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[UNIVERSITY LIBRARI'ES llilmflli illillllil: i H l” il ii H “ii in 31293 01718 8503 This is to certify that the thesis entitled The Effects of Controlled-Burn Management on the Ecology of Forest-Openings Vegetation in Southern Illinois presented by Alicia Suzanne Biagi has been accepted towards fulfillment of the requirements for M.S. degree in ,BQtaDY &_‘_Plant Pathology Major professor Peter G. Murphy Date M 0-7639 MSU is an Affirmative Action/Equal Opportunity Institution ton the Southern 1/0” LIBRARY M'CMQGn State University PLACE IN RETURN BOX to remove this checkout from your record. TO AVOID FINE return on or before date due. DATE DUE MTE DUE DATE DUE SEPZSZUUS 0910'09 1m damages-nu THE EFFECTS OF CONTROLLED-BURN MANAGEMENT ON THE ECOLOGY OF FOREST-OPENINGS VEGETATION IN SOUTHERN ILLINOIS By Alicia Suzanne Biagi A THESIS Submitted to Michigan State University in partial fulfillment of the requirements for the degree of MASTER OF SCIENCE Department of Botany and Plant Pathology 1998 ABSTRACT THE EFFECTS OF CONTROLLED-BURN MANAGEMENT ON THE ECOLOGY OF FOREST-OPENINGS VEGETATION IN SOUTHERN ILLINOIS By Alicia Suzanne Biagi Nine southern Illinois forest-openings examined for classification purposes in 1988 by A. Heikens of Southern Illinois University-Carbondale were re-examined in 1993 to determine the effect of interim fire management on vegetation composition and structure in three zones: opening, opening-forest transition, and adjacent forest. Site boundaries were mapped in 1994 and the vegetation strata of the three zones were compared. Forest-openings ranged from 450 to 7825 m2 in area. In all sites, the ground layer was a forest-prairie mixture with a high percentage of annuals, unusual matrix species and a dominant overstory species which varied by site. At sites which had been fire-managed more than once, openings were characterized by Brickellz‘a eupatoriodes and Silphium terebinthinaceum; these herbs were useful as indicator species. Vegetation of the burned and unburned forest-openings displayed unexpected similarities in herb life-form, life history, and proportion of grasses, forbs, legumes and exotics. However, management maintained the three vegetation zones; increased herb species diversity, richness and cover; and decreased woody species richness and density in the opening. Thus, fire-management can be recommended as a means of retarding the replacement of herbaceous communities by woody species in the region investigated. ACKNOWLEDGEMENTS My sincere thanks are expressed to my advisor, Dr. Peter Murphy for the immeasurable help and advise he has provided and for his generous support. Dr. Murphy earned my admiration and respect as a remarkable role model of a teacher, ecologist, mentor and friend. The project committee members, Dr. Francis Ekem and Dr. Frank Ewers are appreciated for their enthusiasm, review and approachability for consultation. Many thanks to those who helped fund the project, especially the Botany and Plant Pathology Department, the Paul Taylor Fund and the Shawnee National Forest. The study could not have been conducted without the consent of the respective landholders, the Shawnee National Forest Service, The Nature Conservancy and the Illinois Department of Conservation. In particular, K. Andrew West (IDNR) and Max Hutchison (TNC) are thanked for their ready assistance. I would like to thank Dr. Alice Heikens very much for her enthusiasm, constructive comments and sharing of data for the sake of the project. Clearly, her work provided a solid foundation for others interested in forest-openings. Elizabeth Shimp (USFS) aided me during the project in a variety of ways. Her advise, determination and caring attitude for myself and natural systems in no small way inspired the success of the study on the hottest of summer days. Similarly, Jody Shimp and Mark Basinger are thanked, if not for their comic relief, for sharing with me their vast knowledge and appreciation of plant identities in south Illinois. Many individuals are thanked for their involvement with specific aspects of the project. iii Tom Prang and Rob Rubinas helped with the field work. Alice Murphy and Lissa Leege helped plow through several drafts of the thesis with their comments. Several lab collegues were supportive and kindly gave advise. Most especially I would like to thank my parents and family who also shouldered the weight of the project from conception to completion. Achille D. Biagi offered and instructed through long hours the use of AutoCAD making the site maps, probably the most important outcome of the project, a reality. He also donated his computer for thesis completion. H. Eileen B. Biagi helped with field work and no less than six months of labor-intensive child care for my son. Other contributions from my parents included transportation during several moves, generous funding throughout the project, unintrusive but strong encouragement and love. Cosmo M. Bosela’s exuberance and playfulness were perfectly therapeutic and made time pass happily throughout the writing process. Alberta M. Bosela also made this project worthwhile. Above all my husband Michael I. Bosela is thanked for his long hours of companionship throughout the masters process, for his commentary, support and help with the thesis and for his vivacity and good nature. Michael’s acceptance and benevolence were crucial to the completion of this project. iv TI TABLE OF CONTENTS LIST OF TABLES LIST OF FIGURES INTRODUCTION OBJECTIVES LITERATURE REVIEW: The Ecological Dynamics of the Grassland-Forest Interface Grasslands North American Grasslands The Tallgrass Prairie Middle West Savanna Middle West Forest-Openings Disturbance Fire Fire and Tallgrass Prairie Vegetation Fire and Plant Adaptations Fire and the Soil Seed Bank Fire and Nutrient Cycling Fire, the Litter Layer and Water Relations Climatic Drought Lightning Biotic Bison Non-native Plants Project Goal METHODS Overall Plan Site Descriptions Managed Sites Brown Shale Barrens Cave Creek Limestone Glade Gibbons Creek Sandstone Barrens Wildcat Bluff Limestone Glade Unmanaged Sites Page viii xii \IGMM 10 l3 14 15 20 22 23 25 26 26 27 27 27 28 28 30 3O 31 35 35 36 37 37 37 Berryville Shale Glade Cedar Bluff Sandstone Glade Gyp Williams Sandstone Barrens Pounds Hollow Sandstone Glade Round Bluff Sandstone Glade Sampling Methods Vegetation Composition Vegetation Spatial Patterns Soil Measurements Climatic Information Analytical Methods RESULTS Vegetation Composition Herb Species Composition Categorization of Herb Species Grasses, Forbs, Legumes and Exotics Life History Raunkaier’s Life Forms Categorization by Habitat Site Similarity Woody Species Composition Rock Cover Statistical Comparisons: 1988 and 1993 Vegetation Subhabitats Herb Species Composition Rock, Bryophyte and Lichen Cover Categorization of Herb Species Raunkaier’s Life Forms Herb Species Diversity Tree Seedling Composition Shrub and Sapling Composition Tree Composition Statistical Comparisons Among Subhabitats: 1994 Site Open-Area Estimates Soil Measurements: 1993 and 1994 Climatic Data: 1993 and 1994 DISCUSSION Vegetation Composition Herb Species Composition Statistical Comparisons: 1988 and 1993 Site Similarity Categorization of Herb Species Grasses, Forbs, Legumes and Exotics Life History Raunkaier’s Life Forms Categorization by Habitat Woody Species Composition vi 38 38 38 38 39 39 39 44 46 46 46 51 51 51 58 58 58 58 58 63 68 73 73 78 78 83 9O 9O 90 90 99 106 123 123 123 128 136 I36 I36 140 140 142 142 142 I43 144 144 n w: Rock Cover Vegetation Subhabitats Herb Species Composition Categorization of Herb Species Raunkaier’s Life Forms Herb Species Diversity Woody Species Composition Shade Tolerance Density Species Richness Canopy Measurements Soil Measurements: 1993 and 1994 SUMMARY AND CONCLUSIONS RECOMMENDATIONS APPENDIX A: Species lists for woody and herbaceous taxa from floristic survey of the studied forest-openings in 1993 and 1994 for each site and summary of information relating to species importance for sampling in 1993. APPENDIX B: Maps for each of the studied forest-openings. APPENDIX C: Species lists for woody and herbaceous taxa in five subhabitats of the study sites and summary of relative importance information relating to species for the sampling in 1994. LITERATURE CITED vii 204 223 245 n— 1»..IL.-dr._ i... .r. E LIST OF TABLES Table Page 1. Locations of the nine studied forest-openings in southern Illinois. 31 2. Site ownership and management history. 33 3. Number of 50-m2 plots used in 1988 and 1993 vegetation sampling. 42 4. Site visitation dates, 1993 and 1994. 43 5. Summary of 1994 plot-sampling layout. 44 6. Raunkaier’s life form categories. 47 7. Habitat categories and descriptions used to characterize herbaceous vegetation. 49 8. The total number of species in the overall site and the total number of herb species 52 within plots in 1933 and 1993-94. 9. Relative importance (%) of the three most important herb families in 1988 and 1993. 53 10. Relative importance (%) of the three most important herb species in 1988 and 1993. 54 11. Mean (3: 1 SE) number of herb species (per m2) in 1988 and 1993. 56 12. Mean (i 1 SE) herb cover in 1988 and 1993. 57 13. The percentage of grasses, forbs, legumes and exotic herbs in the study sites in 59 1993-94. 14. Herb life history (%) by site in 1993-94. 60 15. Raunkaier’s life forms (%) for herb species in 1993-94. 61 16. Categorization of 1993-94 site herb species into nine habitat types. 62 17. Jaccard site similarity for herbs and the number of common herb species for the 66 study sites in 1988 and 1993-94. viii Table 18. Similarity of herb composition (Jaccard index) among pairs of sites in 1994. 19. Species common to all study sites in 1993 and 1994. 20. Exotic species encountered at the study sites in 1993 and 1994. 21. Relative importance (%) of the three most important woody species in 1988 and 1993. 22. Mean (:1: 1 SE) canopy cover (%) in 1988 and 1993 via two different methods. 23. Mean (1- 1 SE) number of tree species (per 50 mg) in 1988 and 1993. 24. Mean (:1: 1 SE) cover of exposed rock (as a percentage of total ground surface) in 1988 and 1993. 25. Probability values for the comparison (1988 and 1993) of Poaceae, Asteraceae and rock cover in the opening of the study sites. 26. Probability values for site management comparisons of Poaceae, Asteraceae, canopy and rock cover in the site openings in 1993. 27. Relative importance (%) of the three most important herb species in five subhabitats of the study sites in 1994. 28. Relative importance (%) of the Poaceae family in five subhabitats of the study sites in 1994. 29. Relative importance (%) of the Asteraceae family in five subhabitats of the study sites in 1994. 30. Relative importance (%) of the Cyperaceae family in five subhabitats of the study sites in 1994. 31. Mean (i 1 SE) number of herb species (per ml) in five subhabitats of the study sites in 1994. 32. Mean (3.: 1 SE) cover (%) of herb species in five subhabitats of the study sites in 1994. 33. Mean (i 1 SE) cover of exposed rock (as a percentage of total ground surface) in five subhabitats of the study sites in 1994. 34. Mean (:1: 1 SE) bryophyte cover (%) in five subhabitats of the study sites in 1994. ix Page 67 69 7O 72 74 75 76 77 77 79 81 82 84 85 86 87 88 Table 35. 36. 37. 38. 39. 40. 41. 42. 43. 44. 45. 46. 47. 48. 49. 50. 51. Mean (i 1 SE) lichen cover (%) in five subhabitats of the study sites in 1994. Raunkaier’s life form (%) for herbs in five subhabitats of the study sites in 1994. Simpson and Shannon diversity for herb species in five subhabitats of the study sites in 1994. Relative importance (%) of the three most important species of tree seedlings in five subhabitats of the study sites in 1994. Relative importance (%) of the three most important tree seedling genera in five subhabitats of the study sites in 1994. Mean (i: 1 SE) number of tree seedling species (per 25 m2) in five subhabitats of the study sites in 1994. Density (per 25 m2) of tree seedlings in five subhabitats of the study sites in 1994. Tree seedling shade tolerance in five subhabitats of the study sites in 1994. Relative importance (%) for the three most important shrub and sapling species in five subhabitats of the study sites in 1994. Mean (i 1 SE) number of shrub and sapling species (per 25 m2) in five subhabitats of the study sites in 1994. Density (per 25 m2) of shrubs and saplings in five subhabitats of the study sites in 1994. Shrub and sapling shade tolerance in five subhabitats of the study sites in 1994. Relative importance (%) for the three most important tree species in five subhabitats of the study sites in 1994. Relative importance (%) of the three most important tree genera in five subhabitats of the study sites in 1994. Mean (i 1 SE) number of tree species (per 25 m2) in five subhabitats of the study sites in 1994. Density (per 25 m2) of trees in five subhabitats of the study sites in 1994. Tree shade tolerance in five subhabitats of the study sites in 1994. Page 89 91 93 95 97 100 101 102 104 107 108 109 111 113 116 117 118 Table 52. 53. 54. 55. 56. 57. 58. 59. 60. 61. 62. Mean (1: 1 SE) tree diameter at breast height (cm) in five subhabitats of the study sites in 1994. Mean (i 1 SE) tree height (m) in five subhabitats of the study sites in 1994. Mean (i 1 SE) tree crown diameter (m) in five subhabitats of the study sites in 1994. Mean (i 1 SE) diameter at ground level (cm) of stumps in four subhabitats at Brown Shale Barrens-Managed in 1994. Probability values for within-site comparison of Poaceae, Asteraceae, and rock cover and seedling density in 1994. Area (m2) and greatest continuous cardinal dimension (m) of the site opening in 1994. Soil texture summary for the study sites in 1993. Summary of soil characteristics for the study sites in 1993. Mean (i 1 SE) soil depth (cm) for the study sites in 1988 and 1993. Mean (i 1 SE) soil depth (cm) in five subhabitats of the study sites in 1994. Soil moisture (%) in five subhabitats of the study sites in 1994. xi Page 120 121 122 125 125 126 127 129 130 131 132 LIST OF FIGURES Figure 1. Nine study sites and Carbondale, in southern Illinois. 2. Brown Shale Barrens-Managed in May, 1993. 3. Cave Creek Limestone Glade-Managed in August, 1993. 4. Berryville Shale Glade-Unmanaged in August, 1993. 5. Cedar Bluff Sandstone Glade-Unmanaged in June, 1993. 6. Gyp Williams Sandstone Barrens-Unmanaged in July, 1993. 7. Round Bluff Sandstone Glade-Unmanaged in July, 1993. 8a. Map of Berryville Shale Glade (unmanaged). 8b. Map of Brown Shale Barrens (managed). 8c. Map of Cave Creek Limestone Glade (managed). 8d. Map of Cedar Bluff Sandstone Glade (unmanaged). 8e. Map of Gibbons Creek Sandstone Barrens (managed). 8f. Map of Gyp Williams Sandstone Barrens (unmanaged). 8g. Map of Pounds Hollow Sandstone Glade (unmanaged). 8h. Map of Round Bluff Sandstone Glade (unmanaged). 8i. Map of Wildcat Bluff Limestone Glade (managed). 9. Percent association of the herbaceous flora of the study sites, a railroad right-of-way, a hill prairie, and a hardwood forest to nine habitat types. 10. Total number of seedling, shrub and sapling and tree stumps versus distance (from north to south) along a transect with four subhabitats at Brown Shale Barrens-Managed in 1994. xii Page 32 4O 40 40 4O 4O 40 207 209 211 213 215 217 219 221 64 124 Figure Page 11. Mean daily temperature (deg. C) versus month in southern Illinois for three 133 difi‘erent periods of time. 12. Total precipitation (cm) versus month in southern Illinois for three different 135 periods of time. xiii INTRODUCTION Forest-openings (remnant grassland communities) occur throughout the Midwest in the ecotonal region between the tallgrass prairie and the eastern deciduous forest. Forest-openings, also known as savanna remnants, are considered to be a subclass of temperate zone savanna, of which only 0.02% remains (Nuzzo 1986). Savanna remnants (specifically the barrens subclass) are currently the most endangered terrestrial community in the Midwest (White 1984a,b). Savanna decline has resulted from dramatic increases in development and agriculture and the absence of natural fire since c. 1850 (Nuzzo 1986). Historically, fire maintained openings by eliminating woody saplings (Wright and Bailey 1982) and favoring graminoid species (Gibson 1988). Fires were iginited by lightning strikes, Native Americans, and early Euro-American settlers (Wright and Bailey 1982). Due to a growing interest in these unique communities since the early 1960’s increasing efforts have been made to study forest-openings and to reintroduce fire (Seastedt and Rarnundo 1990). In 1988, 22 forest-openings in southern Illinois were sampled and classified into seven types according to vegetation and substrate characteristics, such as the presence of certain herb indicator species and the percent cover of exposed surface rock (Heikens 1991). At the time of the 1988 study, several sites were noted to appear stable, while several others appeared to be undergoing woody encroachment. Therefore, management recommendations were proposed (Heikens 1991). In the ensuing five years, respective land stewards introduced fire via prescribed burn treatments and the mechanical removal of young woody growth at about half of these sites. 2 In 1993, nine of the original 22 sites were resampled, four of which were managed. Sites were chosen on the basis of permit availability, proximity, management history (treated and control sites) and availability of permanent plots. Vegetation composition was then compared between years (1988 and 1993) in the nine sites. Managed and unmanaged sites (1993) were also compared. Forest-opening perimeters were mapped in 1994 to determine opening size and shape and to document site dimensions for future reference. Also in 1994, north-south transects were sampled for herb and woody species structure and composition. This allowed the comparison of opening vegetation and adjacent forest-interior communities. In this way, species unique to the openings, and often uncommon in southern Illinois, were determined to inhabit several of the sites. OBJECTIVES The principal research goal was to determine the changes in managed and unmanaged forest-openings over time, and to evaluate management efficacy in maintaining the structure and composition of forest-opening communities. To achieve this goal, specific project objectives included the following: 1. describe forest-opening herbaceous vegetation composition (in terms of species richness, percent cover, and relative importance) -describe herb composition categorically (1993) by grouping herb species by life history, habitat preference, life form, and family, —develop vascular plant species lists for all nine sites (encompassing the opening and surrounding 15 m of forest vegetation), 2. determine similarities and differences in composition for managed and unmanaged sites, -compare managed sites (1993) with earlier unmanaged condition (1988) using species richness, percent cover, relative importance, and percent similarity, -compare managed sites with unmanaged counterparts (1993) (as above), 3. document vegetation distribution, -elucidate spatial patterns in the occurrence of herbaceous growth from the opening to the forest interior using species richness, percent cover, relative importance, life form and diversity, -e1ucidate spatial patterns in woody seedlings, shrubs and saplings, and trees from the opening to the forest interior using species richness, density, relative importance and shade tolerance, 3 -map the open area of each site, 4. determine soil influences, -compare soil characteristics (i.e., moisture, texture, nutrients, pH, organic matter, depth) between years (1988 and 1993), elucidate trends in soil moisture and depth from the opening to the forest- interior (1994), 5. define climatic variables (i.e., temperature, precipitation) in 1988, 1993 and the average (1910-1993) for southern Illinois, and 6. develop management recommendations for each site based on the size of the opening, the stage of woody succession and information from the literature review. LITERATURE REVIEW: The Ecological Dynamics of the Grassland-Forest Interface Grasslands The grassland biome is ubiquitous. Covering 24% of the vegetated land surface worldwide (Harlan 195 6), they exist on every continent, from the dense bamboo thickets of the tropics to the arctic plains (Risser et a1. 1981). Grasslands are lacking in beta diversity. For example, while over 2000 plant species were listed for the deciduous forest realm in North America (Bazzaz and Parrish 1982), fewer than 300 were listed for the Great Plains (Weaver and Fitzpatrick 1934). Conversely, point diversity may be high. A limestone grassland in Switzerland contained 40 species per square meter (Andreas and Leutert 1996), whereas a mixed-hardwood forest in north- central Virginia contained only 13 herbs per square meter (Gilliam et a1. 1995). Also, ecotypic variation within formations (e. g., the tallgrass prairie) may be extensive (McMillan 1959). Grasslands are generally located in the interior of large land masses (Risser et a1. 1981) on level to rolling topography (Anderson 1982). Although grassland climates vary widely, they are generally subject to seasonal temperature extremes (e.g., —40 to 43°C in North American tallgrass; Nichols and Entine 1978), alternating between a dry and a wet season, and receive about 25 to 100 cm of annual precipitation (Walter 1973, Risser et a1. 1981, Anderson 1990). According to Transeau (1905), the ratio of annual precipitation (P) to annual potential evapotranspiration (PET) can be used to define a grassland region. Where P is only 20 to 60% of the PET, short-grass or mid-grass prairie occurs. Tall-grass prairie occurs in regions in which the ratio of rainfall to PET is 60 to 80%, oak-forest/oak-savanna where the ratio is 80 to 100%, and deciduous forest, about 100 to 110%. North American Grasslands The origin of the grassland environment in North America dates back 25 million years to the Oligocene Epoch, and the climatic changes caused by the uplift of the Rocky Mountain (Risser et a1. 1981). In its approximate present location, the first extensive grassland formation occurred five to seven million years ago during the transition between the Miocene and Pliocene Epochs (Axelrod 1985). At that time, oceanic chilling and Antarctic ice growth contributed to a drying trend in central North America, reinforced by rainshadow from continued Rocky Mountain uplifi, which restricted forest and facilitated an “explosive” evolution of grasses (Axelrod 1985). During the Pleistocene, 10,000-300,000 years before present (YBP) (Risser et al. 1981), central North America was cooler and moister and predominately wooded, although grassland occurred locally as a forest-grassland mosaic (Risser et a1. 