cont'd). COMMON NAME SCIENTIFIC NAME Virginia creeper 20000000010000 001000000110 Waxberry 000000110000000000000000 White pine 21000 0010000 White spruce 21900 910020 Willow 00110 spp- 115 Table A-2. Correlation coefficients (r ) for statistically significant associations befiween small mammal population variables and vegetative characteristics on converted red pine plantations and clearcuts in the Upper Peninsula, Michigan, in 1987, 1988, and 1990. DATE POPULATION VEGETATIVE CORRELATION VARIABLE CHARACTERISTIC COEFFICIENT (rs) 1987 Deer mouse Percent cover -o.84** August abundance <1m in height 1987 Deer mouse Density of woody -0.64* August abundance vegetation 1987 Total small Percent cover -0.65* August mammal abundance <1m in height 1988 Deer mouse Percent cover -0.79** August abundance <1m in height 1988 Deer mouse Percent cover -0.61* August abundance 1-7m in height 1988 Deer mouse Density of woody -O.82** August abundance vegetation 1988 Total small Percent cover -0.70** August mammal abundance <1m in height 1988 Total small Density of woody -0.67** July mammal abundance vegetation 1988 Small mammal Percent cover 0.66** July species richness 1-7m in height 1988 Small mammal FHD 0.70** July species richness 1990 Deer mouse Percent cover -o.78** July abundance <1m in height 1990 Deer mouse Percent cover -O.81** August abundance <1m in height 1990 Deer mouse Percent cover -O.79** July abundance 1-7m in height Table A-2 (cont’d). 116 DATE POPULATION VEGETATIVE CORRELATION VARIABLE CHARACTERISTIC COEFFICIENT (rs) 1990 Deer mouse Percent cover -0.72* August Abundance 1-7m in height 1990 Deer mouse PRO -0 . 67* July abundance 1990 Deer mouse FHD -o . 59* August abundance 1990 Deer mouse Density of woody -0.67* August abundance vegetation 1990 Masked shrew Percent cover 0.60* August abundance <1m in height 1990 Masked shrew Density of woody 0.62* August abundance vegetation 1990 Total small Percent cover -O.68** July mammal abundance <1m in height 1990 Total small Percent cover -0.67** July mammal abundance 1-7m in height 1990 Small mammal Percent cover 0.73** August species diversity 7m in height FHD Percent cover <1m in height Percent cover 1-7m in height Percent cover >7m in height FHD Density of woody vegetation Percent cover <1m in height Table A-4 (cont'd). 121 POPULATION VARIABLE VEGETATIVE CHARACTERISTIC CORRELATION COEFFICIENT (rs) Species diversity Species diversity Species diversity Species diversity Percent cover 1-7m in height Percent cover >7m in height PHD Density of woody vegetation ** *** 'U'U'U AAA OOO OOH HUIO "WEWWILfiflTfllfiflflmflWflflfl‘5