1981, Axelrod 1985). A subsequent warming trend, called the Hypsithennal, began approximately 10,000 YBP with a peak between 7,000 and 8,000 YBP (Anderson 1990). The Hypsithermal, evident from lake sediment cores and paleobotanical data, caused the advance of prairie into boreal and eventually mixed deciduous forest (Anderson 1990). About 11,000 YBP, massive extinctions of large grassland mammals such as the horse, mammoth and ground sloth took place, perhaps owing to the hunting practices of Native Americans (Martin 1975). To facilitate hunting of bison and other animals, aborigines traditionally set fire to North American grasslands beginning at least 10,000 YBP, and thus promoted the advance of prairie to the east (Anderson 1990). For example, the eastward migration of the bison (Bison bison) is largely attributed to anthropogenic fire (Pyne 1983, Hart 1990). Bison crossed the Mississippi River about 1000 AD. and reached Massachussets by the seventeenth century (Roe 1970). Likewise, prairie spread from the base of the Rocky Mountains as far east as present-day Long Island, New York (Blizzard 1931). Although climatic cooling has favored westward expansion of deciduous forest, for the last 5,000 years it has been deterred by fire at the prairie-forest border (Anderson 1990). 7 Prior to Euro-American settlement, grassland was the largest continuous vegetation formation in North America (Risser et a1. 1981), covering one-ninth of the North American continent (Chadwick 1993), and totalling 160 million ha (Chadwick 1995). Spanning approximately 1610 km east-west and 3220 km north-south (Nichols and Entine 1978), the grassland stretched from the central plains of Texas north to the aspen-parkland of Alberta and Saskatchewan and from the Rocky Mountains eastward as a wedge to Illinois, Indiana, and Ohio (Wright and Bailey 1982). Sloping eastward as a catena from the base of the Rocky Mountains at 1829 m above sea level (ASL), the short-grass prairie grades into mid- or mixed-grass at 914 to 1524 m ASL; and mixed-grass prairie grades into tall-grass or true prairie between 274 and 610 m ASL (Wright and Bailey 1982). Short-grasses range in height from 15 to 60 cm, mixed-grasses from 60 to 120 cm, and tall-grasses from 120 to 300 cm (Risser et al. 1981). Annual grassland precipitation increases from 25 cm in the west to 100 cm in the east (Risser et a1. 1981) with a corresponding increase in annual net production of 2 t/ha, 3 t/ha, and 5 t/ha for short-grass, mixed- grass, and tallgrass prairie, respectively (Walter 1973). West to east differences also include an increase in soil organic matter, soil depth, and available nutrients (Bazzaz and Parrish 1982). Given the amount of annual precipitation which occurs in the tallgrass prairie (75 to 100 cm; Risser et al. 1981), a climate-based life zone diagram by Holdridge (1967) (cited in Collins and Gibson 1990) indicates that woody vegetation, not prairie, is clirnatically suited to this region. The tallgrass prairie, therefore, is a dysclirnax to which we now turn our attention for the purposes of this study. The Tallgrass Prairie The tallgrass prairie, or true prairie, differs from the short- and mixed-grass prairie in that it has two peak periods of rainfall (rather than one) and higher plant species richness (Risser et a1. 1981). It is at greater risk of drought than the contiguous northeastern deciduous forest (Risser et 8 a1. 1981) since it does not experience deep soil recharge during periods of extreme drought, such as the great drought of 1933-1934 (Britten and Messenger 1969). Also referred to as the prairie peninsula, it is wedge-shaped in geographic outline, with its base at about the 98th meridian from Manitoba south to Texas, then reaching east to Ohio with remnant disjunct islands or forest- openings scattered throughout the the northeastern deciduous forest (Transeau 1935). Although true prairie once occupied 3% or 575,000 square kilometers of the North American continent, (Knapp and Seastedt 1986), little remains. By 1830, the majority of the true prairie had been settled and cultivated (Risser et a1. 1981). In Illinois, Indiana, and Ohio, 99.9% of the original tallgrass prairie has been eliminated (Chadwick 1995). The largest remaining tract of undisturbed black-soil prairie is a 2 ha pocket in agrarian McLean County, Illinois (Schafer 1990). (“Black-soil” refers to the soil order Mollisol, dominant in temperate grasslands; Foth 1990.) A pristine tallgrass prairie will contain 250 herbaceous species over an area of about 2,600 ha (Risser et a1. 1981), but point diversity is low relative to other grasslands (Collins and Gibson 1990). For example, Peet let a1. (1983) found an average of 18 species per square meter in a mesic tallgrass prairie, whereas Andreas and Leutert (1996) identified 40 species per square meter in a limestone grassland in Switzerland. Ninety five percent of tallgrass indigens are perennials, living up to 20 or more years (Blake 1935, Risser et a1. 1981). Major species of the tallgrass prairie have broad ecological amplitudes and a relatively large geographical range (Risser et a1. 1981). True prairie consists of a matrix of a few dominant warm-season (C4) grasses and many interstitial species (usually C3) (Collins and Gibson 1990). While grasses account for 70 to 98% of ground cover (Lippert and Hopkins 1950), they comprise 10% of the species, composites 26%, legumes 7%, mints 4%, and the Liliaceae 4% (Curtis 1959). Twelve species come into bloom per week from April to September (Chadwick 1993), and 70 at the height of the growing season in June (Walter 1973). Seasonal aboveground biomass production exceeds decomposition by 20% (Golley 9 and Golley 1972). Two-thirds of the prairie biomass occurs belowground (Nichols and Entine 1978), 75% of which is in the top 25 cm of the soil (Risser et a1. 1981). Still, most tallgrass indigens have rooting depths in excess of 1.5 m (Risser et al. 1981). The true prairie is characterized by an association of three dominant genera, i.e., Andropogon-Panicum-Sorghastrum. There are two seral communities which make up 75% of the true prairie, i.e., the Quercus-Andropogon of the Cross Timbers area (Kansas, Texas) and the northern Midwest (Minnesota, North Dakota, Wisconsin) and the Juniperus-Quercus-Sporobolus- Andropogon of Alabama, Arkansas, Michigan, and Tennessee (Risser et a1. 1981). While the prairie-forest interface is “remarkably” abrupt in northern Minnesota and Illinois (Buell and F acey 1960), the transition is a broad macromosaic in southern Minnesota, Iowa, and Illinois (Davis 1977). Middle West Savanna The Midwestern savanna is essentially but “not always” a transitional community between the true prairie and the eastern deciduous forest (N uzzo 1986). Formerly 11,000,000 to 13,000,000 hectares, only 0.02%, or 2, 607 ha remain (Nuzzo 1986, Breining 1993). Today the midwestem savanna (i.e., the barrens subclass) is the most endangered terrestrial community in the Midwest and is listed as globally imperiled (White 1984a,b). The term “savanna” was first used in the sixteenth century by the Taino Indians of the grassy, treeless plains of the Caribbean islands and later by Spanish explorers (Breining 1993). The term applied only to tropical or subtropical grasslands until the 1950’s when ecologists expanded the definition to include temperate plant communities as well (Odum 1953, Oosting 1956, Dyksterhuis 1957). Definitions of Middle West savanna variously describe a grassland sward with “scattered” trees, primarily oak, with nonoverlapping canopies of 10 to 80% (or even 1S 10 90%) cover (Cottam 1949, Haney and Apfelbaum 1990), although 50% represents a suggested upper limit by Nuzzo (1986) and Heikens (1991). The word savanna is sometimes preceeded by the descriptors “oak” and “scrub” to provide additional character information (N uzzo 1986). Savanna trees occur in cohorts with distinct age classes corresponding to precipitation cycles. Each precipitation event may endure for five to 12 years and return every 25 to 50 years. These wet periods may reduce or eliminate fire, and permit the recruitment of Quercus spp. (Haney and Apfelbaum 1990). Tree cohorts usually range between 25 and 250 years in age (Curtis 1959, Haney and Apfelbaum 1990). In Wisconsin, oak savanna herbs had a similarity index of 0.50 to 0.58 relative to typical prairie flora and 0.53 relative to the dry-oak-forest herbaceous flora (Curtis 1959). Bray (195 7) noted that southern Wisconsin prairie harbored both prairie plants and climax forest herbs. In the early 18005 (presettlement), savanna in southern Illinois was an order of magnitude lower than proximate forested areas in tree density, 15.8 trees/ha vs. 159.8 (Anderson and Anderson 1975). Similarly, in Wisconsin, Cottarn (1949) estimated that tree density of a former oak opening had increased from 5.7 trees/ha in 1834 to 57 trees/ha in 1946. Middle West Forest-Openings Savanna remnants (also called forest-openings) occur as relic grassland outliers surrounded by forest; they vary in size from a few square meters (Hanson 1922) to over 65 km2 (McInteer 1942). Although Midwest forest-openings were originally “very extensive” (Braun 1950), they are also “time-transgressive” communities (Heikens and Robertson 1995) known to succeed to closed forest in 10 to 40 years (Schwegman and Anderson 1984, Nuzzo 1986). Accounts after 1825 report the loss of these communities (Sauer 1927). Aerial photographs and research since 1938 document increases in forest herbs and woody species cover and frequency with concommitant decreases in prairie taxa over time (Aldrich et a1. 1982, Heikens and Robertson 1 1 1995) as do forest-opening studies in Wisconsin (Domey and Domey 1989), Ontario (Catling and Catling 1993), Nebraska (Hanson 1922), Missouri (Guyette and McGinnes 1982), Illinois (Robertson and Heikens 1994), Indiana (Bacone and Casebere 1983), Ohio (Hardin 1988) and Kentucky (Baskin and Baskin 1978). The recent literature emphasizes the distinguishing characteristics and classification of forest—openings, their origin and the reasons for their existence. The terms barrens and glades are frequently used to denote certain types of forest-openings. White and Madany (1978) classified barrens and glade forest-openings, also historically known as oak openings, scalds, rock ledges, etc., as savanna subclasses, equivalent to the level of the forest community. Heikens (1991) provides a key for savanna types (and also includes hill prairie and open forest), which separates communities based on vegetation and soil characteristics. For example, barrens have 210% cover of characteristic prairie Species, soil depth 310 to <40 cm, and exposed rock of >1% to 55%. Barrens are grasslands co-dominant with trees, typically Quercus stellata (post oak) or Quercus marilandica (blackjack oak). Furthermore, barrens are identified by substrate type into, chert, shale, and sandstone barrens. The term barrens is known from settlement of the Midwest when pioneers judged local open areas to be too poor to support timber, and therefore sterile and unproductive (Ellesworth 1838). However, the Public Land Survey records of the 1800’s show that barrens occurred on soils that were “good” as well as “poor,” dry and “well-watered” (Hutchison 1982). Although extant barrens usually occur on south to southwest facing slopes (Heikens 1991), they were historically found on all types of topography, including valleys and streamsides (Hutchison 1982, Anderson and Schwegrnan 1991), and on a variety of substrates (Heikens 1991). A common barrens feature includes open grown trees which appear stunted and gnarled. This physiognomy may be due to a combination of moisture deficit, periodic drought, shallow soil and nutrient 12 deficiency (Reich and Hinckley 1980). Barrens communities contain a combination of prairie and woodland herb species, though certain prairie indigens, such as species of Silphium, are generally absent (Vestal 1936). Unglaciated barrens contain a large number of endemic plants. For example, the Mid-Appalachian shale barrens, accessible since the late Tertiary, harbor 18 endemic and six disjunct, near—endemic species (Keener 1983). Forest-openings in Tennessee contain 23 endemics (Baskin and Baskin 1989), and Kentucky, one (Baskin and Baskin 1978). Only two rock outcrop communities in the unglaciated eastern United States are known to have no endemics: the Shawnee Hills of southern Illinois, and the southern Appalachians in western North Carolina and northeastern Georgia (Baskin and Baskin 1988). Barrens are thought to exist for a variety of reasons. Large wintering herds of bison maintained large timber-free areas in Blue Licks, Kentucky and in some cases nearly denuded the land of vegetation (Hutchison et al. 198 6). The excrement from thousands of roosting pigeons killed trees for “several square miles” near Huntington, Indiana (Hutchison et a1. 1986). Drought, oak wilt, tomados, lightning strikes, fire, and isolated edaphic conditions are also factors responsible for local openings in the forest (Hutchison et a1. 1986, Hutchison 1987, Hutchison 1994). Barrens sometimes intergrade with similar habitats known as glades. According to Heikens (1991), however, glades have a greater percentage of exposed rock and shallower soil than barrens. For example, limestone glades contain >10% cover prairie species, 1 to 5% exposed rock, soil depth <10 cm, and scattered Juniperus virginiana and/or Quercus muhlenbergr‘i. Other glades, i.e., sandstone and shale, also have >5% cover exposed rock and soil depth <10 cm, though cover of prairie species is <10% (Heikens 1 991). A study by Jefferies (1987) supports these dichotomies. Mean soil depth in an Arkansas sandstone glade was 5.2 cm, cover of soil and bare rock, 39.6%, and Juniperus virginiana was the most important woody species . 13 Early use of the term glade referred to wet areas (Hutchison et a1. 1986), although in the Midwest they have also come to denote grass-dominated communities that are substrate-controlled (Heikens and Robertson 1995). Rock ledge communities (usually horizontal or broadly rounded bare-rock shelves or slopes at the top of cliffs, less than a meter and up to 18.3 m wide; Winterringer and Vestal 1956) are sometimes considered to be glades. Juniperus virginiana (eastern redcedar) is a common associate of glades, however, dense, even-aged stands have been known to replace some openings since the time of Euro—American settlement (Guyette and McGinnes 1982). Glades often occur on south to southwest facing slopes on shallow, erosive soil with extensive exposed bedrock. Soil water content in summer is frequently below the permanent wilting point (Baskin and Baskin 1988). For example, in a 225 m2 exposed area (albeit in a barrens), of 35,000 seedlings established in April or May, only nine survived by mid-June (Keener 1983). Direct exposure to sun and insolation causes patches of bare, thin soil to heat up 14 to 17°C over air temperature (W interringer and Vestal 195 6, Diboll 1984), and air above the grassland is two to four times as dry as in the surrounding shrubs and woods (Hanson 1922). These phenomena are critical to the development of glade vegetation which consequently blooms in mid and late spring when moisture is available (Harper 1926). Five small cedar glades in Kentucky (glade area not available) contained a total of 148 plant species (Baskin and Baskin 1978). Jeffries (1987) in Arkansas found 76 species in two glades ranging from 0.05 to 0.8 km in area. Baskin and Baskin (1978) suggest that glades act as refugia for prairie flora. However, McCarty and Hassien (1984) report that inconspicuous prairie plants remain in closed woodland understories. Disturbance It has long been recognized that periodic (natural) disturbance events are necessary for the perpetuation of grasslands (T ranseau 1935, Sears 1942, Dyksterhuis 1957, Axelrod 1985). l4 Disturbance has been referred to as a natural component of intrinsic regulatory importance and its role in these communities has been compared to a cyclic occurrence, such as the annual passage of seasons, in which the post-disturbance community resembles the pre—disturbance community, such that it appears stable through time (Loucks et a1. 1985, Collins 1990). This process is known as autosuccession (Loucks et al. 1985) and disturbance is defined as a process which limits “...plant biomass by causing its partial or total destruction (Grime 1979, p. 39).” There are three classes of disturbance in grassland, namely, climatic, pyric, and biotic, all of which vary in size, frequency, and intensity (Malanson 1987). Disturbances historically included fire, drought, windstorrns, grazing, burrowing (Loucks et al. 1985), and local degradative episodes of disease, such as oak wilt (Transeau 1935). Today, these events can be simulated to a limited extent by herbicides, cutting, mowing, conservative grazing and, in some cases, by controlled or prescribed buming. Fire Since the early 1960’s the use of fire for ecosystem management has been accepted by biologists of all disciplines (Wright and Bailey 1982). Its perception as a damaging agent in forests (Miller 1920) and grasslands by scientists (e. g., Weaver and Albertson 1936, Hopkins et a1. 1948) and land stewards (e.g., USDA-Forest Service) has been revised with numerous ecological papers (e.g., the Leopold Report of 1963; Leopold et a1. 1963). Researchers also cite numerous accounts of burning by Native Americans to harvest food (e. g., grain, nuts, fruit), improve forage, drive animals, clear land (for travel, defense, and aggression), and reduce pests such as snakes, flies, and mosquitoes (Pyne 1983, Axelrod 1985). Conflagrations were often expansive and frequent. In 1885 one fire traveled 282 km, another burned an area 32 by 97 km in Texas (Wright and Bailey 1982). Open oak-hickory forests in southern Illinois were burned annually (by Native Americans, then settlers), until 1930 when the Shawnee National Forest was created (Miller 1920, 15 Anderson 1972, DeSelm 1989). Fire frequency was also determined by the roughness of the topography and the presence of fire breaks such as streams or escarpments (Anderson 1990). For example, rock outcrops seldom supported a fire because of their sparse vegetation (Harper 1926). Before settlement, level to rolling topography in the Midwest burned every 5 to 10 years, while dissected topography burned every 20 to 30 (Wright and Bailey 1982). Under natural conditions tallgrass prairie usually burned every two to four years (Aber and Melilo 1991), midgrass every 15 to 30, and shortgrass, not more than every five to 10 years (Wright and Bailey 1982). Peak fire probability is in July and August and secondarily in late spring, although fires can occur at any time of the year (Bragg 1982). Prescribed fires average 102 to 388°C at the soil surface with an extreme range of 83 to 682°C (\N right and Bailey 1982). Sixty degrees Celcius is the standard thermal death point for vegetation (of a given tissue moisture and exposure time, usually about 10 minutes), but grass species (below ground parts) have been known to survive temperatures up to 75°C (Jameson 1961). Dry fuel must reach 346 i 40°C to combust (Wright and Bailey 1982). Over an 18-year period in a Kansas tallgrass prairie an average of 63 to 89% of aboveground biomass (3,090 to 4,350 kg/ha) was removed by combustion (values based on annual burns conducted at four different times of the year) (Ojima et al. 1990). A typical fire will cause the soil to heat to 66 to 79°C at a depth of 0.64 cm for 2 to 4 minutes after passing. Temperature increases below this depth are negligible, regardless of soil texture (Wright and Bailey 1982, Svejcar 1990). Fire and Tallgrass Prairie Vegetation. Pyric events may create patches of bare soil which, during the growing season, heat up 2 to 17°C over surrounding air temperatures (10°C average) (Kucera and Ehrenreich 1962, Wright and Bailey 1982). High soil temperatures stimulate microbial activity, decomposition of organic matter, and nitrogen mineralization (Ojima et al. 1990). After fire, the amount of photosynthetically active radiation reaching emerging shoots 16 increases by 60% (Knapp 1984) and regrowth is therefore precocious (Svejcar 1990). Herb canopy closure may be complete two to three weeks after fire (Eisele et al. 1989, Svejcar 1990). In addition, phenological development is earlier (Bazzaz and Parrish 1982), accompanied by as much as a 60% increase in plant height (Curtis and Partch 1950). Soil moisture is also depleted earlier in the season, causing more rapid vegetation senescence and a reduction of live aboveground biomass to levels comparable to unburned prairie by the end of the growing season (Svejcar 1990). Most prescribed fires are set in spring, between late March and April (Benning and Bragg 1993). “Early spring” burns occur around 20 March and 10 April, “late spring” around 1 May (Benning and Bragg 1993). These burns typically favor “warm-season” C4 species (blooming between July and October) (Howe 1994) and stimulate large increases in cover (Kucera and Koelling 1964, Towne and Owensby 1984), productivity (Svejcar 1990), and, for grasses, tiller number (Svejcar 1990), flowering stems, and caryopse number (Glenn-Lewin et al. 1990). However, the response of a given species may vary across its geographic range (Svejcar 1990). For example, the flowering increase of Andropogon gerardir‘ (big bluestem) ranged from 54 to 3780% at sites in Wisconsin, Illinois, and Iowa (Glenn-Lewin et al. 1990). Furthermore, a species may be an “increaser” at one site and a “decreaser” at another. Sorghastrum nurans (Indian grass) varied from a 663% increase to a 79% decrease in flowering-stem number among burned Wisconsin, Illinois, and Iowa sites (Glenn-Lewin et al. 1990). Moisture regime also contributes to species response to fire. After a dry-year burn in spring, the net productivity of Schizachyrr'um scoparr‘um (little bluestem) was 58% lower than in unburned plots (Hopkins et al. 1948); but in wet-year burn plots it was 81% above the control in mixed-grass prairie (Wink and Wright 1973). Towne and Owensby (1984) in a 56-year study of annually burned tallgrass prairie in Kansas found that a three-week difference in the timing of a spring burn resulted in significant differences in herbage yield and species composition. Consistently, in a re-established tallgrass 17 prairie in Nebraska, Benning and Bragg (1993) concluded that a difference of four days in spring burning determined whether significant increase for both flowering stern number and flowering stem height of Andropogon gerardr‘i was observed. Significant plant response occurred only after 12 May (until 20 May): 8 to 10 days after initiation of plant growth. The ratio of C3 to C4 plants decreases following late spring burns (Towne and Knapp 1996). Late spring burning destroys C3 grass and forb shoots at a period of maximum growth, yet before the initiation of leaf expansion of C4 species, thereby favoring the competitive superiority of C4 plants (Howe 1995). The majority of C4 species in a tallgrass prairie are grasses while most of the C3 plants are forbs (Dickinson and Dodd 1976). Towne and Owensby (1984) found forb yield to be highest and grass yield lowest on unburned prairie plots versus annually burned treatment plots lit in winter, early, mid, or late spring. After 12 years of protection from fire on tallgrass prairie in Missouri, Zimmerman and Kucera (1977) noted large populations of perennial dicots, especially Solidago spp. (goldenrods). Hartnett (1991), studying the tallgrass prairie forb Raribr'da columnifera (prairie coneflower), discovered that plants from sites not burned for many years were 2.6 times larger and produced 50% more stems than counterparts from recently burned sites. Therefore, Svejcar (1990) suggests timing a prescribed burn with the phenological stage of a key prairie species. However, relatively few studies document forb response to fire, although most results are species-specific (Glenn-Lewin et al. 1990). Transitory (one to two year) effects of fire usually include a doubling in tiller number and total aboveground biomass, a variable but positive effect on flowering stem and caryopse number, and a positive second year effect on herbaceous seedling establishment (Glenn-Lewin et al. 1990). Still, exceptions to each of the above trends are reported and sources of variability include site history, species composition, moisture regime, geographic location, timing of burn, and plant growth stage (Svejcar 1990). 18 The majority of studies report an increase in species richness the year after a burn (Collins and Gibson 1990), especially when fire is periodic or absent for ten or more years (Kucera and Koelling 1964). Patterns in richness the year a fire is set are not clear and often correlate with precipitation (Collins and Gibson 1990). However, a second-year effect is often observed since fire stimulates flowering, and establishment results from recently dispersed seed (Rabinowitz and Rapp 1985). Also, where few seedlings are reported in undisturbed prairie (Blake 1935, Goldberg and Werner 1983), fire creates patches of bare soil favorable to colonization (Rabinowitz and Rapp 1985). In mesic grasslands, variations in species richness are primarily a function of the number of forbs (Blankenspoor 1987). However, annual spring burns increase dominance by the matrix grass species such that richness is lower than in unburned prairie, in addition to depleting the soil seed pool (Collins and Gibson 1990). Dormant-season burns have little effect on species richness (and there are few, if any, winter annuals; Collins & Uno 1983) (Collins and Gibson 1990). According to Anderson and Schwegrnan (1991) species diversity is often greatest in ecotones, e.g., prairie to forest. For example, in a mesic southern Illinois barrens, Anderson and Schwegrnan (1991) found prebum species richness to be lowest, increasing the first two years after fire to a peak 15 years later, when shade tolerant forest herbs and woody species were rapidly invading. At Buffalo Beats prairie in southeastern Ohio, herbaceous species richness likewise increased after fire with forest encroachment (Hardin 198 8). Although characteristic prairie species such as Liam's scariosa, Desmodr‘um paniculatum, and Lespedeza repens no longer occurred in unmanaged plots after 22 years, 18 species were new to the prairie opening, six of which were formerly found in the transition zone or forest-opening samples alone. The coefficient of similarity between the former transition area and forest interior was nearly identical. Late spring burns, particularly on an annual basis, also lower species diversity and 19 community heterogeneity (Collins and Gibson 1990). A model by Gibson and Hulbert (1987) shows a gradual increase in species diversity after fire for six to seven years, after which it declines. This inversely correlates with a one to two year peak in the cover and productivity of grasses (and annuals for one year) following fire after which cover and productivity decrease (Risser et al. 1981, Collins and Gibson 1990). For example, following fire in sown swards of Andropogon gerardr‘r‘ and Sorghastrum nutans, productivity was three to four times higher than unburned plots (Hadley and Kieckhefer 1963). After 22 years of postfire succession in southeastern Ohio, cover of Andropogon gerardir' decreased from 50 to 16% (Hardin 1988). Eventually, without periodic fire, woody species dominate prairie sites (Anderson 1983, Anderson and Schwegrnan 1991, Heikens and Robertson 1994). Haney and Apfelbaum (1990) noted the release of oak from rootstocks of unknown age in former oak savanna (now closed forest) following fire cessation in the upper Midwest. The number of trees in unburned tallgrass prairie increased 60% over five years in northeastern Kansas (Briggs and Gibson 1992). Bragg and Hulbert (1976) reported an increase of 40% in woody plant cover in unburned tallgrass prairie over a 30-year period in Kansas. The most obvious change in an oak opening is recruitment to sapling and tree layers, whereas tree seedling numbers decrease over time (Bragg and Hulbert 1976). In a Minnesota savanna, tree (_>_10 cm dbh) recruitment was 5 to 45 stems/ha over a five year period. Over the course of a major drought, the unburned plots had a larger percentage of shrub or sapling stems (plot size=0.375 ha), 58 verus 48%, respectively (Faber-Langendoen and Tester 1993). Between 1834 and 1946 an oak opening in southwestem Wisconsin showed an increase in the frequency of Quercus alba (white oak) and Quercus velutr’na (black oak) of 37 to 83% and 20 to 53%, respectively, while the shade-intermediate pioneer species Quercus macrocarpa (bur oak), declined from 72 to 8% (Cottam 1949). Concurrently, the understory frequency of prairie grasses also declined from 57 to 0%, C eanorhus sp. (redroot) 59 to 14%, and Silphium terebinthr’naceum (prairie dock) from 5 to 0%. 20 Fire and Plant Adaptations. Although tallgrass prairie may quickly succeed to woody vegetation, fire kills or retards woody growth (Bragg and Hulbert 1976, Wright and Bailey 1982, Abrams and Hulbert 1987, Briggs and Gibson 1992). Tree mortality often results not from cambial damage but from root injury and canopy scorching from hot gases (Spurr and Barnes 1992). Fire-sensitive species such as Juniperus virginiana (eastern redcedar) have shallow roots and thin bark (Arend 1950) and are impeded by a fire frequency less than 20 to 30 years (Wright and Bailey 1982). Trees with bark thickness in excess of one centimeter experience little heat damage (Wright and Bailey 1982). Oak, a dominant savanna tree, experiences <4% mortality when not stressed by drought (Faber-Langendoen and Tester 1993). However, losses from oak wilt in the red-oak group exceeded 20% within five years of a fire in an oak woodland in southern Wisconsin (McCune and Cottam 1985). With windstorrn and drought in southeast Texas, oak mortality exceeded 50% (Glitzenstein and Harcombe 1988). But where fire destabilized closed forest in central Illinois, causing 47.6% mortality within three years of a burn, nearby savanna trees experienced no damage (Anderson and Brown 1983). Anderson and Brown (1983) determined that the survival of savanna trees was due to ground layer shading, which restricted herbaceous growth, and to wind action, which removed basal litter, thereby preventing fire from reaching within 30 cm of the base of any savanna tree. Ko and Reich (1993) concur. Although soil moisture, nutrient and organic matter levels were higher beneath savanna oak, total aboveground biomass was 50 to 100% lower than in uncanopied areas. Fire-prone environments harbor a higher proportion of resprouting woody species than non-fire environments (Parker and Kelly 1989). One hundred percent of woody species in California sage (Malanson and Westrnan 1985), 65% in the fynbos of South Africa (Kruger 1977, as cited in Trabaud 1987), and 50% of Califomia sclerophyllous scrub (Mooney and Dunn 1972) are capable of resprouting. Trees regenerate by root suckers, stump sprouts, basal burls (basal stem swellings developed around stem wounds) (Lacey and Johnston 1990), and grubs (oak and 21 hickory sprouts killed by annual fires). Oaks are notorious resprouters (e.g., Quercus macrocarpa, Q. stellata) (Burns and Honkala 1990). In 1913 John Muir noted the presence of grubs about 100 years old in Wisconsin oak savanna (Cottam 1949). Liming and Johnston (1944) discovered sprouts (about 4 years old) from oak “stools” (enlarged callus-like structures at groundline formed from repeated sprout mortality due to periodic fires) in annually burned oak- hickory forest of the Missouri Ozarks. The root systems of these sprouts averaged 23 .9 years in age. Likewise, oak seedling sprouts from southeastern Ohio grew from rootstocks up to 37 years old (Merz and Boyce 195 6). Grassland herbs have been refered to as pyrophytes, particularly in Europe, but the term has recently been deemed “ambiguous” and “inappropriate” as propagative strategies allowing plants to succeed in fire-prone environments are difficult to distinguish from traits allowing regeneration in response to other disturbances such as drought or grazing (Trabaud 1987). Still, many grassland herbs have prepagative traits advantageous to regeneration after fire, grazing, and drought. For example, many species are rhizomatous. Rhizomes are underground stems that can serve as storage organs and sites of water and nutrient absorption, regeneration (via shoots), and anchorage (Risser et a1. 1981). They occur approximately 2.5 cm belowground (for grasses) (Wright and Bailey 1982), safe from the scorching effects of fire, and may be 5 to 10 mm in diameter, and 2 to 4 m long (Risser et al. 1981). Similarly, the depth of subterranean plant organs was found to be species-specific in understory herbs in Acadia forest, New Brunswick (Flinn and Wein 1977). Forbs frequently root deeply, e.g., Amorpha canescens and Liam's punctata roots may exceed 5 m (Weaver and Darland 1949). In general, after fire, root and rhizome biomass increases 50% or less (Svejcar 1990). Seastedt and Rarnundo (1990) found that root length under litter (unburned prairie) was 70% that without litter. Vesicular-arbuscular- (VA) mycorrhizae, a type of endo-mycorrhizae, generally confer a 22 large advantage to C4 grasses (Hartnett et al. 1994). These fungi are symbionts, ubiquitous in tallgrass prairie, which aid a plant in disease resistance, nutrient and water uptake, growth, and interplant linkage (Gibson and Hetrick 1988). With an April burn in Kansas, flowering and stem density of Andropogon gerardir' and Sorghastrum nutans was significantly higher with mycorrhizae than without. C3 grasses and forbs are generally facultative, and show smaller growth responses than C4 grasses (Hartnett et a1. 1994). Fire and the Soil Seed Bank. The primary mode of reproduction in grasslands is vegetative (Abrams 1988, Keddy et al. 1989). Vegetative propagation is, however, costly in terms of energy expenditure per reproductive unit and consequently, the number of propagules is limited (Fenner 1985). The production of a large number of seeds maximizes the potential for dispersal and likelihood of reaching “safe sites” or uncolonized patches (Parker et a1. 1989). This strategy is employed by sparse grasses. Rabinowitz (1978) determined that rare or sparse prairie grasses like Sphenopholis obtusata are light-seeded, 0.06-1.76 mg, and common grasses like Andropogon gerardii are heavy—seeded, 223-2.8 mg. Small seeds are able to germinate more quickly than large seeds and to subsequently preempt patches (Hull 1973, Rabinowitz 1978). Likewise, prodigious seed output allows annuals to heavily stock the seed bank. Seed bank dominion is also based upon seed persistence or the ability to maintain viability over long periods of time (F enner 1985, Levin 1990). This phenomenon has been described as the persistent stage in the life cycle of an otherwise transient species (Parker et al. 1989). Correspondence of species between the aboveground flora and soil seed bank may be due to a rapid turnover of the seed bank subsequent to disturbance. Fenner (1985), Hartnett and Richardson (1989) and Roberts and Vankat (1991) assert that the more frequently a habitat is disturbed, the more closely the species composition of the soil seed bank will resemble the extant vegetation. However, reports of highest consonance between aboveground vegetation and the seed 23 bank come from a regularly burned Chaparral (fire interval 20 years) (Wright and Bailey 1982) which shows an overlap of 50% (Parker and Kelly 1989), not from the tallgrass prairie (fire interval 2-5 years). Most studies report that seed bank species composition is not representative of the existing vegetation (Rabinowitz 1981, Johnson and Anderson 1986, Schiffrnan and Johnson 1992). A seed bank study of a tallgrass prairie (burned every four years) by Rabinowitz and Rapp (1980) confirms this assertion. While the seed bank contained 30 species, a floristic survey named 82 flowering plants. Twenty one species contributed 7.8% of the total seed while the remaining nine contributed 92.2%. The dominants of the site, Andropogon gerardii, Schizachyrr'um scoparium, and Vernom’a baldwim’i, were absent from the seed bank. Fire and Nutrient Cycling. Fire in tallgrass prairie accelerates the oxidative process of organic matter decay (Harvey et al. 1976, cited in Wright and Bailey 1982), volatilizing nitrogen (N) and sulfur, and depositing cations in ash (Wright and Bailey 1982). Ash leachate may stimulate seed germination, as in the Chaparral (Keeley 1987). The percolation of cations in soil has been shown to increase root depth and the equitability of root distribution in the soil horizons (Aber and Melilo 1991). Cations percolate through the soil following rain, displacing hydrogen ions, and raising soil pH slightly in the upper 1 to 10 cm, e.g., from 5.87 to 6.07 (Owensby and Wyrill 1973), for a period of one or two years (Raison 1979). The tallgrass prairie, like most terrestrial ecosystems, is N-limited (Seastedt and Rarnundo 1990). In a typical fire (200°C), over 90% of N in aboveground plant material is volatilized (Wright and Bailey 1982), resulting in a loss of about 1-2 g/m2 (Ojima ct al. 1990). However, postfire vegetation immediately experiences a marked increase in production (Knapp 1985, Svejcar and Browning 1988). With annual burning, postfire productivity is also sustained over time. After 10 years of annual burning at a site with 17% composition Andropogon gerardii, productivity was 24 15 to 20% higher than that of unburned prairie (T owne and Owensby 1984). In Kansas, aboveground productivity was still 30% higher than the unburned control after 18 years of annual burning (Towne and Owensby 1984). These effects are associated with immediate rrricroclirnatic change, and greater plant efficiency (more biomass per g N) (Ojima et al. 1990). Longtenn annual (30 yr) and periodic (4 yr) burns significantly reduced extractable ammonium in the top 5 cm of soil (Vance and Henderson 1984) and soil arrunonium concentration was shown to recover only partially 50 years after a single fire in a Finland forest (V iro 1974). After fire, grasses obtain about 7% of N from bulk precipitation, 7% from mineralization of organic matter, and <17% from atmospheric N-fixation (Sclesinger 1991). The rest is thought to come from retranslocation and root decay (Abbadie et a1. 1992). The N budget for an annually burned prairie has been calculated by Seastedt (1985). One to two grams of N m’zyr" enter via precipitation, 30% of which is absorbed by microbes on standing dead vegetation. Peak live aboveground vegetation uses 4 g N m'zyr", with a 25% turnover, and 1-2 g N m'zyr‘l are volatilized while 05-07 g N m‘zyr'l are deposited on the soil via ash and unburned debris (Ojima et al. unpublished, in Seastedt 1988). Roots use 5-6 g N m'zyr". Live roots have a 30-40% tumover rate . Longtenn burning may favor N-fixation. Additions of ash following fire provide inorganic phosphorus (P) in amounts comparable to those bound organically in the original standing material. Phosphorus stimulates N-fixation by terrestrial cyanobacteria (i.e., Nostoc muscorum) and litter removal increases soil temperature favorable for algal growth until canopy closure (about three weeks) (Eisele et a1. 1989). N-fixing legumes may also increase in number following fire. For example, after 15 years of annual late spring fires, legume density was significantly higher (8 i 1 stems/m2) than in unburned plots (3.0 i 0.3 stems/m2), although total biomass was not significantly different (Towne 25 and Knapp 1996). Unlike most forbs, Towne and Knapp (1996) discovered that legume biomass increased from about 11% to about 25% after 10 years of annual burning. Anderson and VanValkenburg (1977) found net density of legumes increased from 17,554 to 63,320 stems/ha following fire in a successional southern Illinois forest-opening and legume production was at least seven times greater on burned than on unburned plots. Large increases in the exotic legumes, Lespedeza striata and L. stipulacea (from 0 to 2,366 and 486 to 4,364 stems/ha, respectively) were found subsequent to a forest-opening burn in southern Illinois (Van Valkenburg 1977). Thompson and Heineke (1977) also found significant increases in frequency of the exotic legumes, L. stipulacea and L. striata, as well as Coronilla varia, Glycine max, Lespea’eza cuneata, Medicago lupulina and M sativa along periodically burned railroad right-of-ways in southern Illinois while Diboll (1984) noted an increase in the European legume T rifolium repens following an experimental prairie burn in east-central Wisconsin. Martin et al. (1975) found native Cassia, Desmodium, and exotic Lespedeza spp. to increase in bumed—over forest cuts in the Piedmont (Virginia to Georgia). Fire, the Litter Layer and Water Relations. Longtenn unburned prairie is said to be energy- lirnited as standing dead plant material and litter reduce usable solar inputs significantly relative to burned prairie (Seastedt and Rarnundo 1990). Seastedt (1988) suggests a 1-2 year lag time between foliage production and litter deposition on a burned site. Consequently, large increases in production are followed by litter accumulation about three years postfire (Wright and Bailey 1982). An unburned prairie planting in Wisconsin (planted and left untreated for six years) bore thatch with 40.0 i 9.4% cover, 10-40 cm deep, while a site bumed in April bore 29.0 i 6.0% cover thatch, 2 to 10 cm deep, the following June. Although litter does not release allelochemics (Rice and Parenti 1978), decreases in production with time are widely attributed to a layer of thatch. For example, Curtis and Partch (1950) found that the most important factor affecting 26 flowering of Andropogon gerardii was the presence of litter. Burned plots which were recovered with litter were not significantly different from controls in the number of flowering stems, basal area per clump, or average height of flowering stems whereas those which were not recovered following a fire had significantly taller (60%) flowering stems and greater production (six times greater) of flowers than the control. With litter removal and the release from light limitation, increased plant growth decreases soil moisture for one to two months after fire (Ojima et a1. 1990). Soil moisture losses in summer can be greater in tallgrass prairie than under woodland canopy (Kucera 1952). For example, in spring, Minnesota savanna soil moisture was intermediate between a mesic oak woods and xeric prairie (Ovington et a1. 1963). Beneath savanna trees in Wisconsin, soil moisture was significantly higher at the 5-30 cm depth than surrounding open savanna during dry periods. It was, however, similar after a period of rain, despite a 33% reduction of rain under tree canopies (Ko and Reich 1993). Climatic Drought. Drought, like fire, is an important environmental control of woody species. Transeau (1935) recounted the great drought of 1913-1914 in which thousands of trees along the prairie- forest border died. Albertson and Weaver (1945) documented 30 to 93% mortality of native deciduous trees (Ulmus spp., Fraxinus spp., Celtis occidentalr’s), 35 to 80% for Juniperus virginiana from Oklahoma to Nebraska in hedgerows, timberbelts, and the like under the drought conditions of the 19305. Hanson (1922) attributed the presence of dead Quercus macrocarpa saplings and shrubs (0.3-1.5 m high) in prairie inclusions in Nebraska to a past xerophytic period. Walter (1973) noted that the effects of the 1934-1941 drought were still evident in 1953, in so far as recurrent drought every century was partially responsible for the treeless prairie. Furthermore, woodlands had lower tree mortality rates than open savanna during drought with 21.4 and 6.1% for 27 Quercus ellipsoidalis and Quercus macrocarpa, respectively (Faber-Langendoen and Tester 1993). Lightning. From the tropical forest to the tundra, lightning is an important source of fire ignition (Wright and Bailey 1982, Trabaud 1987, Hart 1990). Ten thousand wild fires occur in the United States each year, 80% of them in the Rocky Mountain and Pacific coast states (Spurr and Barnes 1992). Lightning without precipitation is less common in the eastern reaches of the tallgrass prairie (Howe 1994). A study in the northern Great Plains mixed-grass prairie (Montana, North Dakota, and South Dakota) determined that 293 lightning-caused fires between 1940 and 1981 averaged 10.8 ha in area, even though most were suppressed (Higgins 1984). Higgins (1984) deduced a fire frequency of 6.0 yr '1 10,000 km'z, with 73% of fire events occurring in July and August. Eighty-eight percent of lightning fires burned 3.64 ha or less and strikes were most common on top of buttes. Howe (1994) attributed dormant season and spring burns to anthropogenic sources. He fiirther demonstrated that rnidseason burns were important in increasing the C32C4 ratio, species diversity and community heterogeneity in prairie. When lumping species into flowering guilds, the late-flowering dominants (flowering between July and October) had 47% cover after a mid-July burn in a tallgrass prairie planting in Wisconsin, 92% after a March burn, and 80% on the control while perennials flowering before mid-July showed 46%, 6%, and 17% cover, respectively. Biotic Bison. Historically, the bison-grassland relationship was significant. Risser et al. (1981) estimated a pre—Euroamerican settlement population of 50 to 125 million bison (Bison bison). Like mid-season burns, these large native ungulates reduce dominance of matrix (graminoid) species. By preferentially grazing graminoids instead of forbs, they enhance species diversity (Collins and 28 Gibson 1990). Bison were observed three times more frequently on watersheds dominated by C4 grasses that were burned in spring than in unburned areas (V inton et al. 1993). Light grazing removed about 15% of aboveground material (Collins and Barber 1985), moderate, 45%, and heavy, 77% (Shariff et al. 1994). Laboratory-simulated herbivory showed that the grass genus Zea can compensate for up to 50% of tissue loss (Dyer et a1. 1982). And Vickery (1972) (cited in Dyer et al. 1982) found that pasture net primary production under light sheep stocking was 40% higher than without. Bison also created wallows (concave depressions 3-5 m in diameter) which disrupted matrix species (Collins and Barber 1985). Non-native Plants. Non-native plants such as Melilotus spp. (sweetclover), Alliaria petiolata (garlic mustard), Rhamnus spp. (buckthom), and Lonicera spp. (shrubby honeysuckle) are known to overtake Midwestern forest-opening remnants, leading to the demise of many prairie indigens (Haney and Apfelbaum 1990). Drew (1947) called the presence of the exotic legume Melilorus spp. in north-central Missouri prairie “devastating.” Likewise, Melilorus has threatened to overtake portions of Simpson barrens in southern Illinois, despite periodic fires (A. Biagi, pers. observation). Heitlinger (1975) reported the invasion and takeover of degraded prairies by Melilotus alba in the absence of fire disturbance. Anderson and Schwegman (1971) reported a decrease in the exotic vine, Lonicerajaponica (Japanese honeysuckle), after a spring burn which proceeded after L. japonica leaf emergence in a mesic southern Illinois barrens. The prolific Eurasian plant, Alliaria petiolata has recently found its way to southern Illinois along railroad right-of-ways (A. Biagi, pers. observation). It is known to form dense thickets in Midwestem savannas, and to green-up early in the season and remain so well into the fall (Packard 1990). Project Goal Given the loss of savanna remnants throughout the Midwest, their imperiled nature and novelty to ecological research until very recent times, critical questions concerning the 29 requirements for their persistence, particularly the role of fire, stand to be addressed. Management (cutting and burning) of forest-openings in southern Illinois since a 1988 pre-management reconnaissance has presented the opportunity to compare pre- and post-management and control site vegetation over time. Mapping of site boundaries and transect plotting was done to provide further indication of the effect of fire in the comparison of managed and unmanaged site opening, transition and forest interior vegetation subhabitats. METHODS Overall Plan In 1993, nine of 22 forest-openings examined by Heikens (1991) in 1988 were chosen for study (Figure l and Table l). The study was limited to nine sites because of time and logistical constraints; site selection was based on the factors described below. In the five years following the 1988 vegetation sampling of Heikens (1991), four of the nine sites were managed via prescribed buming and the mechanical removal of woody vegetation. In the present study the sites were resampled in order to determine the effects of management, as well as the nature of changes occurring due to purely natural processes. Random-plot sampling in 1993 was facilitated by the permanent plot stakes installed by A. Heikens in 1989 at four of the nine study sites. A comparison was made between managed and unmanaged sites in 1993, especially between sites which were of the same substrate and classification type (e.g., barrens, glades), and were close in proximity. Site variables for all nine sites were also compared to their former (198 8) condition. Sampling in 1994 was conducted using north-south transects which spanned the opening area and extended into the forest-interior. F crest-interior vegetation was compared with opening vegetation and north-south gradients were examined for vegetation patterns. In 1994 the dimensions of the nine forest-openings were mapped using canopy cover and dbh limits to distinguish the opening from the forest. Research permission, as well as records documenting site management history (Table 2), were obtained from respective land owners, viz., the Shawnee National Forest-Forest Service, The Nature Conservancy, and the Illinois Department of Natural Resources (formerly the Illinois 30 31 Table 1. Locations of the nine studied forest-openings in southern Illinois. Site County Quadrangle Township Range Section Managed: Brown Union Jonesboro 12$ 2W 23 (N 1/2 NW1/4 NE1/4) Cave Johnson Kamak 13S 3E 28 (SE1/4 NWl/4) Gibbons Pope Herod 1 18 7E 4 (SI/2 NW1/4 SW1/4) Wildcat Johnson Kamak 13S 2E 24 (S 1/2 NW1/4) Unmanaged: Berryville Union Jonesboro 12$ 2W 26 (W 1/2 NE1/4 NEl/4) Cedar Johnson Lick Creek 115 2E 31 (N 1/2 SW1/4 NEl/4) Gyp Pope Herod 1 IS 7E 17 (NW 1/4 NEl/4) Pounds Gallatin Karbers Ridge 108 SE 36 (N 1/2 SW1/4) Round Johnson Goreville 1 13 2E 27 (SI/2 NEl/4 SW1/4) Berryville=Berryville Shale Glade, Brown=Brown Shale Barrens, Cave=Cave Creek Limestone Glade, Cedar=Cedar Bluff Sandstone Glade, Gibbons=Gibbons Creek Sandstone Barrens, Gyp=Gyp Williams Sandstone Barrens, Pounds=Pounds Hollow Sandstone Glade, Round=Round Bluff Sandstone Glade, Wildcat=Wildcat Bluff Limestone Glade. Department of Conservation). Site Descriptions The study sites are located in southern Illinois on rolling topography known as the Shawnee Hills (Mohlenbrock 1982). The average growing season is 200 days in length, with a continental (cool winter, warm summer) climate (Mohlenbrock 1982). All nine forest-opening study sites are on slopes of 20 to 50% inclination (Heikens 1991) at elevations of 122 to 229 m. In fact, four are delimited by sheer bluffs and two others by ledges or steep slopes. All openings are irregular in outline and have an aspect of south or west. As described by Heikens (1991), the glade type forest—opening has >5% cover of exposed rock, soil depth <10 cm, and canopy cover <80%. Conversely, barrens have <5% rock cover and soil is 10 to 40 cm in depth (Heikens 1991). In general, forest-opening canopy cover does not exceed 80%, and Heikens (1991) suggests that a canopy cover of 50% is probably the best 32 NORTH Corbondole ® Pounds ollow GibbonsOCreek . . Round Bluff Gyp Willioms . Brown . Cedar Bluff 0 Berryville Wildcat Bluff 0 Cove Creek 1.0 r-Ur Ur 315 515 Kilometers Scole Figure 1. Nine study sites ond Corbondole, in southern Illinois. O—Monoged sites, 0 —Unmonoged sites. 33 cum—O 253254 $25 Laoozzznug .250 ocoomncam tEm vasomnmzx doe—O ocoomocam 30:21— mpcsomnman. .mcotam ocozmocam mEEEB Q>OHQ>O £8:sz ocoaocmm x620 muonflOumHO .820 ocean—Em tam EBUHMDU don—D ocoomoEfi x020 e>aUn>>temnu~>m 63383 oo2n<2 838m $20,; 33% oBEDHmLmD .xocatomcou 2:62 eckuuzrr £853.32 .8252 he EoEtameQ 4.6553229 mZA= mmmD mmmD mmmD mze 02m mam QED ~50 A>m ”82w oomacaEcb .EEE m. E NEE m. .3m-m exemv Doom BEoeoE esow oQoQS .EEE m E v; ”(E v .3m-m some 06: Lo; esco— wwtkm .EEE N E +w-_ mE _ .m came Doom EstooE osoo $2me mze 04>» <2 <2 <2 <2 <2 <2 <2 voE—R <2 <2 <2 <2 <2 <2 <2 mm: m: _ 02,—. BO .EEE m E N; mE m .m 6605 Dow: BEeeoE 66mm $23k GQQNC am? a mafia 8:. ES mo E _ ”it v .322 6605 Doc _ooU esov Eta—R N229 >25 m. Transects in which this additional data on trees and soil were taken are listed under “subsample transect” in Table 5; the transect numbers correspond to those depicted in Appendix B, Figures 8a-8i. At Brown-MGD, density and diameter of stumps at ground level were also recorded in the subsample transect. Herbaceous vegetation, bryophytes and lichens were sampled in noncontiguous l-m2 nested plots placed interior to the middle of the 5 m2 western transect edge. Dominance was recorded using the cover classes according to Menges et al. (1987). Exposed rock was measured in the same way as herbaceous vegetation at seven of nine forest-openings. In 1994 all nine study sites were mapped using the previously described “opening” boundaries (Appendix B, Figures 8a-8i). Site maps are arranged on grids with 5 m between vertices. Scale and orientation is provided in the legend. Heikens’s (1991) permanent plots, marker trees, and other significant features are provided for orientation. The points marking the perimeter of the opening (estimated as the point at which a canopy cover of 375% and a tree size of 36.6 cm dbh are reached) are represented with “XXX.” Where no estimation symbols or “establish ” bounds are given, the connections were interpolated. Significant features including bluff edges, rock shelves, thickets, footpaths, fences, roads and streams were originally located 46 relative to 1-7 arbitrary east-west baselines with a compass accurate to one degree. The maps are depicted in plan view (from above looking down) and were generated using AutoCAD, Version 13. Soil Measurements In 1993, following Heikens (1991), soil probes from five randomly chosen locations at each site were combined, dried, mixed, and then sent to A & L Laboratory, Memphis, Tennessee where the following analyses were performed: pH, buffer pH, estimated nitrogen release, percent organic matter, cation exchange capacity, texture (percent sand, silt, and clay), and parts per million of phosphorus, potassium, calcium, magnesium, manganese, and zinc. The probes extended to the maximum depth penetrable (532 cm in 1993). Soil depth was measured for each probe. In 1994, samples were drawn from the mid-opening, forest-edge boundary and forest- interior (15 m beyond forest-edge boundary) of the subsample transect for eight of the nine forest- openings. Samples were immediately sealed in air-tight containers and subsequently weighed, dried, then reweighed to determine percent soil moisture. Soil depth was again measured for the first five probes. Climatic Information The following clirnatic information was obtained from the Midwestern Climate Center, Champaign, Illinois for Carbondale, Illinois (590 km from the study sites), for the years 1910 through 1994: monthly precipitation (with water equivalent for snow included), monthly snowfall and monthly average temperature. Analytical Methods Vegetation composition in 1993 was assessed in a variety of ways. As in Heikens (1991), the rnidpoints of the species cover classes were used for calculations. The “trace” cover class received a value of 0. l. Dominance consisted of the total of areal coverage values divided by the 47 area sampled (Cox 1990). Relative Importance (R1) was computed by dividing the sum of relative dominance or relative density and relative frequency by two then multiplying by 100 (dividing by 2 provided an average value for, e. g., relative dominance and relative frequency). R1 was calculated separately for woody and herbaceous taxa. R1 was then used as a weight in a physiognornic classification scheme by Raunkaier (1934) (cited in Smith 1966) in which species were placed into one of five nominal groups according to the position of the perennating bud. A definition of each life form catagory is given in Table 6. Bud position was determined by referring to floristic manuals, i.e., Gray’s Manual of Botany (Femald 1950), Manual of the Vascular Plants of the Northeastern United States and Canada (Gleason and Cronquist 1991), and Flora of the Great Plains (Barkley et al. 1986), or, in a few instances, by inspection of numerous herbarium specimens. Table 6. Raunkaier’s life form categories. Life Form Description“ Therophyte Annuals, completing their life cycle in one season. Geophyte Buds buried belowground, on a bulb or rhizome. Hemicryptophyte Perennial shoots or buds near ground level, often covered with litter. Chamaephyte Perennial shoots or buds above ground level up to 25 cm. Phanerophyte Perennial buds over 25 cm above ground level. *Descriptions according to Raunkaier (1934), cited in Smith (1966). The cumulative species list of herbs for each site was evaluated with Raunkaier’s life form categories, as was the sampling data of 1988 and 1993, in which species were assigned to a life form category and then summed by RI. For data of 1988, 1993 and 1994, herb families were also summed by RI. Ferns (members of Pteridophyta) were grouped for comparison with other 48 vascular plant families. All herbaceous taxa for a site were classified by life history (annual, biennial, perennial). Similarly, each herbaceous species was classified by life form: grass, forb (an herbaceous dicot), legume and exotic. Note that the first two categories are mutually exclusive whereas the legume and exotic categories are not. A single species may belong to as many as three of four categories. For example, Kummerowia striata is an exotic legume which is also a forb. This classification scheme also excludes certain groups such as ferns and sedges. Herbs were also placed in up to four of nine habitat types. Table 7 provides a summary of the habitats derived from Mohlenbrock (1986). Again, each species was given an equal “vote,” receiving a maximum value of one. A low-fidelity species occurring in three habitats, for example, would contribute the value one-third to each habitat type. Species values for each habitat type were then summed. Habitat and life history summaries are based upon the cumulative species list for each site. Woody taxa in 1988 and 1993 were grouped according to their shade tolerance, viz., tolerant, intermediate, and intolerant, and then group R1 values were tallied. Shade tolerance for each species was preferentially assigned using Silvics of North America, Volumes I and II (Burns and Honkala 1990) when possible, then Michigan Trees (Barnes and Wagner 1981), and finally using habitat descriptions in Guide to the Vascular Flora of Illinois (Mohlenbrock 1986). Comparison of site similarity between 1988 and 1993, and between sites in 1993, was calculated using Jaccard’s index. Comparisons of management responses were made with the four managed sites, Brown-MGD, Cave-MGD, Gibbons—MGD, and Wildcat-MGD throughout. Site sampling data for 1988 and 1993 were not normally distributed, nor could they be transformed to approximate the normal distribution. Therefore, analyses of a site between years were made using the Mann-Whitney U test via the Statistical Package for the Social Sciences (SPSS), Version 6.1. Within-year statistical comparisons were also made between proximate sites 49 Table 7. Habitat categories and descriptions used to characterize herbaceous vegetation. Habitat Descriptions" Bluff Includes exposed slopes, cliffs, outcroppings, and ridges. Disturbed Includes disturbed soil, disturbed places, and roadsides. Edge Includes edges of woods and edges of fields. Open Includes open areas, openings, clearings, fields, pastures, and meadows. Open Woods Includes woodlands with continuous canopies, usually not >85°/o cover. Prairie Includes relatively undisturbed native grasslands which are not typically flooded. Lowland Includes streambanks, wet areas, wetlands, swamps, low ground, and floodplains. Thicket Includes any area characterized by dense woody growth, usually <3 m high. Woods Includes woods which are moist, rich, dry, rocky, flat, and upland. *Descriptions according to Mohlenbrock (1986). with the same substrate and forest-opening classification with 1993 data. Poaceae, Asteraceae, and exposed rock cover were analyzed in this way. A correlation analysis for total herb number and site opening size was performed using Excel, Version 5.0. Floristic spatial patterns in the 1994 data, if present, were assessed by dividing the transects into five sections or subhabitats, forest interior-north, transition zone-north, opening, transition zone-south and forest interior-south. Plots for the transition zones were selected relative to the forest boundary, one plot interior and one exterior, for all sites. Forest interior plots were the remaining two plots, distal to the forest-opening. Appendix C gives RI by subhabitat for woody and herbaceous species located in the 1994 plot sampling. R1 was also used to weight herb species classed by Raunkaier’s life forms and to weight shade tolerance categories for woody seedlings, shrub/saplings, and trees. No subhabitats were assigned for Gyp-UMG in which no continuous open area was present. Shannon and Simpson diversity indiees were calculated for herbs within each subhabitat. Species-area curves were used to ensure that comparisons were conducted using an adequate proportion of herb species inhabiting each subhabitat. Data for 1994 were not normally distributed, despite transformation, and therefore, analysis was pursued using the Kruskal-Wallis test via SPSS. Seedling density and percent cover 50 for Poaceae, Asteraceae, and exposed rock were compared statistically. Subhabitats with less than five plots were omitted from analysis. RESULTS Vegetation Composition Herb Species Composition Belonging to 88 families, 472 plant species were identified in this study. The largest families, the Asteraceae, Poaceae, Fabaceae, Rosaceae, and Cyperaceae comprised 68, 60, 28, 24, and 22 species, respectively, out of the 375 herb taxa encountered across all sites. The average number of species per site increased from 68 in 1988 to 192 in 1993-94, representing an increase of 182% (Table 8); however, given that 1988 surveys occurred in the opening while the 1993-94 surveys occurred in the opening and transitional area, one might expect an increase in total species number. Unmanaged sites averaged an increase of 154.5% in total species number between sampling years while managed sites increased 223.2%. The difference in the total number of herb species between years ranged from 63 (at Pounds-UMG) to 140 (at Cave-MGD). The average difference for woody species was 32. Cave-MGD had the highest species richness in 1994 with 261 species, while Pounds-UMG had the lowest, with 149. In an effort to facilitate assessment of herb cover and the observation of inconspicuous herb species, four nested l-m2 plots were used in 1994. When mean species richness in the 198 8, 50-m2 plots were compared to that of the mean found in four nested l-m2 plots, herb number increased 38.1% (4.5 species) (Table 9). Species richness increase between sampling periods was greater for unmanaged sites (58.4%) than for managed sites (23.5%). More species were found at managed sites in both years. However, the difference between managed and unmanaged sites was greater in 1988 (6.4 species), before treatment, than afterwards (4.8 species). The Poaceae and Asteraceae held the top two positions of relative importance (R1) for six sites in 1988 and 1993 (Table 10). The managed sites averaged a 0.7% increase in Asteraceae R1 51 52 2. H 8 mm H cm as H N2 S H we 3 H mm m _ H mm 3 H a: 3 H 3 mm _ H :82 .22 __< 3 H c... N m. H mm 3 H we. 3. H 8 m. m H 2. 3. H mm 3. H 2. 3 H 3. 72:. 82522.3 mm _ H :32 8 2 5 mm. 2. a 2 _ mm 2820 22.2% :2: 2:5: 2. cm 2: S S. 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H w... :32 32m __< YN H :1 m _ H m o o._ H ow $25 wowmcmEczv 532 ad H m4; mm md H No mo v.0 H m.m mm 23:0 2552mm tam venom _._ H vs om md H Wm 02 wd H m6 Cm 2220 2532mm Bozo: mussel 0.0 H v.9 om md H f0 02 v.0 H v.3 Om £5:me ocofincmm 25:55 “:0 9.0 H m6. em md H md 02 0.0 H We 0m own—O ocoancwm $25 830 wd H as mm Go H Ham m: 2 H 0.0 mm 250 22m oEitom SEED v; H QM: md H 2: 2m H m.m_ $23 nowmcmEv :82 0.. H 5.2 x. md H w. 9 mm 5.0 H _.m M: ova—O 2532:: tim 233:5 . H 0.5 m. m. o H n w co Yo H m._ _ m_ mcotmm ocoancam x85 2955 o._ H m.m~ mm m .o H _. _ _ we 9o H 0.3 mm 250 ocoumoES x35 o>mU wd H 52 cm m. o H o o 03 We H 0.2 om mantam Exam 5505 33.82 $5 v 1885 mai Z $5 .83 32 Z ANS cm .55 £3 Z 3mm .23: 38:03 :8: waist was $2 hoh 325/ .89 E 32a NE; n88: So“. be 2353 2: ES 32 E NE ow Ba .33 E NE 5m 860% €2— .Ho 55:5: Amm _ HV :32 d 2an Table 10. Relative importance (%) of the three most important herb families in 1988 and 1993. 54 Values for 1988 are derived from Heikens (1991). Site 1988 1993 Berryville Shale Glade- Asteraceae 23.9 Poaceae 31.3 unmanaged Poaceae 2 l .3 Asteraceae 17.0 Fabaceae 16.3 Lamiaceae 1 5 .2 Brown Shale Barrens- Poaceae 50.9 Poaceae 37.9 managed Asteraceae 23.7 Asteraceae 22.7 Fabaceae 6.3 Fabaceae 7.9 Cave Creek Limestone Asteraceae 49.7 Asteraceae 39.0 Glade-managed Poaceae 20. 1 Poaceae l3 .7 Smilacaceae 5 .9 Lamiaceae 5 .9 Cedar Bluff Sandstone Poaceae 47.4 Poaceae 33 .9 Glade-unmanaged Euphorbiaceae 9.2 Vitaceae 9.7 Smilacaeeae 9. 1 Anacardiaceae 7. 3 Gibbons Creek Sandstone Poaceae 49.4 Poaceae 33.2 Barrens-managed Asteraceae 12.0 Asteraceae 18.9 Euphorbiaceae 7.9 Fabaceae 9.6 Gyp Williams Sandstone Poaceae 39.3 Asteraceae 28.9 Barrens-unmanaged Asteraceae 14.8 Poaceae 20.3 Euphorbiaceae 14.4 Vitaceae 1 l .7 Pounds Hollow Sandstone Poaceae 35.9 Poaceae 40.9 Glade-unmanaged Euphorbiaceae 10.5 Euphorbiaceae 9.2 Ferns 8.5 Ferns 6.5 Round Bluff Sandstone Poaceae 49.1 Poaceae 35 .3 Glade-unmanaged Euphorbiaceae 1 5 .9 Buphorbiaceae 9. l Hypericaceae 8.9 Rubiaceae 7.7 Wildcat Bluff Limestone Asteraceae 37.2 Asteraceae 44.9 Glade-managed Poaceae 22.0 Poaceae 10.6 Lamiaceae 6.2 Fabaceae 9.9 55 between 1988 and 1993, and an 11.8% decrease for Poaceae. Unmanaged sites increased 3.6% for Asteraceae and decreased 6.3% for Poaceae RI over time. The Fabaceae and Euphorbiaceae also appeared among the t0p three most important families at four sites on at least one sampling date. The grasses, Schizachyrium scoparium and Danthom'a spicata, were the most important forest-opening species at five of nine sites in both 1988 and 1993 (Table 11). At unmanaged sites, Danthom‘a spicata was 4.5% (1988: pretreatment) to 6.8% (1993: post-treatment) more important than at managed sites. Although it did decrease in RI over time, this decrease was less for unmanaged sites (-23.6%) than for managed sites (-64.0%). Schizachyrium scoparium was slightly more important (2.0 to 2.6%) at managed sites than at unmanaged sites for both years despite a 53.1% decrease at the managed sites over time. At the two managed limestone glades, Cave-MGD and Wildcat-MGD, Silphium terebinthinaceum of the Asteraceae family was among the most important species for both sampling dates. However, it too decreased over time (-36.9% at Cave-MOD and -42.9% at Wildcat-MOD). Helianthus divaricatus was present among the top three most important herb species at five sites for at least one sampling year. At the managed sites it increased 33.3% from 1988 (RI=6.2%) to 1993 (RI=9.3%), whereas at the unmanaged sites a slight decrease (RI=-3.7%) was observed. It is notable that at Gyp-UMG, Parthenocissus quinquefolia, a characteristic woodland species, replaced Schizachyrium scoparium, a dominant prairie species, in the first R1 position in 1993. Mean herb cover (%) increased 61.9% from 1988 to 1993 (Table 12). Although herb cover increased over time at the unmanaged sites, it did not increase at the managed sites. Still, managed sites exceeded unmanaged sites in both years, averaging 21.4% (1993) to 24.0% (1988) more herb cover. The limestone glades, Cave-MGD and Wildcat-MOD, had the highest cover for both years. 56 Table 11. Relative importance (%) of the three most important herb species in 1988 and 1993. Values for 1988 are derived from Heikens (1991). Site 1988 1993 Berryville Shale Glade- Danthom'a spicata 10.5 Danthonia spicata 15.7 unmanaged Helanthus divaricatus 10.3 Cum'la origanoides 15.1 C unila origanoides 9.6 Schizachyrium scoparium 8.8 Brown Shale Barrens- Schizachyrium scoparium 28.7 Schizachyrium scoparium 16.2 managed Helianthus divaricatus 1 1.3 Dichanthelium Iaxiflorum 1 1.3 Danthom‘a spicata 10.2 Helianthus divaricatus 9.8 Cave Creek Limestone Schizachyrium scoparium 18.8 Silphium terebinthmaceum 8.2 Glade-managed Silphium terebinthinaceum 13.0 Schizachyrium scoparium 7.9 Aster oblongifolius 9.6 Verbesina virginica 5.8 Cedar Bluff Sandstone Danthom’a spicata 23.9 Damhonia spicata 9.9 Glade-unmanaged Schizachyrium scoparium 14.3 Parthenocissus quinquefolia 9.5 Dichamhelium acuminatum 9.2 T oxicodendron radicans 7.3 Gibbons Creek Danthom'a spicata 21.9 Helianthus divan'catus 15 .6 Sandstone Barrens- Schizachyrium scoparium 15.4 Dichanthelium acummatum 12.7 managed Dichamhelium laxiflorum 7.0 Schizachyrium scoparium 9.0 Gyp Williams Schizachyrium scoparium 22.9 Parthenocissus quinquefolz’a l 1.3 Sandstone Barrens- Helianthus divaricatus 5.8 Antennaria plantagim‘folia 9.9 unmanaged Lespedeza virginica 5.2 Helianthus divaricatus 7.9 Pounds Hollow Schizachyrium scaparium 18.9 Schizachyrium scoparium 10.9 Sandstone Glade- Danthom'a spicata 8.7 Danthom'a spicata 8.4 unmanaged Cheilamhes lanosa 7.9 C rotonopsis elliptica 8.2 Round Bluff Sandstone Schizachyrium scoparium 25.4 Schizachyrium scoparium 10.8 Glade-unmanaged Danthom'a spicata 17.1 Danthom'a spicata 10.5 Crotonopsis elliptica 15.9 Diodia teres 6.5 Wildcat Bluff Silphium terebmthinaceum 17.5 Silphium terebimhinaceum 10.0 Limestone Glade- Schizachyrium scoparium 15.4 Helianthus divaricatus 8.1 managed Smilax bona-nox 4.4 Solidago sp. 5.3 57 0.: 3 H 3m 3 H 0.: :82 8% =< 7: am Hi: 3 H3 92638585502 3 2 H N2 No I H 3 2 086 ocoamuswm tam 253. 4H 3 H 3 cs .3 H 3 CM €20 82%:sz 5%: €58 v8 2 H n: 9: no H 2: 3 823m 82%3 85:; 90 w: 2 H «.2 ca 2 H cm 8 820 ”Seesaw mam .86 T: 2 H3: N: 3H 2 mm ”use 29% 2.598 Bmgga 3w m.m H m3 E H 0.2 as; Sweeney :82 2: mm H 2:. Q 2 H o? M: 030 ”cough $3 “825» 3: i H g 8 S H on 2 805m ecosmufim 6.86 2896 2- M: H 8% we 2. H eon R 030 888.51. x86 38 to f H Nam ed a; H gm om 228m 22m .565 Ewan—2 855.5 82 z 32 z 85 .28: Sufism sec Escov as $2 E 83:3 .82 Ba £2 a .38 e3 Amm _ HV :8: .2 22¢ 58 Categorization of Herb Species Grasses, Forbs, Legumes and Exotics. In the forest-openings studied, forbs comprised the largest percentage of species (66.1% i 1.6%), followed by grasses (17.3% i 1.2%), legumes (included in the forb category) (9.2% i 0.9%) and exotics (6.6% i 1.6%) (Table 13). Unmanaged sites had a slightly higher percentage of grasses and exotics than managed sites (Ax=4.3% and 1.5%, respectively). However, managed and unmanaged sites were similar in the percentage of legumes and exotics. Sites proximate to human activity, Cave-MGD, Cedar-UMG, Pounds-UMG and Round-UMG, had the highest percentage of exotic species, together averaging 4.5% higher than the overall average. Life History. Perennials were the dominant life history type (as a proportion of all species) among forest-opening herbs (75.8% i: 2.5%), followed by annuals (20.6% i 2.7%) then biennials (3.6% i 0.4%) (Table 14). Managed- and unmanaged-site herb floras were also dominated by perennials and did not differ in the percentages of life history types. Raunkaier’s Life Forms. When classified by life form, it was determined that most forest- opening herb species were geophytes (40.4% i 3.8%), hemicryptophytes (38.0% i 5.6%) or therophytes (20.6% i 8.1%) (Table 15). Managed and unmanaged sites did not differ in the proportion of any life form except the geophytes, which were higher for managed (42.6% i 0.9%) than for unmanaged sites (38.8% i 2.1%). Also, managed sites had more geophytes (42.6% i 0.9%) than hemicryptophytes (39.4% i 1.9%). Categorization by Habitat. The herbaceous flora of the forest-openings was most highly associated with the woodland habitat (47.7% i 2.2%), followed by disturbed, Open, bluff, prairie and open woods habitats (Table 16). All other habitat association percentages were 31.8%. Managed sites had a higher percentage of characteristic prairie species (9.8% i 1.8%) than umnanaged sites (6.9% i 0.6%) while unmanaged sites were higher for species normally found in a 59 0._ H 0.0 0.0 H N0 0._ H _.00 N; H W: mm _ H 502 002m :< w.— H m n 0._ H Wm 0._ H m.m0 0._ H m.0_ 000:0. 000055.50 mm _ H :002 0.0. m0 0.00 0.0m 000.0 0:05.050 0025 0:200 .8 0.0 0.00 Sum 0030 0:900:0m Bozo: m0:000 Wm 0.2 0.00 0.0_ 0:095 0,5305% "WEE—:3 000 0._ _ m0 0.00 0.0m 000.0 0:90.050 0025 5000 :0. fix m.m0 0.3 0020 030m 0E>bc0m 009050850 _ m H N. m. m0 H _.0_ 0._ H 000 m0 H 0.3 $03. 0000525 mm _ H :002 0._ m0. W00 :1 0020 0:200:05 bin 800:? NN 00— 0.00 mm. 0:055 0co~m0cmm x0000 £5005 0.0. m .0 0:00 m0 _ 000—0 0:200:05 0.0000 0>0U 0.0 _.0_ 0.00 _.m_ 2.05m 0.00m 5505 00058—2 053m— 00 0:531— 00 0.80 00 00.26 00 03m 030280 5.022: 000 000000.00 0:502 0:0 3me 05 0:0 0020 030000000: :3 080 0:0 330 0001 00:03:00 05 00 005. 8 0: 85 00020 003 006000 seam .20: 006000 0.0; $00003 03:20:50 05 0o Bo 006000 00 590080 0.0 03 0008:0800 b00030 40.0.00. 5 00:0 00:3 05 E 00.0; 0098 0:0 00:500. .0050 603.0% 00 0w8=0800 0:0. .2 0.000. 60 Table 14. Herb life history (%) by site in 1993-94. Category percentages are the proportion of species out of the cumulative (1993-94) herb species lists. Site % Annual % Biennial % Perennial Managed Brown Shale Barrens 23.9 3.1 72.9 Cave Creek Limestone Glade 20.5 4.7 74.7 Gibbons Creek Sandstone Barrens 17.4 3.9 78.7 Wildcat Bluff Limestone Glade 6.3 3.9 89.8 Mean: 1 SE (managed sites) 17.0i3.8 39:03 79.0i3.8 Unmanaged Berryville Shale Glade 23.9 2.5 73.6 Cedar Bluff Sandstone Glade 25.6 3.9 70.5 Gyp Williams Sandstone Barrens 10.4 5.6 84.0 Pounds Hollow Sandstone Glade 25.0 0.9 74.1 Round Bluff Sandstone Glade 32.7 3.6 63.6 Mean :1: 1 SE (unmanaged sites) 23.5 i 3.6 3.3 i 0.8 73.2 i 3.3 All Sites Mean i 1 SE 20.6 i 2.7 3.6 _+. 0.4 75.8 i 2.5 61 0:300:0001um20. .003000052802n2m: .002000On0m0 .0:b_000E0;UH>t0m 00m0:0E:D 0.0 H 0.: 0._ H 0.00 0.0 H 0:00 N0 H 0.0 00:0 00w0c0E0 mm _ H :002 0.0 0&0 N00 0._ 0005 00000055 0005 H000:2, 0:0t0m 0.: 0.00 0. _0 0.0 0:900:0m 0.00:0 0:0006 0.0m 0.00 0.00 _._ 0:800:51— x0000 0>0U 0.0m 0.0.0 0.0 0.0 0:0t0m 0.00m :30:m 00w0002 Mir—.00 20:30 Om000 420000 03m .000: 000.500 0: w:_0:0000 00 0280000 8:00 005 00:00:00.5 0300—0: 3 0005000 00 000000000 0:00 005 .00-m00_ E 006000 0:00 :00 A000 00:00 0.:— 0.:0_00_:=0m .0. 0.000. .0005 0:200:00 002m :000..3n:000..3 .0005 0:80:00 .005 0:00.0n0:00m .0005 0:800:0m 30:0... 00:000n00::00 .0:0::0m 0:800:0m 00:00.5? §5u§5 .0:0::0m 0:900:0m 0.00:0 0:0005u0:0005 .0005 0:800:0m 00:5 :0005 ”:0000 .0005 0:988... 0.00:0 0>0UH0>0U .0:0::0m 0.00m 85050305 .0005 0.00m 0...>b:0mn0.:>.¢:0m 0083qu}, .00x02FnFU... 00.30.7031. .0.:.0:0u.:00 0:0 000.»: 00:00... .E 0. 03:00:00 0:0 m:.:000 00: .00 00300 0.00000 0 :00: 0000 0:0 0000 006000 .00»: 23.000 0:0 :00: 0:9: :. :0000 00:: 00:0... 00.0 0000 :0 00.. 006000 03:03:50 05 .00 :00 00.»: 000.000 :05 :. 0:00:80 006000 0:00 .00 5000020 05 0:0 0008:0800 000.»: 08.000 0:.: 8:. 006000 0:00 00.0 00000. .00 5000000000 .0. 0.00... 63 blufl" type habitat. Glades differed from barrens in having a greater proportion of bluff and disturbed herb associates, while barrens had a greater proportion of woods and open woods herbs than glades. The forest-opening habitat category means (above) were compared with those of three other communities, i.e., an undisturbed hill prairie in southern Illinois (V oigt and Mohlenbrock 1964), an invaded railroad right-of-way remnant prairie in west-central Kentucky (Bryant 1977), and a pristine, lowland mixed-hardwood forest in east-central Illinois (Aikman and Ebinger 1991) (Figure 9). The categories permitting the greatest discrimination among the aforementioned studies were the woodland and disturbed habitats. For the woodland habitat, forest-openings placed below the hardwood forest and above the hillprairie and railroad right-of-way sites. Forest-opening herbs were second only to the railroad right-of-way community for percent association to the disturbed habitat. Site Similarity Forest-opening similarity of the herbaceous flora between 1988 and 1994 was 19.3% i 1.1% (Table 17). Although managed and unmanaged sites both averaged 219% similarity, managed sites had more herb species in common (37.0 i 2.0) than unmanaged sites (30.8 i 2.7) over time. The mean Jaccard index (%) for all possible site pairs was 5.5% higher and there were 35.6 more common herb species than the same-site, 1988-1994 comparison in Table 17 (Table 18). Managed site pairs and sites with the same classification (i.e., the same substrate and forest- opening type) had more herb species in common than unmanaged sites and site combinations with dissimilar management histories. Sites with the same classification also had the highest Jaccard similarity (30.8% i 1.4%). 64 Figure 9. Percent association of the herbaceous flora of the study sites. a railroad right-of-way, a hill prairie, and a hardwood forest to nine habitat types. Habitat associations follow Mohlenbrock (1986). Sources: Railroad right-of-way (n=96 species) Bryant 1977, Hill prairie (n=39 species) Voigt and Mohlenbrock 1964, Hardwood forest (n=21 species) Aikman and Ebingcr 1991. 65 \\ \\\\\\ \\ _ \\\\\\\\\- , K\\\\\ ' ' ' ' ‘ ‘ a, :3 c; 8 8 r? 8 S g 8 N .- 66 Table 17. J accard site similarity for herbs and the number of common herb species for the study sites in 1988 and 1993-94. Data for 1988 are derived from Heikens (1991). The Jaccard index is described in Mueller-Dombois and Ellenberg (1974). The mean i 1 SE is reported. Site °/o Similarity No. Common Herbs Managed Brown Shale Barrens 19.5 39 Cave Creek Limestone Glade 17.4 39 Gibbons Creek Sandstone Barrens 14.9 31 Wildcat Bluff Limestone Glade 22.3 39 Mean i1 SE (managed sites) 18.5 i 1.6 37.0 i 2.0 Unmanaged Berryville Shale Glade 23.0 35 Cedar Bluff Sandstone Glade 15.4 24 Gyp Williams Sandstone Barrens 19.5 33 Pounds Hollow Sandstone Glade ' 23.6 37 Round Bluff Sandstone Glade 18.4 25 Mean i 1 SE (unmanaged sites) 19.9 i 1.5 30.8 i 2.7 All Sites Mean i1 SE 19.3 i 1.1 33.6 i 1.9 67 Table 18. Similarity of herb composition (Jaccard index) among pairs of sites in 1994. Site combinations are ranked from most to least similar. The Jaccard index described in Mueller- Dombois and Ellenberg (1974) was used. The mean i 1 SE is reported. Sites °/o Similarity No. Common Herbs GIBm & GYP“ 33.9 103 PDS" & CDRu 31.9 77 RNDu & CDRu 31.8 76 CAV"n & WLCm 30.8 98 GIB’“ & BRNm 30.6 103 GYPu & BRNm 30.3 86 BRNm & BVLu 30.0 84 GIBm & BVLu 29.1 87 BVL“ & CDRu 28.4 71 GIBm & WLCm 28.1 86 BRNm & CDRu 27.4 79 GYPu & WLCm 27.3 69 CAV"n & GIBm 27.2 100 GYPu & BVLu 26.0 64 GYPu & CDR“ 25.9 66 PDSu & BRNm 25.8 70 PDS“ & RND“ 25.7 57 RND & BRNm 25.3 68 RNDu & BVL“ 24.7 57 PDSu & BVLu 24.5 57 PDSu & GYP“ 24.1 57 GIBm & CDR“ 23.8 73 PDSu & GIBm 23.4 68 WLCm & BRN‘m 23.3 67 GIBm & RND" 22.6 65 GYPu & RNDu 22.6 53 CAV‘“ & GYPu 22.2 70 CAV'“ & BVLu 22.2 69 WLCm & BVL“ 21.7 54 CAVm & BRNm 20.6 72 CAV & CDRu 19.3 62 WLCm & CDRu 18.7 48 CAVm & RND“ 17.3 52 PDSu & WLCm 16.3 39 PDS“ & CAV'" 15.9 48 RND“ & WLC'“ 15.1 36 Mean (both sites managed) 26.8 i 1.6 (n=6) 87.7 i 6.2 (n=6) Mean (both sites unmanaged) 26.6 i 1.0 (n=10) 63.5 i 2.7 (n=10) Mean (dissimilar mgt. histories) 23.4 i 1.2 (n=20) 66.5 i 3.8 (n=20) 68 Table 18 (cont’d). Mean (same substrate and for.- opening classification) 30.8 i 1.4 (n=5) 82.2 i 8.3 (n=5) Mean (all combinations) 24.8 i 0.8 69.2 i 2.8 mManaged site. “Unmanaged site. BVL=Berryville Shale Glade, BRN=Brown Shale Barrens, CAV=Cave Creek Limestone Glade, CDR=Cedar Bluff Sandstone Glade, GIB=Gibbons Creek Sandstone Barrens, GYP=Gyp Williams Sandstone Barrens, PDS=Pounds Hollow Sandstone Glade, RND=Round Bluff Sandstone Glade, WLC=Wildcat Bluff Limestone Glade. Eleven herbs and 16 woody species were common to all nine study sites (Table 19). A11 11 herbs are native, seven are forbs, and eight are perennials. All 16 woody species are also native. However, exotic species were also shared by all sites (Table 20). Exotics found in the forest-openings included 40 herbs and eight woody species. Fifteen of these were found at Cave- MGD alone. Seven exotic plant species were found at Berryville-UMG, 11 at Brown-MGD, 31 at Cave-MGD, 19 at Cedar-UMG, 5 at Gibbons-MGD, 3 at Gyp-UMG, 11 at Pounds-UMG, 14 at Round-UMG, and 4 at Wildcat-MGD. Woody Species Composition Despite the different methods used to estimate canopy cover in 1988 (Heikens 1991) and 1993, the top two most important species were the same over time at five of the sites (i.e., Berryville-UMG, Cave-MGD, Cedar-UMG, Gyp-UMG, and Pounds—UMG) (Table 21). At Cave- MGD, Quercus rubra (red oak) and Quercus shumardii (Shumard’s oak) are believed to be the same species, although diagnosed differently, as they are difficult to distinguish. Juniperus virginiana (eastern redcedar) was conspicuous as the most important overstory member at all of the sandstone glades for both years. All other forest-openings, viz., the shale glade and barrens sites, were characterized by Quercus stellata, often accompanied by Vaccim‘um arboreum, Ulmus alata, and Quercus marilandica. 69 Table 19. Species common to all study sites in 1993 and 1994. LSpecies Common Name Woody: Amelanchier arborea Shadbush Carya ovata Shagbark Hickory Carya texana Black Hickory Celtis tenuifolia Dwarf Hickory Cornusflorida Flowering Dogwood Diospyros virginiana Common Persimmon Fraxinus americana White Ash Juglans nigra Black Walnut Parthenocissus quinquefolia Virginia Creeper Prunus seron'na Wild Black Cherry Quercus rubra Red Oak Quercus stellata Post Oak Quercus velutina Black Oak T oxicodendron radicans Poison Ivy Ulmus alata Winged Elm Vitis aesn‘valis Herbaceous: Acalypha gracilens Ambrosia artemisiifolia Carex umbellata C unila origanoides Danthom’a spicata Dichanthelium laxrflorum Helianthus divan'calus Lespedeza repens Ruellia humilis Schizachyrium scoparium Solidago ulmifolia Summer Grape Three-seeded Mercury Common Ragweed Sedge Dittany Curly Oat Grass Panic Grass Woodland Sunflower Creeping Bush Clover Wild Petunia Little Bluestem Elm-leaved Goldenrod 70 Table 20. Exotic species encountered at the study sites in 1993 and 1994. Species Woody: Campsis radicans Elaeagnus umbellata Ligustrum vulgare Lonicera japom'ca Lonicera sp. (shrub) Moms alba Pinus echinata Rosa multiflora Herbaceous: Abun’lon theophrastii Achillea millefolium Allium vineale Anagallis arvensis Asparagus oflicinalz’s Bromus commutatus Bromus racemosus Capsella bursa-pastoris Cardamine hirsuta Cardamine sp. Corom'lla varia Cosmos bipinnatus Daucus carota Dianthus armeria Digitaria sanguinalis Digitaria sp. F estuca arundinacea Kummerowia stipulacea Kummerowia striata Lactuca sem‘ola Lespedeza cuneata Leucanthemum vulgare Matricaria matricarioides Medicago lupulina Melilotus alba Phleum pratense Plantago lanceolata Poa compressa Poa pratensis Polygonum convolvulus Rumex acetosella Setariafaben‘ Sida spinosa Com_mon Nye Trumpet Creeper Autumn Olive Common Privet Japanese Honeysuckle White Mulberry Shortleaf Pine Multiflora Rose Velvet-leaf Common Yarrow Field Garlic Scarlet Pimpemel Asparagus Hairy Chess Chess Shepherd’s-purse Spring Cress Crown Vetch Cosmos Wild Carrot Deptford Pink Crab Grass Large Fescue Korean Bush Clover Japanese Bush Clover Prickly Lettuce Sericea Lespedeza Ox-eye Daisy Pineapple-weed Black Medic White Sweet Clover Timothy Buckhom Canadian Bluegrass Kentucky Bluegrass Black Bindweed Sour Dock Giant Foxtail Prickly Sida Sites of Occurrence“ BN, CV, RD WC BN, CD BV, BN, CV, CD, PD, RD BN, CD, WC BV, GB PD BV, CV, CD, PD CV BN, CD, GP BV, CV, CD, PD, RD CV BN CV, CD, RD, WC CV, CD, PD CV CD BV, BN, CV, GB, GP, RD BN CV RD CV, CD PD CD CV, CD, PD, RD CV, RD CD, GB, PD, RD, WC BV, BN, CV, CD, GB, RD CV, CD, GP, PD CV CV CV CV CV BN, CV CV, CD, PD, RD CD BV CD CV CV 71 Table 20 (cont’d). T araxacum oflicinale Common Dandelion CV T orilis japom'ca Hedge Parsley CV T rifolium campestre Low Hop Clover CV T rifolium pratense Red Clover CV Verbascum thapsus Woolly Mullein CV, CD, RD Veronica arvensis Corn Speedwell RD Viola raphanesquii Johnny-iump-up BV, BNLCV. GB, PD, RD *BV=Berryville Shale Glade-Unmanaged, BN=Brown Shale Barrens-Managed, CV=Cave Creek Limestone Glade-Managed, CD=Cedar Bluff Sandstone Glade-Unmanaged, GB=Gibbons Creek Sandstone Barrens-Managed, GP=Gyp Williams Sandstone Barrens-Unmanaged, PD=Pounds Hollow Sandstone Glade-Unmanaged, RD=Round Bluff Sandstone Glade-Unmanaged, WC=Wi1dcat Bluff Limestone Glade-Managed. Table 21. Relative importance (%) of the three most important woody species in 1988 and 1993. Canopy cover in 1993 was estimated according to Daubenmire (1959). In 1988 the sum of all overhead canopy was used to estimate cover. Values for 1988 are derived from Heikens (1991). 72 Site 1988 1993 Species “/o RI Species % RI Berryville Shale Glade- Quercus stellata 22.6 Quercus stellata 26.8 unmanaged Quercus marilandica 17.5 Quercus marilandica 14.9 Vaccim'um arboreum 14.7 Vaccim'um arboreum 8.8 Brown Shale Barrens- Quercus stellata 40.9 Quercus stellata 36.5 managed Ulmus alata 22.1 Ligustrum vulgare 14.8 Vaccinium arboreum 12.4 Vaccim'um arboreum 10.5 Cave Creek Limestone Quercus prinoides var.* 40.9 Quercus prinoides var.‘ 17.6 Glade-managed Quercus rubra 12.7 Quercus shumardi'i 14.7 Diospyros virginiana 8.9 Acer saccharum 9.9 Cedar Bluff Sandstone Juniperus virginiana 25.3 Juniperus virginiana 23.1 Glade-unmanaged Quercus stellata 20.9 Quercus stellata 19.5 Quercus marilana’ica 18.7 Ulmus alata 8.8 Gibbons Creek Sandstone Quercua stellata 29.7 Quercus stellata 24.7 Glade-managed Ulmus alala 24.1 Fraxmus americana 12.8 Carjya texana 17.1 Ulmus alata 10.3 Gyp Williams Sandstone Quercus stellata 28.1 Quercus stellata 19.9 Barrens-unmanaged Ulmus alata 20.5 Ulmus alata 17.0 Quercus marilandica 17.2 Carya texana 15.9 Pounds Hollow Sandstone Juniperus virginiana 30.9 Juniperus virginiana 32.6 Glade-unmanaged Ulmus alata 17.9 Ulmus alata 13.7 Vaccim'um arboreum 16.7 Vaccim'um arboreum 13.3 Round Bluff Sandstone Juniperus virginiana 34.9 Juniperus virgim'ana 43.3 Glade-unmanaged Quercus marilandica 15.7 Ulmus alata 15.2 Vaccim'um arboreum 13.1 Quercus stellata 12.2 Wildcat Bluff Limestone Quercus prinoides var.* 28.7 Quercus prinoides var.‘ 14.9 Glade-managed Juniperus virginiana 14.5 Quercus stellata 10.2 Querecus rubra 10.3 Diospyros virginiana 9.1 RI=Relative Importance. *Quepri =Quercus prinoides var. acuminata 73 showed comparable values for sites with expansive openings (i.e., Cave-MGD, Pounds-UMG, and Wildcat-MGD) or with very little opening remaining at all (i.e., Cedar-UMG and Gyp-UMG) (Table 22). The canopy cover values in 1993 were lower than in 1988 at sites in which the opening was not expansive (i.e., Berryville-UMG, Brown-MGD, and Round-UMG). For both years (pre- and post-management), canopy cover was greater at the umnanaged than at the managed sites. However, mean forest-opening canopy cover for all sites in 1988 (38.9% i 7.5%) and 1993 (29.0% i 5.2%) did not differ. The mean number of tree species (per 50 mg) was lower in 1988 (4.7 i 0.3) than in 1993 (7.2 .+_ 0.6) (Table 23). In 1993 there were nearly the same number of trees encountered at managed sites (7.1 i 0.4) as at unmanaged sites (7.3 i 0.6). However, managed and unmanaged site tree species number increased 77.5% and 40.4%, respectively, over time. Rock Cover Forest-opening rock cover (expressed as a percentage of total ground surface) for all sites did not differ between 1988 (20.3% i 7.1%) and 1993 ( 14.4% i 6.7%) (Table 24). However, exposed rock cover was greater at unmanaged sites than at managed sites for both sampling years. Statistical Comparisons: 1988 and 1993 Within-site statistical comparison of each forest-opening (1988 and 1993) showed that all sites were significantly different (0t=0.05) for Poaceae and Asteraceae cover except Berryville- UMG (Table 25). Percent cover of exposed rock was significant for about half of the forest- openings, i.e., Cave-MGD, Cedar-UMG, Gibbons-MGD, and Gyp-UMG. Between-site management comparisons for 1993 show that the unmanaged sandstone glade combination was significantly different in percent cover of exposed rock (on=0.05) (Table 26). Conversely, the managed-unmanaged site comparison was significant for Poaceae and Asteraceae cover. Neither management combination showed significance for canopy cover. 74 Table 22. Mean (i 1 SE) canopy cover (%) in 1988 and 1993 via two different methods. Canopy cover in 1993 was estimated according to Daubemnire (1959). In 1988 the sum of all overhead canopy was used to estimate cover. Values for 1988 are derived from Heikens (1991). Site N 1988 N 1993 Managed Brown Shale Barrens 30 29.5 :1: 0.2 30 9.4 :1: 2.4 Cave Creek Limestone Glade 27 180 i 3.9 27 15.0 1: 3.5 Gibbons Creek Sandstone Barrens 15 26.8 i 5.8 15 45,8 i 11,4 Wildcat Bluff Limestone Glade 18 166 i 33 18 12.2 i 2.8 Mean (managed sites) 22.7 i 3.2 20.6 i 8.5 Unmanaged Berryville Shale Glade 23 79.6 i 7.4 28 37.7 i 4.5 Cedar Bluff Sandstone Glade 30 49.1 t 4,9 30 42.9 i 3.5 Gyp Williams Sandstone Barrens 30 67.8 i 6.0 30 50.0 i 3.5 Pounds Hollow Sandstone Glade 30 21_8 3; 3,2 30 18.1 i 3.2 Round Bluff Sandstone Glade 23 41.4 i 4.1 23 30.1 i 5.1 Mean (unmanaged sites) 51.9 i 10.1 35.8 .+_ 5.5 All Sites Mean 38.9 i 7.5 29.0 i 5.2 75 Table 23. Mean (i 1 SE) number of tree species (per 50 m2) in 1988 and 1993. Values for 1988 are derived from Heikens (1991). Site N 1988 N 1993 Difference Managed Brown Shale Barrens 30 3.7 i 0.2 30 6.0 i 0.5 2.3 Cave Creek Limestone Glade 27 3.3 i 0.3 27 6.7 i 08 3.4 Gibbons Creek Sandstone Barrens 15 4.1 i 0.3 15 7.9 i: 0.4 3.7 Wildcat Bluff Limestone Glade 18 4.9 i 0.4 18 7.6 i 1.0 2.6 Mean (managed sites) 4.0 i 0.3 7.1 i 0.4 3.1 Unmanaged Berryville Shale Glade 23 5.5 i 0.4 28 8.8 :t 0.6 3.3 Cedar Bluff Sandstone Glade 30 5.7 i 0.3 30 9.1 i: 0.6 3.4 Gyp Williams Sandstone Barrens 30 5.9 i 0.3 30 9.3 i 0.5 3.4 Pounds Hollow Sandstone Glade 30 4.0 i 0.3 30 4.0 i 0.5 0.0 Round Bluff Sandstone Glade 23 4.8 i 0.4 23 5.2 i 0.6 0.4 Mean (unmanaged sites) 5.2 i 0.3 7.3 i 1.1 2.1 All Sites Mean 4.7 i 0.3 7.2 i 0.6 2.5 76 Wm- nd H v.3 :. H mdN :82 $35 :< 5N- N.: H MEN dd. H mdN $2: cowmcsEcsv :32 EN _- dd H w.mm mN 0v H Wow N «620 ocoamncam tam venom o. _- 06. H ddo dm 0.0 H 0.8 dm 2630 2.2855 BozoI messed Om- Nd H dd dm v; H we dm acotam ocoumncnm 35:23 9A0 we- wd H m.m om 5N H 0._ _ dm ode—O ocoamdcmm $35 .550 VNT _.m H 0.9 N Nd H 3N N 0320 22m o:_>ztom dowacchb 9m- _ m H mh d.m H w.w $26. dowmcmEv :82 Nd- N._ H 9m. m: _._ H wd M2 2620 ocodmoEfi tim Eon—<5 Va- m. d H m._ 2 m._ H new 2 mayhem ocoancmm #35 3255 0.0. d _ H Q: N 5N H OmN N 2620 ocoamoES x85 o>aU d. T Nd H _d dm Nd H _._ om mcotam 22m .555 domain: 855:5 mag Z 33 Z 85 .23: 28.6: EC?— UD>C®U 0.5 ww©_ hOm m®:_w> .m©©_ was wwmi a: Avomhhfi UCDOHW :30» “O Dwmucoouom w mwv xUOh Dumomxv mo uo>oo Amm _ HV flaw: .VN gawk. 77 Table 25. Probability values for the comparison (1988 and 1993) of Poaceae, Asteraceae, and rock cover in the opening of the study sites. The Mann-Whitney U test was used and an asterisk indicates a significant difference between sites (or=0.05). Data for 1988 were obtained from Heikens (1991, unpublished data). Site p-Value Poaceae Asteraceae Rock Managed Brown Shale Barrens 0.0000“ 0.0000“ 0.3103 Cave Creek Limestone Glade 0.0000“ 0.0000“ 0.0000“ Gibbons Creek Sandstone Barrens 0.0000“ 0.0000“ 0.0032“ Wildcat Bluff Limestone Glade 0.0000“ 0.0000“ 0.7540 Unmanaged Berryville Shale Glade 0.7753 0.0786 0.2070 Cedar Bluff Sandstone Glade 0.0087“ NA 0.0137“ Gyp Williams Sandstone Barrens 0.0000“ 0.0021“ 0.0001“ Pounds Hollow Sandstone Glade 0.0000“ 0.0000“ 0.4681 Round Bluff Sandstone Glade 0.0000“ NA 0.0621 NA=Not available. No Asteraceae were located in the 1988 sampling at Cedar Bluff Sandstone Glade-Unmanaged or Round Bluff Sandstone Glade-Unmanaged. Table 26. Probability values for site management comparisons of Poaceae, Asteraceae, canopy and rock cover in the site openings in 1993. The Mann-Whitney U test was used and an asterisk indicates a significant difference between sites (0L=0.05). Site Combination p-Value Poaceae Asteraceae Canopy Rock Unmanaged- Unmanaged Cedar-Round (Glades) 0.7377 0.7420 0.6804 0.0000“ Managed- Unmanaged Gibbons-Gyp (Barrens) 0.0000“ 0.0002“ 0.1253 0.3514 Cedar=Cedar Bluff Sandstone Glade, Gibbons=Gibbons Creek Sandstone Barrens, Gyp=Gyp Williams Sandstone Barrens, Round=Round Bluff Sandstone Glade. 78 Correlation analysis between total herb number and site Opening area yielded 0.00052 .% :w 3. an at m: an e: i: an em :55 2: a: .3 a: so: 3m 2 E as m: as ad 2: an Ex 02 an a: ”.2 an em as E 56 m: E 56 EN to 56 m: E 56 ed to 56 0.3 as. 55 3; as. 55 2m as. 55 a>m m: mmh .5 2-8. 2.; am 602 E 8% beam 0% mo 385935 95 E 86on as: “antenna “8:. 8:: 05 mo AX; 85:85: 0233M NN 28¢. 80 SaxoxmzmTaeu mam .mBEQaS 2:2me #5 A5 .5302: tomficumheQu 005 th .m0.§ ShSQHaS SQ $020323 EEKSEfiEQH RS 2Q .3083. 3:055ch Em «SQ .m03020wt0 02531.5 55 60:00.30 230200391? 0gb 06:35: 20:09“ 05: SD ram 08.59“ .3 EU .mmhaeteaumg 05003.:mn 08 tm 0052.300 Szotmamu Q00 Mam 508% SBVHEQ EV ram 03.051. .3 mt 00355000303 0:02:03"? 3% EV 032050 ckxuuexmsvxu Ed REV .0005 0000085 0005 “wow—LBHUJB n000—0 0,5605% tam nanoanZM .0020 0,5605% 30:03 mason—umna— amag—am 0:00m000m x0000 mconflOum—O .0020 0:00m000m .mBm EBUHMDU 000—0 0:03.065 x0000 0>00H>Eu0mux_>m .Ezomiotgfi “moaomnmdm .500m000N :ozacfibuméh £50000an .ntoZ-0=0N :0_u_m§.._.uz-mh .SLOZLotBE ~m000mHZ-E .008 003502... 0.x $00 Mam do 000 Mam do #5 .00 dd BREW dd 0.3 045V 0.0 .30 3% N6 30 BE m0 >00 N0: ms 53 New mm— :0 New m. 2 55 a0; 0.0 .50 BR 0. 2 a8 2% 0.2 .50 BR 0: .50 ask *0; .8083 R 03.? 81 Adam-00:85 0m000mum-E .fisom-000N 00_:m000-_-nm-m._- $00000“an .5002-000N 00:5005-HZ-m-r 0:02-00:25 00000muZ-E dd H 0.2 9m H de 9m H N. _ m 0.0 H mdN _.m H N.0N mm _ H 000.2 0.00m :< wd H _.m_ d.m H _d_ :0 H 0.3 0.2 H ddm 0.0 H mNm 303 00300003 mm _ H 000E -- -- Ndm 0N0 Wm. 000.0 00000003 tam 00:0m 0.mN _._N mNm 0d. _dm 000.0 000.0000m 30:03 000000 -- -- -- odd own 0006 00000008 tam 00000 md _._ _ vdv Ywm 0.3 000—0 30.5 0E>>t0m 00w000E03 0.x H 0.2 v m H dNN Nd H 0.0N _.0 H d. _N 0.0 H 0.9 $006 00.000000: mm _ H 000E 0.0 r... _ 0.0 0.3 0.2 0005 00050003 “Cs—m “000:3 0N. dNN Ndm m.wm m. _ N 2055 0000m000m 0.0000 000090 0.0 wd 0.3 Ym 0N 0005 00000065 x0000 0>0U 0. 3 N90 mdm m.mN m.mN 000t0m 200m 0305 00w000<4 m-E m-mb A—O Z-mh. 2.: 03m 000— E 0000 >020 05 00 20:00:03 030 E 2:000 0000000 05 00 $00 00000000: 0350M .mN 030E A. 3... L305 82 .500m000000— 00000mum-E .500m000N 00:3000rrum-m,r .w0_00mOnLO 0:02-000N 00:5000Fnz-mrr 000200.505 00000072400 .0 H w.mm _.m H 5mm m0 H 0.8 0.0 H _.om m.m H 0.0. mm _ H 000.2 000m :< M2: H QNN m m. H m m. _.m H 2. Wm. H _.m_ _.m H 0:2 @000 00w000E03 mm _ H 000.2 -: --- --- 0.0 02m 000.0 000.0055 tim 00:30 min.— 00 oh 0.0m 0. _ m 000.0 00000000m 30:0: 000000 --- --- :- v.— 0.m 0005 000%000m :05 .0000 5mm 0.0m No 0._ _ ndm 00000 0.00m 0_:>>t0m 00w000E0D we H 0 mm 0N H wsm N0 H fwm 0.0 H w.wm Wm H 0.8 $000 00w0000: mm _ H 0002 NE 0.0m 0.0m 0.0m _.m_ 000.0 00000000: :05 000003 Gmm mdm 0.: 0.: a. _ N 000.00m 000.0000m 0.0000 000006 0.0.0. 0.00“ «:00. Nov v. 3 000—0 000000003 0.0000 0>0U Ndm 0.0m o.wm 0.0.0. 0:. 000b0m 0.00m 0305 00w0002 méh m-m,—. m0 Zip—L 2.; 8% .000. E 0000 x020 0E 00 0000000000 03.0 E 30000 000000000< 00. 00 A000 00000000: 0200.0“ .3 0.00.0 83 Cyperaceae were more important in the opening of unmanaged sites than of managed sites (Table 30). For all sites, mean Cyperaceae R1 in the opening exceeded that in the transition zone- north and forest interior-north. Managed sites exceeded unmanaged sites in mean number of herb species (per m2) in all subhabitats except the transition zone-north (Table 31). In the opening (and transition zone-south) of unmanaged sites, there were fewer herb species than in the other subhabitats. There was, however, no difference in the mean number of herb species among subhabitats for the managed sites or for all sites. Mean herb cover (%) for all of the forest-openings did not differ among subhabitats (Table 32). However, cover at managed sites exceeded that of unmanaged sites for all subhabitats. Herb cover at managed sites also peaked in the opening subhabitat (48.3% i 8.4%). Rock, Bryophyte and Lichen Cover Mean cover of exposed rock (expressed as a percentage of total ground surface), was highest in the opening (17.2% i 3.6%) and transition zone-south ( 10.7 i 3.8) subhabitats of all of the forest-openings (Table 33). At unmanaged sites the opening had the highest cover value ' (16.6% i 0.0%), however, the transition zone-south and forest interior-south values were not available (due to truncation of unmanaged sites by bluffs). Managed sites had a higher percent cover of exposed rock in the opening than all of the other subhabitats except the transition zone- south. Mean bryophyte cover for all sites (5.6% to 9.0%) did not differ among subhabitats (Table 34). However, unmanaged site bryophyte cover exceeded that for managed sites in the opening subhabitat. Unmanaged sites also exceeded managed sites in opening subhabitat for lichen cover (Table 35). At managed sites, the opening and northern subhabitats exceeded that in the southern subhabitats. 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H 0.6 vd H ad $0.5 d0wacmEcs. 5.02 --- --- Wm H dd .H dd H m. a ..d H md d. 0.....0 0:985...“ .....m 953. a... H Vm. v ..d H ..d d Qm H 5w 0 ..N H N.m .. wd H ..d m. 0.00.0 05.02% Bozo... 0.05.9. --- --- --- mN H ...V d ..d H ..d o. 0.00.0 0:90....0m .....m 00.000 m. H m. .0 Nd H Nd v md H 0N m. dd H md. .0 N. H d.N .0 0.00.0 0.0.5 0:.>>t0m. d0wmancb ..d H Nd ..d H Nd dd H ... ..d H .0. md H ... 30:0 d0wmcaE. 5003. ..d H Nd o ..d H Nd d md H dd mm dd H wd o md H N.N o 0.00.0 25.00:... :25 .00....3 ..d H md c ..d H Nd d ... H 5N dN dd H wd d .d H ..d c 2.05%. 0.5.09.8 0.00.0 0.52.0 dd H dd 0 dd H dd 0 dd H ..d mm o. H 9N w bd H d. m 0.00.0 05.00:... 0.00.0 0300 ..d H ..d v ..d H Nd m vd H wd m. dd H m. w ..d H vd d. 0:055 0:28 5.55 3&0:an WE Z Wma. Z m0 Z Z-m.—. Z 2.; Z 0.5 .82 ... 8... 8.... 2.. .o 398.38 8.. ... ...... .88 88.. .mm . H. :82 .mm 0.89 90 Categorization of Herb Species Raunkaier’s Life Forms. Hemicryptophytes and geophytes were the most important forest- opening life forms (29.9% to 47.2%), followed by phanerophytes and therophytes (5.6% to 18.7%), then chamaephytes (23 .4%) (Table 36). Most forest-opening subhabitat values for hemicryptophytes and geophytes did not differ. The same was true for managed and unmanaged life form RI, in which standard error showed overlap in values for at least four of five subhabitats. Herb Species Diversity Simpson diversity (Ds) for all of the forest—openings did not differ among subhabitats (Table 37). However, managed sites had greater Ds diversity than unmanaged sites in the opening, and southern subhabitats. Similarly, Shannon diversity (l-l’) was greater for managed sites than for unmanaged sites in all subhabitats except the forest interior-north. Mean Shannon diversity did not differ among subhabitats for all of the forest-openings. Tree Seedling Composition Six forest-opening sites (i.e., Cave-MGD, Cedar-UMG, Gibbons-MGD, Pounds-UMG, Round-UMG and Wildcat-MOD) showed interesting trends in seedling importance (Table 38). For example, at Wildcat-MOD, Rhus aromatica occurred in the first position in the opening, Ulmus rubra was most important in both of the southern subhabitats and Ostrya virgihiana in both of the northern subhabitats. Seedlings of mesophytic species (i.e., Ulmus sp.) did not occur in the first position in the Opening of any site but were first in importance in the transition and forest interior subhabitats. Seedlings of the genus Quercus occurred in 22 of the 35 (62.9%) of the first RI positions and ofien accounted for 230% of tree seedling importance (Table 39). Cumulative seedling R1 of Quercus spp., however, did not show consistent peaks or troughs in the opening. Throughout the limestone glades, Cave-MGD and Wildcat-MOD, Ulmus was an important genus of tree seedling. 91 Table 36. Raunkaier’s life form (%) for herbs in five subhabitats of the study sites in 1994. Life form categories are summed by relative importance. Categories are according to Raunkaier (1934). The mean i 1 SE is reported. Site FI-N TS-N 0P TS-S Fl-S Chamaephyte: Berryville 21.4 15.9 22.5 12.8 15.3 Brown* 0.6 0.0 0.0 8.8 0.0 Cave“ 0.8 0.6 0.1 0.0 0.0 Cedar 0.9 2. 5 --— --- --- Gibbons* 0.0 0.0 0.0 0.0 3.6 Pounds 1.8 1.7 0.0 0.0 4.1 Round 0.0 0.0 0.0 --- «- Wildcat* 2.6 2.5 0.7 0.0 0.0 Geophyte: Berryville 20.7 11.6 17.9 35.4 40.6 Brown* 40.6 42.1 52.2 61.8 61.9 Cave’“ 44.1 40.8 44.8 41.1 17.2 Cedar 2 5.9 22.4 --- --- --- Gibbons* 37.8 43.3 49.1 41.6 43.6 Pounds 20.2 35.1 30.5 60.0 39.9 Round 27.9 38.3 41.9 --- «- Wildcat* 21.6 45.4 36.6 43.3 43.3 Hemicryptophyte: Berryville 52.9 72.4 57.8 33.7 38.4 Brown* 38.2 49.7 35.5 26.6 31.5 Cave* 36.1 35.5 40.0 39.6 37.9 Cedar 36.6 62.2 --- --- «- Gibbons“ 41.7 45.5 31.9 29.9 19.8 Pounds 47.4 31.4 33.4 20.0 39.9 Round 38.5 25.3 16.8 --- «- Wildcat* 45.1 33.3 45.3 36.0 36.0 Phanerophyte: Berryville 4.9 0.0 0.0 18.1 0.0 Brown* 13.6 4.8 8.9 1.4 0.0 Cave“ 18.1 21.8 11.9 19.3 44.9 Cedar 33.6 2.5 --- --- --- Gibbons* 11.6 3.7 1.6 8.4 24.4 Pounds 21.7 19.6 20.9 10.0 0.0 Round 25.6 6.9 0.0 --- «- Wildcat* 20.1 18.8 15.1 18.3 21.6 Therophyte: Benyville 0.0 0.0 1.8 0.0 5.8 Brown* 7.0 3.4 3.4 1.4 6.6 Cave“ 0.8 1.2 3.2 0.0 0.0 Cedar 2.9 10.4 --- --- «- Gibbons“ 8.8 7.5 17.5 20.1 8.7 Pounds 8.9 12.2 15.1 10.0 16.2 Table 36 (cont'd). Round 7.9 29.5 41.3 --- --- Wildcat* 10.6 0.0 2.4 2.3 2.8 Mean (managed sites): Chamaephyte 1.0 i 0.6 0.8 i 0.6 0.2 i 0.2 2.2 i 2.2 0.9 i 0.9 Geophyte 36.0 j; 4.9 42.9 i 0.9 45.7 j; 3.4 46.9 i 4.9 41.5 i 9.2 Hemicryptophyte 40.3 j: 1.9 41.0 i 3.9 38.2 i 2.9 33.0 i 2.9 31.3 i 4.1 Phanerophyte 15.9 i 1.9 12.3 i 4.7 9.4 i 2.9 11.9 i 4.3 22.7 3; 9.2 Therophyte 6.8 i 2.1 3.0 i 1.6 6.6 i 3.6 5.9 i 4.7 4.5 11.9 Mean (unmanaged sites): Chamaephyte 6.0 i 5.1 5.0 i 3.7 7.5 i 7.5 6.4 i 6.4 9.7 i 5.6 Geophyte 23.7 i 1.9 26.9 i 6.1 30.1 _+_ 6.9 47.7 :12.3 40.3 i 0.4 Hemicryptophyte 43.9 i 3.8 47.8 i 11.5 36.0 i 11.9 26.9 i 6.9 39.2 i 0.7 Phanerophyte 21.5 i 6.0 7.3 i 4.4 6.9 i 6.9 14.1 j; 4.1 0.0 i 0.0 Therophyte 4.9 i 2.1 13.0 : 6.1 19.4 3: 11.6 5.0 _+_ 5.0 11.0 i 5.2 Mean (all sites): Chamaephyte 3.5 _+_ 0.3 2.9 i 0.4 3.3 i 3.2 3.6 i 2.3 3.8 i 2.4 Geophyte 29.9 i 3.4 34.9 i 4.2 39.0 i 4.5 47.2 i 4.5 41.1 i 5.8 Hemicryptophyte 42.1 i 2.1 44.4 _+_ 5.8 37.2 i 4.8 30.9 3: 2.9 33.9 i 3.1 Phanerophyte 18.7 i 3.1 9.8 i 3.1 8.3 i 3.1 12.6 i 2.9 15.2 i 7.5 Therophyte 5.9 i 1.4 8.0 i 3.5 12.1 i 5.4 5.6 i 3.3 6.7 i 2.3 92 F l-N=Forest Interior-North. TS-N=Transition Zone-North. OP=Opening, TS-S= Transition Zone-South, Fl-S=Forest Interior-South. *Managed site. 93 8. 0.. 0.0 28.0 82880 0.20 8.50 ...N 2. 0.. 0.. 0.0 28.0 8988.0 32.0: 28.5.. ...m ...N 0.. a... 0.. 2.20 22.0 2.30.80 80885.... SH 2 SHE N... H 2 ....H 0.0 ....HZ 888088.... 00 . H82). 0.0 0.0 0.0 ..m 0.0 2.20 898...... 020 822.3 3- 2 0.0 2 ..m 08.80 89880 28.0 880.0 0.0 0.0 0.0 0.0 0.0 2.20 888...... 28.0 88 0.0 N.N ..N 4.0 0.0 8050 280 88.0 8088.2 ”hm. 00:50.5 N. H40 0.30... .... H: .03.... 0.. H00 00 . H58: 8.0.? a... H .s S H 0.0 a... H .... 0... H .... 0.0 H 0... .8... 8088...... 00 . H 82>. 0.0 0.0 0.0. 28.0 898.80 .020 .880 0.0 0.. ..m 0... .s 2.20 882.80 32.0.. 28.5.. o... m... ...m E. 2 28.0 22.0 2.30.80 8858.... 0.. H0... EH3. 0.. Ho... 2 H4... 0.. Ho... 888885.00 . H582 ...N. a. . N 0.0 4.0. S. 2.2.. 828...... .020 8.2.3 0.0 ...N. 0... 0.0 0.0. 88.80 89880 28.0 82.06 0.0. E. 0.0. 2.. 0.0 28.0 888...... 28.0 86 0.0 0.0 0.0 3. 0.0 80.80 280 88.0 8028.). A00. 509:5 0-... 0-0.. .5 2-0.- 2-... 8.0 .voo. ... 000.0 .630 05 mo 0.850.320 0>w ... 00.0000 0.0.. 8.. 0.0.0.»... 5:55 0:0 509:6 ..m 0.00-.- Ill AU. ~CCUC A". O\C~...N. 94 508-55.:— .0o.on.nm-_m .5:om-2.oN co...0:0.._.um-m._. .wcfioncnmo .atoZ-o=oN so...0§.HHZ-m,_. ...toz-.oc8£ 0020..."??— m.o H ON vd H ..N Nd H N.N m.o H N.N Nd H Wm Mm _ H :32 00am =< N... H N.N a... H 0.. 3 H 0.. M... H 0.. ...o H N.N .8... 808.85.. 00 . H 58.2 .2558 um 030,—. 95 Table 38. Relative importance (%) of the three most important species of tree seedlings in five subhabitats of the study sites in 1994. Site F I-N TS-N OP TS-S FI-S BVL Ame arb 24.5 Que ste 21.9 Que ste 17.9 Que ste 26.0 Que vel 13.8 Vac pal 22.1 Que mar 18.8 Que mar 14.4 Que vel 12.3 Car tex 12.'6 Que vel 8.3 Vac arb 10.9 Rhu cop 12.6 Rhu aro 11.4 Sas alb 12.2 BRN“ 0st vir 17.5 Que ste 14.9 Que ste 22.2 Vac arb 26.0 Ulm ala 23.7 Que ste 15.2 Ost vir 13.7 Vac arb 13.2 Ulm ala 16.6 Vac arb 20.4 Que vel 7.8 Car gla 8.3 Ulm ala 12.0 Que ste 15.8 Que ste 10.3 CAV“ Ulm rub 14.6 Rhu aro 19.8 Que pri 12.8 Ulm rub 14.0 Ulm rub 22.4 Cel ten 9.4 Ulm rub 11.1 Rhu aro 12.3 Cel ten 11.3 Cel ten 15.1 Rhu aro 9.3 Cel ten 6.8 Ulm rub 9.9 Rhu aro 10.8 Ace sac 9.0 CDR Que ste 11.8 Que ste 25.1 Car ovt 11.3 Ulm ala 18.6 Rhu aro 9.9 Jun vir 9.8 GIB“ Sym orb 23.1 Ulm ala 25.6 Que ste 16.3 Que ste 16.3 Que ste 16.1 Car tex 12.3 Sym orb 21.2 Car tex 14.1 Que mar 10.8 Car tex 10.6 Ulm ala 10.3 Car tex 12.9 Vac pal 9.5 Vac arb 9.5 Que vel 9.9 PDS Que ste 21.6 Que ste 19.9 Ame arb 19.2 Que alb 21.5 Que alb 23 .4 Vac arb 14.0 Ame arb 13.9 Que ste 16.2 Ame arb 13.8 Ame arb 13.0 Que mar 8.4 Vac arb 12.6 Vac arb 15.7 Que ste 7.7 Que rub 9.9 RN D Ulm ala 13 .9 Ulm ala 16.6 Vac arb Jun vir 11.0 Rhu aro 16.1 Ulm ala Car ovt 9.3 Car ovt 10.7 Que ste WLC“ 0st vir 19.3 05! vir 19.9 Rhu aro 17.0 Ulm rub 24.9 Ulm rub 16.8 Ulm rub 17.6 Ulm rub 11.1 Ulm rub 12.9 Rhu aro 8.8 Os! vir 11.2 Ace sac 8.9 Ace sac 7.2 Os! vir 11.6 Ulm ala 8.6 Cer can 9.3 ‘Managed site. ' FI-N=Forest1nterior-North, TS-N=Transition Zone-North, OP=Opening, TS-S=Transition Zone- South, F1-S=Forest Interior-South. BVL=Berryville Shale Glade, BRN=Brown Shale Barrens, CAV=Cave Creek Limestone Glade, CDR=Cedar Bluff Sandstone Glade, GIB=Gibbons Creek Sandstone Barrens, PDS=Pounds Hollow Sandstone Glade, RND=Round Bluff Sandstone Glade, WLC=Wildcat Bluff Limestone Glade. Ace sac =Acer saccharum, Ame arb =Amelanchier arborea, Car gla =Carya glabra, Car ovt=Carya ovata, Car tex=Carya texana, Cel ten=Celtis tenuifolia, Cer can=Cercis canadensis, Jun vir=Juniperus virginiana, Os! vir=03trya virginiana, Que mar=Quercus marilandica, Que pri =Quercus prinoides var. acummata, Que rub =Quercus rubra, Que ste =Quercus stellata, Que 96 Table 38 (cont’d). vel=Quercus velutz'na, Rhu aro =Rhus aromatica, Rhu cop =Rhus copallina, Sas alb =Sassafras albidum, Sym orb =Symphon‘carpos orbiculatus, Ulm ala=Ulmus alata, Ulm rub=Ulmus rubra, Vac arb =Vaccim'um arboreum, Vac pal =Vaccim’um pallidum. 97 ..N. ...... ES 0... .... ...... .... .... 8.. ..N. .... 80 .... 8.. ...... .... .8. 55 N... 8.. 8. 0..... a... ...... a... .... ...-o 020 6.x Q...» 93A 0.x .Qm .69 h.m~ 9.6 03; .©.N_ 9.6 USA 0.x .Qm LSD ...... .... 8... .... .... 8... N... ...... 8... a... 8.. 8... ...... .8 8. .... .... 8m 0..... .... 8m ....m .... 8m a... .... 8m. ..m. .... 8w 00. 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A0000 00w000E03 mm . H 000.). 000.0 000.0000m ......m 0000: 000.0 00800000; 30:02 00000; 000.0 000.0000m .005 .0000 0.020 220 2.350 .0000 00300.... mm . H 000.). 0.0.0 2.902.... ......0 .8205 000..0m. 000.0000m 0.00.0 00000.0 000.0 000.020.: 0.00.0 0>0U 000..0m 0.0:m 038m .0000 00w000E00. mm . H 0003. ”.003. :0. mm . H 000.>. 00.00000. 0 D 00w000.). SEW—20h. A0280. NV 050,: 104 Table 43. Relative importance (%) for the three most important shrub and sapling species in five subhabitats of the study sites in 1994. Site FI-N TS—N OP TS-S FI-S BVL Vac arb 45.5 Que mar 47.2 Que mar 59.0 Fra ame 25.0 Car tex 50.0 Vac pal 23.7 Vac arb 34.7 Vac arb 29.2 Que ste 25.0 Ulm ala 50.0 Que mar 20.5 Que ste 18.1 Que ste 11.8 Sas alb 25.0 Ulm ala 25.0 BRN“ none none Que ste 50.0 Vac arb 58.3 Vac arb 76.2 Vac arb 50.0 Ulm ala 41.7 Ulm ala 23.8 CAV“ Ost vir 28.8 Cor a’ru 39.4 Fra ame 23.9 Cer can 70.8 none Fra ame 22.5 Ace sac 21.9 Cer can 19.9 Jun vir 29.2 Car tex 16.3 Fra ame 12.9 Que pri 15.7 Que shu 12.9 Ulm rub 12.9 CDR Jun vir 33.5 Jun vir 100 Ulm ala 31.6 Fra ame 9.7 GIB” Ulm ala 50.0 Ulm ala 69.0 Ulm ala 48.1 Vac arb 55.0 Car ovt 50.0 Car tex 25.0 Car tex 15.5 Ame arb 14.8 Ulm ala 45.0 Ulm ala 50.0 Fra ame 25.0 Jun vir 15.5 Car tex 14.8 PDS Vac arb 39.7 Jun vir 41.7 Jun vir 31.7 Jun vir 32.5 Dio vir 33.3 Que mar 22.6 Ame arb 26.3 Ulm ala 23 Bac arb 17.5 Ame arb 16.7 Ulm ala 18.8 Vac arb 22.5 Vac arb 21.7 Que alb 15.0 Que alb 16.7 Vac arb 16.7 RND Car gla 50.0 Jun vir 70.8 Jun vir 73.3 Ulm ala 22.5 Ulm ala 29.2 Ulm ala 26.7 Jun vir 16.3 WLC" Ost vir 46.0 Os! vir 75.0 03! vir 55.6 Ost vir 50.0 Ulm rub 40.2 Ace sac 21.6 Ace sac 25.0 Ulm rub 19.0 Cel ten 25.0 Cra sp. 29.4 Ame arb 10.8 Ace sac 6.3 Ulm rub 25.0 Os! vir 25.3 *Managed site. F I-N=Forest Interior-North, TS-N=Transition Zone-North, OP=Opening, TS-S=Transition Zone- South, FI-S=Forest Interior-South. BVL=Berryville Shale Glade, BRN=Brown Shale Barrens, CAV=Cave Creek Limestone Glade, CDR=Cedar Bluff Sandstone Glade, GIB=Gibbons Creek Sandstone Barrens, PDS=Pounds Hollow Sandstone Glade, RND=Round Bluff Sandstone Glade, WLC=Wildcat Bluff Limestone Glade. Ace sac =Acer saccharum, Ame arb =Amelanchier arborea, Car gla=Carya glabra, Car 105 Table 43 (cont’d). ovt=Carya ovata, Car tex=Carya texana, Cel ten=CeItis tenuifolia, Cer can=Cercis canadensis, Cor dru=C0mus drummondii, Cra sp. =Crataegus sp., Fra ame =Fraxinus amen'cana, Jun vir=Juniperus virginiana, Ost vir=03trya virginiana, Que alb =Quercus alba, Que mar=Quercus man'landica, Que pn‘ =Quercus prinoides var. acuminata, Que shu=Quercus shumardii, Que ste =Quercus stellata, Que vel=Quercus velun'na, Sas alb =Sassafras albidum, Ulm ala=Ulmus alata, Ulm rub =Ulmus rubra, Vac arb=Vaccinium arboreum, Vac pa! = Vaccini um pallidum. 106 subhabitats except the forest interior-south. The mean number of shrub and sapling species (per 25 ml) for all sites ranged from 0.6 i 0.1 to 0.9 i 0.2 (Table 44). There was no difference among forest-openings for mean shrub and sapling species richness. Likewise, managed and unmanaged sites did not differ in the number of shrub and sapling species among subhabitats. Shrub and sapling density (per 25 m2) for all sites was lower in the opening than in the northern subhabitats (Table 45). Except for the forest interior-north, managed and unmanaged site subhabitats did not differ in density. Shade-tolerant and shade-intolerant shrubs and saplings were more important than the shade-intermediate category in all subhabitats except the forest interior-south (Table 46). Unmanaged sites exceeded managed sites for shade intolerance in the opening and northern subhabitats. However, managed sites exceeded umnanaged sites in the opening in the shade- intermediate category and in both of the transition zones for shade tolerance. Tree Composition Fire-tolerant Quercus species were the most important trees in the opening of all of the managed sites (Table 47). Although the opening at one unmanaged site (Berryville-UMG) was dominated by Quercus marilandica, the openings of the two other unmanaged sites were dominated by Juniperus virginiana, a fire-intolerant species. At all managed sites, mesophytic species such as Fraxinus americana, Ulmus alata, and Acer saccharum increased in R1 in the transition and forest interior subhabitats. Species of Quercus occurred in 22 of the 35 (62.9%) first positions of R1 and often accounted for 235% of forest-opening RI (Table 48). Relative importance for this genus also peaked in the opening, accounting for 56.7% of opening R1 for trees. Juniperus virginiana RI also peaked in the opening, at =24%. 107 .5.=OW-..0CD:: .mo..0n~.llmu~n_ ...—.SOWiUZON F.O_._m:m..r—tflmumr~. .wcmcvmmvflnwo ...—...OZuDCON :O_:m:m..r~tflz..mr_t .~_...OZu..Om..D:—— .m®.OlHZi—n~ md H 0d ..d H wd ..d H 0d ..d H 0d Nd H dd 000.). 000m :< Nd H 0d Nd H N._ . d H ..d ..d H 0.0 md H 0... .00.... 00300003 000.). --- --- md H wd v Nd H ..d d Nd H wd d. 000.0 000.0000m 0.0:. 0003. md H wd 0 0d H 0:. 0 md H ..d 0 md H 0d .. vdH 0d m 000.0 000.0000m 30:0: 000000 --- Nd H md 0 Nd H 0._ 0. 000.0 000.0000m tam. .0000 md H md 0 0d H d. v Nd H 0d 0.. 0d H d._ v md H w. v 000.0 0.00m 0:3»..0m 00w000E0D 0d H wd dd H 0d ..d H md Nd H 0d md H ..d .0000 00w000EV 000$. md H d.N 0 md H 0d 0 ..d H Nd mm. md H md 0 0d H m. 0 000.0 000.000..1...0_m..000_.>> Nd H md 0 md H 0.0 0 Nd H ..d dN 0d H d. 0 md H 0d 0 m00..0m. 000.0000m 0.00.0 00000.0 dd H dd 0 Nd H md 0 ..d H 0d mm md H wd w md H 0d m 000.0 000.0065 0.00.0 0>0U md H wd 0 0d H 0d m ..d H Nd m. dd H dd w dd H dd d. m00..0m. 0.00m 0305 00.3003. m...— Z m.m... 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H 0.0. 0.0 H 0.0 0 m H 0 H o 0 H 0.0 0.0 H 00. 0.2... 8022900522. .022 0.0 0.0 0.0 0.0 0. _ N 2.20 8232: 020 022.3 0.00 0.0 0.: 0.0. 0.00 2.20 2.22250 28.0 293.0 0.0 0.0 N.N 0.0 0.0. 2.20 2.235: 2020 2:00 0.0 0.0 0.0 0.0 0.0 22.00 22.0 55.0 302.02 no.0.fiDE.o.:. 0.0 H 0.0 0.: H 0.00 0.0. H 0.0. 0.0 H 3.0 0.0. H 0.9 02.0 8022220 :32 00. 0.0. 0.00 2.20 25.0050 020 2:50 ...0. 0.0.. 0.00 20 N. . 0 2.20 82002.0 32.0: 2.55. 0.: 0.0.. 2.20 222.50 020 .2000 0.0 0.0. 00. 0.: 00. 2.20 22.0 2.350 80205:: .0. H .0. 0.0 H 0. 0. 3. H 0.2. 0.0. H 0.00 0.0 H 0.0. 02.0 8022... :82 0.0 0.00 0.0. 0.0 o... 2.20 820225 020 022.3 0.0 3.0 0.: 0.0. 0.00 22.00 822.50 V.020 2.80.0 0.0 000 0.00 0.00 0.00 2.0.0 82865 28.0 2,00 0.0. 0.00 00. 00 0.0 22:00 22.0 55.0 802.02 NEED—OE. 0-.... 0-0.0 .0 2.0.. 2.... 2.0 000000. 0. mm . 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H m.00 0.d H d..N WNN ..d. 0.00 ..mH0.m m... 00.0.0... 0.0.00E.0.0. 00.0.0.0. .0000 00d0000.00-.00.0.0.. 000.). 2.20 25.22.00 .020 .0550 000.0 000.0000m 32.0.. 000000 000.0 000.0000m .....m. .0000 2.20 22.0 2.3050 .0000 00w0000.-.00.0.0.. 0002 0000 00800001. ..0.m. 000:3 000.0 000.0000m 000.0 00000.0 000.0 000.000.... 0.00.0 0>00 000..0m. 0.00m 038m. .0000 00w000E00-.000..0.0.. 000.). 2.20 2.20050 .020 .253. ”.0000 ..0. 000.). 00w000003 00m000<0 000.20... ..0..000. 00 0.00... 111 Table 47. Relative importance (%) for the three most important tree species in five subhabitats of the study sites in 1994. Site FI-N TS-N OP TS-S FI-S BVL Que mar 62.5 Que mar 50.0 Que mar 66.8 Car tex 50.0 Que vel 42.9 Que ste 37.5 Que ste 50.0 Que ste 33.2 Que ste 50.0 Car gla 14.3 Car tex 14.3 BRN” Que ste 50.0 Que ste 63.3 Que ste 100 Que ste 100 Ulm ala 100 Car gla 25.0 Car gla 15.6 Ulm ala 25.0 Ulm ala 10.6 CAV“ Fra ame 35.4 Que ste 27.2 Que pri 33.3 Que pri 36.7 Ace sac 20.8 Que pri 35.4 Fra ame 17.2 Que shu 25.0 Corflo 26.7 Que pri 20.8 Que ste 17.4 Que shu 14.2 Fra ame 16.7 Fra ame 18.3 Fra ame 14.6 CDR Jun vir 40.2 Jun vir 45.3 Ulm ala 32.1 Ulm ala 32.8 Fra ame 10.6 Que ste 21.9 GIB“ Que ste 28.6 Que ste 33.2 Que ste 44.9 Que ste 43.7 Que ste 73.3 Car tex 19.1 Ulm ala 28.6 Car tex 17.8 Ulm ala 25.4 Car tex 17.8 Fra ame 14.1 Car tex 19.1 Que mar 16.9 Car tex 18.3 Car gla 8.9 PDS Que ste 32.6 Jun vir 66.7 Jun vir 68.3 Jun vir 56.3 Jun vir 37.5 Jun vir 21.8 Que ste 16.7 Que mar 18.3 Que ste 21.8 Dio vir 12.5 Que mar 21.8 Que mar 8.3 Ulm ala 13.3 Car gla 12.7 Que alb 12.5 RND Jun vir 47.2 Ulm ala 48.9 Jun vir 100 Ulm ala 28.2 Jun vir 31.3 Fra ame 15.5 Car gla 19.8 WLC"I Ost vir 27.5 Ace sac 66.8 Que pri 20.9 Ulm rub 26.7 Car ovt 24.7 Car ovt 16.3 Que alb 19.6 Que shu 16.9 Car ovt 18.3 Cra sp. 24.7 Fra ame 11.3 Car gla 13.4 Que ste 15.5 Ost vir 18.3 Fra ame 16.7 *Managed site. FI-N=Forest Interior-North, TS-N=Transition Zone-North, OP=Opening, TS-S=Transition Zone- South, FI-S=Forest Interior-South. BVL=Berryville Shale Glade, BRN=Brown Shale Barrens, CAV=Cave Creek Limestone Glade, CDR=Cedar Bluff Sandstone Glade, GIB=Gibbons Creek Sandstone Barrens, PDS=Pounds Hollow Sandstone Glade, RND=Round Bluff Sandstone Glade, WLC=Wildcat Bluff Limestone Glade. Ace sac =Acer saccharum, Car gla=Carya glabra, Car ovl =Catya ovalis, Car ovt=Carya ovala, Car tex=Carya texana, Cra sp. =Crataegus sp., Dio vir=Diospyros virginiana, Fra ame =Fraxinus americana, Jug m'g =Juglans nigra, Jun vir=Juniperus virginiana, Os! vir=03trya virginiana, Que alb =Quercus alba, Que coc=Quercus coccinea, Quemar=Quercus 112 Table 47 (cont’d). marilandica, Que pri =Quercus prinoides var. acummata, Que shu =Quercus shumardii, Que ste =Quercus stellata, Que vel=Quercus velun'na, Ulm ala=UImus alata, Ulm rub=Ulmus rubra 113 Table 48. Relative importance (%) for the three most important tree genera in five subhabitats of the study sites in 1994. Trees are listed to species when possible. Site FI-N TS-N OP TS-S FI-S BVL Que spp. 100 Que spp. 100 Que spp. 100 Car tex 50.0 Que vel 71.5 Que ste 50.0 Car spp. 28.6 BRN“ Que ste 50.0 Que ste 63.3 Que ste 100 Que ste 100 Ulm ala 100 Car gla 25.0 Car gla 26.2 Ulm ala 25.0 Ulm ala 10.6 CAV" Que spp. 64.6 Que spp. 48.5 Que spp. 58.3 Que spp. 55.0 Que spp. 35.4 Fra ame 35.4 Fra ame 17.2 Fra ame 16.7 Corflo 26.7 Ace sac 20.8 Dio vir 10.0 Cer can 8.3 Fra ame 18.3 Fra ame 14.6 Os! vir 10.0 Cor dru 8.3 Jug mg 14.6 Ost vir 8.3 Ulm rub 14.6 CDR Jun vir 40.2 Jun vir 45.3 Ulm ala 32.1 Ulm ala 32.8 Fra ame 10.6 Que ste 21.9 GIB* Car spp. 38.1 Que spp. 47.2 Que spp. 61.8 Que ste 43.7 Que ste 73.3 Car tex 28.6 Ulm ala 28.6 Car tex 17.8 Car spp. 31.0 Car spp. 26.7 Fra ame 14.1 Car (ex 19.1 Ulm ala 10.2 Ulm ala 25.4 PDS Que spp. 54.4 Jun vir 66.7 Jun vir 68.3 Jun vir 56.3 Jun vir 37.5 Jun vir 21.8 Que spp. 25.0 Que mar 18.3 Que spp. 30.9 Que spp. 37.5 Jug nig 15.9 Ulm ala 8.3 Ulm ala 13.3 Car gla 12.7 Dio vir 12.5 Ulm ala 12.5 RND Jun vir 47.2 Ulm ala 48.9 Jun vir 100 Ulm ala 28.2 Jun vir 31.3 Fra ame 15.5 Car gla 19.8 WLC“ Que spp. 45.2 Que spp. 33.0 Que spp. 58.5 Que spp. 36.6 Car ovt 24.7 0st vir 27.5 Ace sac 26.8 Car tex 11.7 Ulm rub 26.7 Cra spp. 24.7 Car ovt 16.3 Car spp. 26.8 Ace sac 10.4 Car ovt 18.3 Que spp. 20.2 Ost vir 18.3 *Managed site. FI-N=Forest Interior-North, TS-N=Transition Zone-North, OP=Opening, TS-S=Transition Zone- South, FI-S=Forest Interior-South. BVL=Berryville Shale Glade, BRN=Brown Shale Barrens, CAV=Cave Creek Limestone Glade, CDR=Cedar Blufl‘ Sandstone Glade, GIB=Gibbons Creek Sandstone Barrens, PDS=Pounds Hollow Sandstone Glade, RND=Round Bluff Sandstone Glade, WLC=Wildcat Bluff Limestone Glade. 114 Table 48 (cont’d). Ace sac =Acer saccharum, Car gla =Carya glabra, Car ovt=Carya ovata, Car spp. =Carya spp., Car tex=Carya texana, Cor dru =C0mus drummondii, Corflo =Cornusflon'da, Cra spp.=Crataegus spp., Dio vir=Diospyros virginiana, Fra ame =Fraxinus amen'cana, Jug nig=Juglans nigra, Jun vir=Juniperus virginiana, Ost vir=Ostrya virginiana, Que alb =Quercus alba, Que coc=Quercus coccinea, Que mar=Quercus marilandica, Que spp. =Quercus spp., Que ste =Quercus stellata, Ulm ala=UImus alata, Ulm rub=UImus rubra. 115 The mean number of tree species (per 25 m2) for all sites and for unmanaged sites was lower in opening than in any other subhabitat except the transition zone-south (Table 49). Managed and umnanaged sites did not differ in tree species number in any subhabitat. Density of forest—opening trees (per 25 m2) ranged from 0.9 i 0.2 to 2.0 i 0.3 and was greater in the northern subhabitats than in the opening (Table 50). Managed and unmanaged site tree density was similar among subhabitats except in the forest interior-north (which was greater for umnanaged sites). The majority of forest-opening trees were shade-intolerant in all subhabitats except the forest interior-south (Table 51). Shade intolerance was greater in the opening (83.4% i 6.7%) than any other subhabitat except the forest interior-north (69.8% i 8.7%). Over 95% (95.6% i 4.4%) of trees in the opening of unmanaged sites were shade-intolerant, versus 74.3% i 9.0% in managed sites. Managed sites exceeded unmanaged sites for shade-intermediate tree values in the opening and both of the northern subhabitats. Forest-opening subhabitats were similar in mean tree diameter at breast height (dbh) (cm) (except the forest interior-south which was lower than the transition zone-north) (Table 52). Managed and unmanaged sites showed overlap of dbh values for all subhabitats as well as little variation among subhabitats. Mean forest-opening tree height (m) (=13.8 m) did not differ among subhabitats (Table 53). Surprisingly, trees which occurred in the opening at managed sites were =10.2 m taller than those in the opening at unmanaged sites. However, unmanaged sites were represented by trees from a single site. Managed sites also exceeded unmanaged sites in tree height in the northern and southern forest interior zones. Mean crown diameter (m) for all sites did not differ among subhabitats (Table 54). At managed sites, crown diameter tended to be greater in one or both of the northern subhabitats than 116 .500m-.0..000. 000.0...Hm-.,.. .500m000N 00...000...Hm-m... .m0.0000un.0 n0:07.000N 00...000....HZ.m... a5.0200020. 000.0“.HZ-.... No H N. Nd H 0.0 No H 0.0 No H 0.. No H m. 00.0.). 000m =< v.0H 0.. NoHfio NOH 0.0 00H ... 00H 0.. .0000 000000.000. 000.). m... H m0 0 m0 H 0.0 0 m0 H 0.. 0. 000.0 000.0000m .005 0003. 00 H 0.. 0 m... H 0.0 0 N... 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Z 3% .000 0000 .0 0.0.0 .00000.. 0.9000000 00. 0. 00.00000. 00)» .. .00.w:0. 003.000 000 0.000-000: 00. :00 0.00.00.00.02: 030.0 00.00.000.000 03. ..0 0w0.0>0 00. 0. .0.0E0.0 030.0 .000. 0. 0000 000.0 00. .0 0.00000000 0>.. 0. .0: 8.00.0.0 030.0 00.. .mm . H. 000.). .3 0.00... 123 in the opening and southern subhabitats. Managed site crown diameter exceeded that for unmanaged sites in the opening and both of the forest interior subhabitats. Cutting of seedlings, shrubs and saplings, and trees at Brown-MGD (in a north-south 5 m x 40 m transect) consisted of the removal of a total of 13 seedlings, 10 shrubs and/or saplings, and 9 trees (Figure 10). The majority of woody species removal occurred in the transition and forest interior subhabitats, not in the opening. Mean diameter at ground level (dgl) (cm) was lowest in the opening for shrubs and saplings (due to an absence of stumps therein) (Table 55). Mean tree dgl was similar in all subhabitats. Statistical Comparisons among Subhabitats: 1994 All of the managed sites were significantly different (0t=0.05) among subhabitats for Poaceae cover while only one of the three unmanaged sites was significantly different for Poaceae cover (i.e., Pounds-UMG) (Table 56). None of the unmanaged sites were significant for Asteraceae cover. While most of the forest-openings were not significant for seedling density, most were significant for cover of exposed rock at the soil surface. Site Open-Area Estimates Total open area per site ranged from 450 to 7825 m2, with a mean of 3956 _+_ 839.1 m2 (Table 57). The managed sites had the four largest open areas. Openings at some of the sites occurred as discontinuous fragments, i.e., Cedar-UMG, Gibbons-MGD, Gyp-UMG, and Round- UMG. Therefore, length (m) estimates are also provided for the longest continuous east-west and north-south site dimensions. The east-west axis of five of eight sites was longer than the north- south axis. Soil Measurements: 1993 and 1994 All soils of forest-opening sites analyzed were a silty loam or loam textural class (Table 58). Together the sites averaged more silt than sand or clay. Unmanaged sites had a greater 124 12 104. — --. .— — — --. Total Number of Stumps I I I I I r \ -_ --- _-__ 0 : . 0 7 0-5 m FI- 5-10 m Fl- 10-15 m 15-20 m 20-25 m 25-30 m 30-35 m 3540 m N N TS-N OP OP OP OP TS-S Distance and Subhabitat + Seedlings + Shrubs/Saplings + Trees Figure 10. Total number of seedling, shrub and sapling and tree stumps versus distance (from north to south) along a transect with four subhabitats at Brown Shale Barrens—Managed in 1994. The subsample transect terminated in a bluff in the transition zone. 125 Table 55. Mean (’0 1 SE) diameter at ground level (cm) of stumps in four subhabitats at Brown Shale Barrens-Managed in 1994. Cut stumps were measured in the subsample transect which terminated in a bluff edge in the transition zone-south subhabitat. FI-N TS-N OP TS—S No. of 25 m2 plots: 2 l 1 Mean stump diameter at ground: Shrubs/Saplings 5.6 4.8 4.4 Trees 10.6 14.5 10.0 FI-N=Forest Interior-North, TS-N=Transition Zone-North, OP=Opening, TS-S=Transition Zone- South. Table 56. Probability values for within-site comparison of Poaceae, Asteraceae, and rock cover and seedling density in 1994. Subhabitat zones with less than five plots were omitted from analysis. The Mann-Whitney U test was used and an asterisk indicates a significant difference between sites (Ot=0.05). Site Zones p-Value Seedling Exposed Used: Poaceae Asteraceae Density Rock Managed Brown Shale Barrens 1-4 0.0002* 0.1112 0.0000* 0.0391* Cave Creek Limestone Glade 1-5 00008“ 0.0000* 0.0507 0.4133 Gibbons Creek Sandstone Barrens 1-5 00353" 0.0000" 0.0869 0.0002‘ Wildcat Bluff Limestone Glade 1-5 00000" 0.1348 0.3023 0.0255" Unmanaged Cedar Bluff Sandstone Glade 1-2 0.0784 0.6001 0.5797 0.0161“ Pounds Hollow Sandstone Glade 1-4 00012" 0.4826 0.1268 NA Round Bluff Sandstone Glade 1-2 0.7375 0.9362 0.0271" 0.0917 Subhabitat Zone l=Forest Interior-North, 2=Transition Zone-North, 3=Opening, 4=Transition Zone-South, 5=Forest Interior-South. NA=Not available. 126 Table 57. Area (m2) and greatest continuous cardinal dimension (m) of the site opening in 1994. Site Area Length N orth-South East-West Managed Brown Shale Barrens 4350 40 170 Cave Creek Limestone Glade 6650 145 110 Gibbons Creek Sandstone Barrens 5675 65 150 Wildcat Bluff Limestone Glade 7825 125 100 Mean i 1 SE (managed sites) 6125 i 737.0 94 i 24.7 133 i 33.0 Unmanaged Berryville Shale Glade 2700 95 45 Cedar Bluff Sandstone Glade 450 8 40 Gyp Williams Sandstone Barrens 725 20 20 Pounds Hollow Sandstone Glade 4250 40 180 Round Bluff Sandstone Glade 2975 60 95 Mean 1': 1 SE (unmanaged sites) 2220 3: 717.3 45 i 15.4 76 i 28.8 All Sites Mean i 1 SE 3956 i 839.1 66 i 15.6 101i 19.3 127 m._ H om m m H or. NM H 6N mm _ H :00: 0,005 ..< cd H QM: md H Woo ad H NLN $0.7. Ema—ESE; mm _ H 5002 E004 bzm o. S cm mc0t0m 0coumncmm 25:23 go anA bzm 2 co mm 0.00—O 0coamv=0m .cEm 3.000 E001— bzm w_ oo mm 0020 0.9% 0_:>>t0m EmmcmEcb w _ H m.mm w.m H m. _ m w.o H mom $05 003525 mm _ H :82 E004 bzm 2 0m _m mc0tam 08.85% 0.005 0:935 E004 mm Nv mm 0020 0:238: 0.00.0 0>00 Ego; bzm mm mo 2 0:055 22m .565 009282 00.0.0 235,—. tan—U «we Em as ...—am Q0 25 29:05“ 3:32 5.. :95 02 0003 0020 0.53254 tam 300:3 0:0 0920 2.88:3 :55 055% .0020 0:20.023 30:0: massed .0 $5.30 05 Eat :0me 836% 6052800 0003 A0920; :0 @000 E0 N._ H W8 00083 :9. 0>E .89 5 00:0 beam 05 08 @0863 0:50. =om .mm 035. 128 proportion of sand than clay due to the higher proportion of unmanaged than managed sites with a sandstone substrate and exposed rock. The limestone glades, Cave-MGD and Wildcat-MGD, had the highest pH values of any forest-opening site, at 7.4 and 6.9, respectively (Table 59). The average pH for the remaining sites was 4.9 i 0.2. Cation exchange capacity (CEC) was also highest at the two limestone glades. Mean soil depth for all forest-openings (managed and unmanaged) did not differ between 1988 and 1993 (Table 60). Forest-opening subhabitats did not differ in mean soil depth (cm) (Table 61). Unmanaged sites exceeded managed sites for depth in the opening; however, the unmanaged comparison value was based on a single site. The opening of unmanaged sites also had the greatest soil depth among unmanaged site subhabitats. Mean soil moisture (z14.5%) for all of the study sites, did not differ among subhabitats. All managed and unmanaged site subhabitats were similar in percent soil moisture (Table 62). Climatic Data: 1993 and 1994 Mean monthly temperature (°C) during the spring and growing season of 1988 and 1993 was consistent with the 1910-1993 average (Figure 11). Monthly precipitation values (cm) (1988 and 1993), however, diverged greatly from the 1910-1993 mean during the growing season (Figure 12). From April through August, 1988 rainfall was below the 1910-1993 average. In 1988, only during March 1988 did precipitation surpass the 1910-1993 average. Conversely, in June and July of 1993, precipitation exceeded average rainfall. The difference between June 1988 and 1993 rainfall was 16.9 cm (6.6 inches). 129 Table 59. Summary of soil characteristics for the study sites in 1993. Nutrient units are in parts per million. Samples taken from the opening at Pounds Hollow Sandstone Glade were too small for analysis. Character BVI-I BRN" CAV“ CDR GIB“ GYP RND WLC“ pH 5.4 5.2 7.4 4.8 4.4 4.5 5.2 6.9 Buffer pH 6.70 6.53 NA 6.48 6.49 6.56 6.82 NA Phosphorus 8 5 5 5 11 5 9 6 Potassium 100 104 280 78 64 72 70 346 Calcium 1060 l 130 6000 740 280 290 400 4860 Magnesium 65 142 94 92 52 72 74 145 Manganese 70 89 42 1 18 72 74 208 157 Zinc 3.0 3.1 0.4 3.7 1.6 2.3 3.3 5.5 %Org. Matter 7.4 4.0 5.6 3.9 3.0 2.6 7.9 6.7 ENR 192 124 156 122 104 96 202 178 CEC 8.5 10.8 31.5 8.8 6.1 5.8 4.3 26.8 *Managed site. BVH=Berryville Shale Glade, BRN=Brown Shale Barrens, CAV=Cave Creek Limestone Glade, CDR=Cedar Bluff Sandstone Glade, GIB=Gibbons Creek Sandstone Barrens, GYP=Gyp Williams Sandstone Barrens, RND=Round Bluff Sandstone Glade, WLC=Wildcat Blufl‘ Limestone Glade. ENR=Estimated nitrogen release, CEC=Cation exchange capacity (meq/ 100g). NA=Not available. 130 Table 60. Mean (i 1 SE) soil depth (cm) for the study sites in 1988 and 1993. Five soil depth measurements were taken at all of the sites. Values for 1988 are from Heikens (1991). Standard error values for 1988 data were not available. Site 1988 1993 Difference Managed Brown Shale Barrens 7.9 12.4 i 4.9 4.5 Cave Creek Limestone Glade 6.8 13,1 i 29 6.3 Gibbons Creek Sandstone Barrens 9.5 9,1 t 2.8 -0.4 Wildcat Bluff Limestone Glade 8.6 2.6 i 1.2 -6.0 Mean (managed sites) 8.2 i 0.6 9.3 i 2.4 1.1 Unmanaged Berryville Shale Glade 8.5 4.2 i 0.7 -4.3 Cedar Bluff Sandstone Glade 5.7 18.4 i 2.9 12.7 Gyp Williams Sandstone Barrens 10.8 1 1.1 i 4.5 0.3 Pounds Hollow Sandstone Glade 4.0 1.0 i 0.7 -3 .0 Round Bluff Sandstone Glade 4.3 2.7 i 0.3 -l .6 Mean (unmanaged sites) 6.7 i 1.3 7.5 i 3.2 0.8 All Sites Mean 7.3 i 0.8 8.3 i 1.9 1.0 131 Wm H 0... Em H c.» Em H o... w. H m... N... H 0.x 200.2 00.5 :< 06 H No 0.0 H wh o.o H w...— oN H 0.0. 0.. H 0.0 00.0 v0w0c0Ecsv c002 --- --- --- 0.0 H 0.0. N. H N. _ 0.00.0 0:20....0m .cEm venom m. H 0.0 0.005 0:200:0m tam .0000 ... H No Yo H w.m m... 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Source: Midwestern Climate Center for Carbondale, Illinois. 134 30 25 4. 0 2 » 4 5 1 10 «- 020.00 00500 Mar Apr May Jun Jul Aug Sep Oct Nov lec Feb Month l l I | 1910-1993 (+ 1 SE) : 135 30 25 .. 35 20 -» (D g, as 0 ‘, E 15 I: C . ' a; . o ‘I 10 «~ 9 x ‘ N 5 O # ‘r t i § # , . t 1 Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec Months —o—1988 ----x --1993 -n—191o-1993 (+ 1 SE) Figure 12. Total precipitation (cm) versus month in southern Illinois for three different periods of time. Source: Midwestern Climate Center for Carbondale, Illinois. DISCUSSION Vegetation Composition Herb Species Composition The reasons for the changes in herb species composition between 1988 (Heikens 1991) and 1993-94 were difficult to elucidate as they were confounded by various factors, including the effects of the drought of 1988, management treatment, successional changes, and differences in project sampling emphasis. Heikens’s (1991) sampling and floristic surveys were confined to the open area of each studied site (A. Heikens, pers. communication). As the 1993-94 floristic surveys included the openings and an 8.5 m border area, these floristic survey findings are not directly comparable. However, plot sampling for both years was confined to the Opening. Therefore, a better between-year comparison is made using the total number of herb species found within plots (Table 8) and the average number of herb species per plot (up to 30 50-m2 plots were used for herbs in 1988 versus four nested l-m2 plots located within each of the same number of 50-m2 plots in 1993) (Table 11). Plot sampling data showed an overall increase of 36 herb species in all plots collectively between years, despite the greater total area sampled in 1988 (Table 8). The total number of herb species increased both over time and after management. The mean number of herb species per plot also increased from 11.8 i 1.7 to 16.3 i 1.6 over time (Table 11). The smaller plots used in 1993 may have facilitated the observation of small and sparse herbs at the expense of missing a greater variety of microhabitats (and therefore different species) potentially encountered in the larger plots used in the earlier study. Using fewer but larger plots might decrease the amount of time spent in sampling herb species; however, difficulty of accurately assessing herb cover, especially of small or sparse individuals, is an inevitable consequence for a 50 m2 plot size for herbs. 136 137 Herb species number was greater at managed sites than at unmanaged sites in both 1988 (before management) and 1993 (after management) (Table 8). Percent increase of herb number between sampling periods for unmanaged sites, 58.4% exceeded that for managed sites, 23.5%, when 50 m2 plots (1988) were compared with four nested l-m2 plots (1993). As the increase in species number was not simply a response to management treatment, temporal differences in herb composition may possibly be attributed to climate factors. The 1988 growing-season was characterized by a severe drought, “...one of the worst in the century,” (Anonymous 1988) during which precipitation was 41.1% (April-August) lower than average (1910-1993) in southern Illinois (Figure 4). In contrast, 1993 growing-season precipitation was 11.6% above average, and rainfall for June was 16.9 cm higher than in 1988. Piper (1995), studying four tallgrass prairie sites in Kansas found that, in general, species number was lowest in the drought year (1989) and higher before and afterward (1986 to 1992). Tilman and El Haddi (1992) reported that the number of species in native prairie and European-grass fields in Minnesota declined 37% in response to the 1988 drought. Furthermore, a major portion of this loss was found to be annual species (up to 7.3 per 20 x 60 m field) and species loss was independent of pre-drought abundance. The loss of perennial species was inversely correlated with pre-drought abundance. To determine whether the proportion of annuals was lower in 1988 than in 1993, the percent life history composition of two sites, Cave-MGD and Pounds-UMG, was calculated for 1988. In 1988 the proportion of annuals was 2.4% for Cave-MGD (20.5% in 1993) and 19.4% for Pounds-UMG (25.0% in 1993) (Table 14). Therefore, both sites did have a lower (5.6% to 18.1%) proportion of annual species during conditions of severe moisture-stress than afiemard. However, without data on pre—drought herb composition, the effect of drought on herb composition, if any, is impossible to ascertain. An increase in herbaceous cover at the unmanaged sites (173.9%) and not at the managed sites (the means were within one standard error of each other) may indicate succession in the herb layer (Table 12). According to Barbour et a1. (1980), vegetative cover generally increases with 138 increasing succession. Likewise, forest-opening sites which appeared to be incorporating woodland herb species into the top three positions of relative importance (RI) (Table 10), were also those which exhibited the greatest increase in herb cover between sample periods; for example, the longterm unmanaged sites (i.e., no anthropogenic fire since c. 1930 in southern Illinois, Anderson and Schwegman 1991), Cedar and Gyp, both showed an increase in herb cover in the opening, together averaging a 260.9% increase in cover (versus the mean for all unmanaged sites, 173.9%), as well as an increase in RI position of broadleaf woodland herbs such as Parthenocissus quinquefolia, Toxicodendron radicans, or of successional, open-woods species like Helianthus divan'catus and Solidago sp. Anderson and Schwegman (1991) noted an increase in the frequency of woodland herbs with succession in a mesic southern Illinois barrens (sampled 15 years postfire) and a decrease in frequency of prairie species (e. g., Schizachyrium scoparium and Sorghastrum nutans). Hardin (1988) similarly reported an increase in cover and frequency of forest interior herb species after 22 years of succession in a prairie inclusion in southeastern Ohio. Still, the 1993 cover values of herbs (28.5% i 4.5%) were much lower than values reported for similar habitats in the literature. For example, Jeffries (1987) found an average herb cover of 52.6% for three sandstone glades in Arkansas. Adjacent to the glade proper at Cedar-UMG, Pounds-UMG and Round-UMG there was a dense, even-aged stand (a “dog-hair” stand) of woody species in. the sapling to small tree size class. The presence of a few sporadic prairie species within this area (e.g., Liam's sp. at Pounds-UMG) suggests that succession had progressed to the point that the opening was relegated to the rock outcrop alone. Poaceae RI declined more between sample periods at managed sites (-11.8%) than at umnanaged sites (-6.3%) (Table 9). Schizachyrium scoparium, a characteristic prairie grass, appeared to be particularly sensitive to management and was on average 10.4% lower in R1 for managed sites in 1993 (Table 10, Appendix A). Asteraceae in the managed sites also decreased in RI (-2.0%), while increasing in R1 in the unmanaged sites (2.6%) (Table 9, Appendix A). The 139 increase in Asteraceae R1 at unmanaged sites from 1988 to 1993 and decrease in characteristic prairie Poaceae species at the managed sites with time (averaging 4.8 years postfire for the 1993 sampling) was consistent with studies by Zimmerman and Kucera (1977), Towne and Owensby (1984), Abrams et al. (1986), and Hartnett (1991) report that nongraminoid species increase (e.g., biomass, number of stems) with time after disturbance. Piper (1995) stated that there was an inverse relationship between productivity and species diversity (diversity primarily being a function of the number of forb species) in grasslands. As total biomass of the dominant species decreases, competition between grasses and forbs presumably decreases and species richness and evenness increase. According to Dhillion and Anderson (1994), grass productivity was higher on burned than on unburned sites during the growing season following the fire in both drought and nondrought years. Svejcar (1990) found aboveground biomass to achieve levels comparable to unburned prairie by the fall after a spring burn. Risser et a1. (1981) and Collins and Gibson (1990) also state that cover and productivity return to prebum levels in one to two years following fire in tallgrass prairie. A model by Gibson and Hulbert (1987) predicts a peak in diversity six to seven years following fire after which diversity declines. Therefore, a thorough assessment of forest-opening productivity and species diversity in response to management may shed light on the hypothesis regarding dominance/productivity and competition cycles in forest-Openings. Even so, the Asteraceae or Poaceae remained in the first RI position for all sites in both years and for seven of nine sites, the same family remained in the first position both years. Three of four managed sites (excepting Cave-MGD) exhibited an increase in RI (30.5%) of the Fabaceae, whereas four of five unmanaged sites (excepting Round-UMG) showed a decrease in RI (466%) for this family between 1988 and 1993 (Appendix A). This complements the findings of Martin et al. (1975), Anderson and VanValkenburg (1977), and Towne and Knapp (1996), who found increases in legumes (e. g., density, biomass) afier fire management. 140 Statistical Comparisons: 1988 and 1993. Mann-Whitney U tests showed significant differences between years for Poaceae and Asteraceae cover (Table 25), in all sites except Berryville, a xeric unmanaged glade which did not show signs of undergoing successional change as seen in rank of percent similarity values (Table 17), woody species RI (Table 21) or herb species RI (Table 10). Between-year differences in herb cover (Table 12), presumably due to succession, may be responsible for the nearly uniform significance results for Poaceae and Asteraceae. To test for differences in Poaceae and Asteraceae cover between managed and unmanaged sites, the Gibbons-MGD and Gyp-UMG data sets were compared (Table 26). Gibbons-MGD and Gyp-UMG are both sandstone barrens which occur within five miles of each other and differ primarily in management history. These sites showed significant differences in Poaceae and Asteraceae cover but not in rock cover in 1993. In 1993, Gibbons, which was fire-managed in 1989, was visually more open with a nearly continuous graminoid layer. In 1993, Gyp-UMG resembled an Open forest with isolated patches of Schizachyrium scoparium and Sorghastrum nutans under canopy openings roughly 5 x 10 m. Poaceae R1 was 12.9% higher at Gibbons-MGD than at Gyp-UMG in 1993, but Asteraceae RI was 10% lower (Table 9). This further supports the contention that increases in Asteraceae and decreases in Poaceae occur with successional progression. Two unmanaged sites, Cedar and Round, were also compared to each other for Poaceae and Asteraceae cover and no significant differences were found (Table 26). Both sites are unmanaged sandstone glades which occur within ten miles of each other. At both glades, Poaceae and Asteraceae cover was sparse, and consisted largely of isolated clumps of Danthom‘a spicata or Schizachyrium scoparium with few specimens of Asteraceae. Site Similarity. Although 34 of the 1988 opening herb species (74% of the total for 1988) were relocated in the opening and/or transition in 1993-94 (Table 17), all of the possible combinations of forest—opening site pairs were more alike in 1994 (24.8% i 0.8%) (Table 18) than the same site compared over time (1988 and 1993-94) (19.3% i 1.1%) (Table 17). Presumably then, drought I41 and successional change were more important in determining herb species similarity than differences in management, substrate or forest-opening type (Table 18). Also, since the transition zone was included in the 1993-94 surveys and not in 1988, sampling methods may have distinguished the sampling years. Managed sites and managed site pairs had a greater number of common herbs than unmanaged sites. However, sites of a common classification (i.e., the same substrate and forest-opening type) were more similar (30.8% i 1.4%) than sites with the same management history. These sites were close in proximity, i.e., Gibbons-MGD and Gyp-MGD, Cedar-UMG and Round-UMG, Cave-MGD and Wildcat-MGD (Figure l), but did not necessarily have similar (or any) management histories. Compared to herb species (11), more woody species (16) were shared among all forest- opening sites (Table 19). All of these woody and herbaceous taxa are typical of the oak-hickory forests of this region. Exotic plants were also found at all of the study sites (11.7 on average) despite the fact that a few sites, i.e., Gyp-UMG and Wildcat-MGD, were surrounded by forest and were not easily or frequently accessed by humans (Table 20). The number of exotics appeared to correlate strongly with proximity and/or frequency of human visitation rather than with site management history (if any), forest-opening type or substrate. In most instances, forest-opening exotics were found in small isolated patches or as solitary individuals. However, a few exceptions existed at Cave-MGD which had more exotics (31) than any other study site. F estuca amndinacea was common along the road at Cave-MGD, and appeared to be extending north into the glade. Robust individuals of Campsis radicans also appeared to be successfully established at the site. In addition, Medicago lupulma and Kummerowia stipulacea occurred at the southern perimeter of the opening but have been known to increase with fire-disturbance (Van Valkenburg 1977, Thompson and Heineke 1977). Although infrequent, individuals of Verbascum thapsus grew to >12 m and bore large seed heads. Only a single individual of Melilotus alba (Fabaceae) was found in the glade proper at Cave-MGD. 142 The unmanaged sites bore a suite of small, solitary exotics most likely introduced by hikers and campers (and horse riders at Cedar-UMG). Only two exotic species were found at Cedar- UMG in 1988 versus 19 in 1994. For example, at Cedar-MGD heavy visitor use in 1993 exposed sizable (>5 m2) patches of bare soil. Exotics increased from two to 11 at Pounds-UMG, from one to 14 at Round-UMG and from zero to eight at Berryville-UMG. The managed sites, Brown, Gibbons, and Wildcat, also contained relatively few exotic species with eleven, five and four species, respectively. The exotics at these sites were uncommon except for Viola raphanesquii, a tiny spring ephemeral, found throughout the opening at Gibbons- MGD and an extensive ground cover of Corom‘lla varia at Brown-MGD found along the northeastern slope overlooking the roadway. Categorization of Herb Species Grasses, Forbs, Legumes and Exotics. The variability in the proportion of grasses, forbs, legumes and exotics was unexpectedly low among forest-opening sites in 1993 (Table 13). For example, the percentage of legume species varied little among sites (9.2% i 0.9%), despite their disparate management histories. Other studies of Midwestern forest-openings have found dramatic increases in native and exotic legumes after management (e. g., Martin et al. 1975, Anderson and VanValkenburg 1977). However, these effects were immediate and not studied beyond the first year postfire. The site with the greatest percentage of legumes, Gyp, was a long- term unmanaged site. The sampled forest-openings contained 9.2% i 0.9% legumes and 17.3% i 1.2% grasses. In comparison, tallgrass prairies contain a similar percentage of legumes (9.2%), but a lower proportion of grass species (10%) (Curtis 1959). Life History. The percentages of annuals, biennials and perennials (for managed and unmanaged sites and on a site by site basis) were unexpectedly similar, with the exception of Wildcat-MGD (Table 14). Wildcat-MGD had the highest percentage of perennials (89.8%) and had also received more bum management treatments (3) than any other site. However, Gyp-UMG 143 also had a high percentage of perennials (84.0%) but had received no management. The sampled forest-openings differed in the estimate for perennials of Risser et a1. (1981) (95%) for the tallgrass prairie by about 20%, averaging 75.8% i 2.5%. Due to their xeric nature, the southern Illinois forest-openings examined in this study had a substantial component of annuals, averaging 20.6% i 2.7% of all species. Raunkaier’s Life Forms. According to Walter (1973), “most” prairie and steppe species in temperate climates are hemicryptophytes. However, in the sampled forest-openings, hemicryptophytes and geophytes were found in comparable proportions, at 38.0% i 5.6% and 40.4% _+_ 3.8%, respectively (Table 15). In fact, managed sites had a greater pr0portion of geophytes than hemicryptophytes and a higher percentage of geophytes than unmanaged sites. In light of Walter (1973), the high percentage of geophytes may be a consequence of the high proportion of woodland species found in these forest-openings. Bray (1957) attributed the occurrence of 21 non-aggressive climax forest herbs (e.g., Claytonia virginica, Erythrom‘um albidum, Trillium recurvatum) located in Middle West prairies to their geophytic physiognomy which allowed them to escape late spring fires through regeneration from underground parts. When the forest-opening herbs were categorized by habitat, 47.7 i 6.6% of the species had a woodland habitat affinity (Table 16). In addition, 10.8% of herbs were characteristic of an open woods habitat, giving these forest-openings a composite 58.5% woodland/open woods type herb flora. However, when geophytes from the most species-rich site, Cave-MGD, were analyzed by habitat, 42.6% were associated with woodlands and 11.3% with open woods, about the same proportions as would be expected from a random sample of all forest-opening herbs. The high percentage of geOphytes, therefore, was not attributable to the high proportion of woodland herbs in the forest-openings. In addition, many authors (e.g., Anderson 1982, Anderson 1990, and Seastedt and Ramundo 1990) underscore the importance of rhizomatous (geophytic) herb species in regularly burned tallgrass prairie. 144 Categorization by Habitat Although managed sites exceeded unmanaged sites in the proportion of herb species typically occurring in a prairie habitat, the difference was small (2.9%) (Table 16). Overall, the proportion of species occurring in each habitat type was unexpectedly similar for managed and unmanaged sites. The barrens flora has been described as a mixture of forest and prairie species (White and Madany 1978, Packard 1990, Hutchison 1994). For example, in Wisconsin, oak savanna had a similarity index of 50% to 58% with prairie, and 53% with dry oak forest (Curtis 195 9). However, the forest-opening herb flora was primarily typical of woodlands (47.7% i 2.2%) with disturbed, bluff, open, and prairie habitat association percentages between five and 15%. Heikens’s (1991) dichotomous key separates barrens, limestone glades, and loess hill prairie from sandstone glades, shale glades, and open forests as sites with _>_10% prairie species, such as Aster sp., Brickellia eupatorioides, Euphorbia corollata, Helianthus sp., Lespedeza sp., Schizachyrium scoparium, Silphium terebinthinaceum and Solidago sp. Categorization of the sampled forest-openings into habitat types supports Heikens’s (1991) key for limestone glades with 12.9% characteristic prairie species, and sandstone and shale glades with 6.6%, but not barrens which averaged 7.3%. The forest-openings were intermediate between the hill prairie and hardwood forest for species affinity to a woodland habitat and had a larger proportion of prairie associate species than the hardwood forest (Figure 9). This also supports the assertion that forest-openings harbor a flora that is transitional between forest and prairie. Woody Species Composition Many of the same overstory species were found at the study sites in the same RI position in 1988 and 1993 despite the different methodologies used to estimate canopy cover (Table 21). As indicated in the key by Heikens (1991), different forest—opening types were characterized by particular woody species. Both limestone glades were dominated by Quercus prinoides var. acuminata (yellow chestnut oak), all sandstone glades by Juniperus virginiana (eastern redcedar) 145 and the barrens and shale glade by Quercus stellata (post oak) for both years. Results for the sandstone glades concur with Jeffries (1987) who also determined Juniperus virginiana to be the most important tree species in three Arkansas sandstone glades. Consecutive burns at Wildcat (spring 1982, spring 1988, fall 1990) undoubtedly eliminated Juniperus virginiana, a fire-sensitive species, as no individuals were found in 1993 although Heikens (1991) noted it in 9 of 18 plots (RI= 14.5%) in 1988. Anderson and Schwegman (1991) found that redcedar was eliminated from a mesic southern Illinois barrens after one fire. A greater number of woody species (per 50 m2) were present in 1993 than in 1988 at both managed and unmanaged sites (Table 23). No difference in woody species number was found between managed and unmanaged sites in 1993, whereas unmanaged sites had more woody species than managed sites in 1988. A greater number of species of tree seedlings were encountered in the forest interior subhabitats (and transition zone-north) than in the opening of managed sites in 1994 (Table 40). Areas with permanent plot stakes from the 1988 sampling in some instances overlapped area defined as transition in 1994 (see Appendix B: Figures 8f-h). These finds suggest that succession was occurring at both managed and unmanaged sites for woody species (versus herb succession which occurred at unmanaged sites only). The length of time since management (averaging 4.8 years at the 1993 sampling) may have been long enough to allow woody species establishment. Data from a mesic barrens in southern Illinois suggests that management encourages resprouting and woody growth. Anderson and Schwegman (1991) found an increase (216,000 stems/ha to 30,000 stems/ha) in seedling stems (between 1.4 m and 31.2 cm dbh) one year after fire cessation. Shearin et al. (1972) reported enhanced germination of Liriodendron rulipifera in hardwood and pine plantations in South Carolina for three growing seasons following a prescribed burn. The between-year increase in woody species may also involve the survival rate of tree seedlings through the severe drought of 1988. Keener (1983) noted the desiccation and mortality of nearly 35,000 seedlings (albeit herbacous) in a 225 mZ-barrens between April/May 146 and mid-June (nine seedlings survived). Mesophytic tree seedlings, observed in 1990 (two years post-drought) by Heikens et al. (1994) at Brown-MGD (i.e., Acer saccharum), were not observed in 1988 (Heikens 1991). Between-site canopy cover comparisons were not significantly different for managed- unmanaged and unmanaged-unmanaged site combinations in 1993 (Table 26). It was unexpected that the Gibbons-MGD and Gyp-UMG canopy comparison was not significant as Gyp-UMG had few canopy openings whereas Gibbons-MGD was a visibly open mosaic of openings (>5 m2) and canopy. However, Gibbons-MGD had only recently received a single burn (fall 1989) and the shrub and sapling layer appeared vigorous in 1993. Rock Cover The mean percentage of exposed rock for all forest-openings decreased 5.8% between 1988 and 1993 (Table 24). Since average herb cover was 11.9% higher in 1993 than in 1988 (Table 12), a concomitant decrease in exposed rock presumably resulted from colonization by plants, especially those successional species which spread by runners, covering the rock surface and trapping litter (e. g., Parthenocissus quinquefolia, T oxicodendron radicans, and Antennaria plantaginifolia). The sites with the greatest estimated percent increase in herb cover (i.e., Berryville—UMG, Cedar-UMG, Gibbons-MGD, Gyp-UMG, and Round-UMG), were also among sites with the greatest decline in rock cover since 1988. This generalization however, was not true for Cave (burned in 1986) which had a decrease in herb cover and did not have a build up of litter in the opening in 1993-94. Barrens had less exposed rock than glades in 1993 (at 0.8% and 21.3%, respectively) (Table 24). This result concurs with the key by Heikens (1991), in which barrens have 55% exposed rock cover, (averaging 3.1% for 4 barrens sites in Heikens 1991) and glades have at least five percent cover of exposed rock (averaging 22.3% for 12 sites in Heikens 1991). Therefore, the 147 ratio of barrens to glades, 2:2 for managed sites and 1:4 for unmanaged sites, undoubtedly contributed to the greater mean percentage of exposed rock for unmanaged sites in both years. Between-year statistical comparisons of percent cover of exposed rock were significantly different for two of the managed sites (Cave and Gibbons) and two unmanaged sites (Cedar and Gyp) (Table 25). Cave-MGD and Gibbons-MGD were burned one year prior to sampling while the other managed sites, burned three years earlier, were sampled afier a longer postfire period. Fire management at Cave and Gibbons may have caused significant change in the herb layer (e.g., increased dominance by grasses) and decreased litter, thereby altering exposure of the surface rock. The absence of fire may have contributed to a combination of herb growth and litter buildup responsible for the significant change in rock cover at Cedar-UMG and Gyp-UMG. Significantly different percentages of rock cover between 1988 and 1993 at Cave-MGD (burned in 1989) may be attributable to the decrease in herb cover (Table 12). The decrease in herb cover since 1988, however, is difficult to explain as Gibbons-MGD was burned in 1989 as well and showed an increase in herbaceous cover. However, Gibbons-MGD is a more mesic site with greater (30.8%) canopy cover (Table 22) whereas Cave-MGD is a xeric, more exposed site. Significant differences in percent cover of rock cover between two unmanaged sites in 1993 may be attributed to the fact that at Cedar, no extensive rock shelves existed, whereas at Round a 5- to 15-m shelf extended along about 80 m of the glade (Table 26). Despite the fall 1989 burn at Gibbons, rock cover values between sites (Gibbons-MGD and Gyp-UMG) were not significantly different in 1993 (Table 26). Both sites showed significant decreases in rock cover since 1988 (average loss of 3.5%) (Table 25). Although Gyp-UMG appeared to be undergoing herb and overstory succession, exposed stones and bedrock were still evident at the soil surface. This may be due to its hilltop location and to weathering. Fire management did appear to remove litter on and around surface rock at Gibbons-MGD. Therefore, it is not entirely clear why these sites were not significantly different. 148 Vegetation Subhabitats Herb Species Composition Herbaceous species distribution patterns appeared to correspond with spatial gradients in the forest-openings (Table 27). These changes were especially pronounced at the managed sites which had large continuous tracts of open area. For example, Cave-MGD had an exemplary continuum of species. An Open area approximately 145 x 110 m (north-south x east-west) (Table 57) graded from dry woods at the northern hilltop through the opening to mesic woodland in the south. Species of dry woods, such as Helianthus divaricarus, Solidago ulmifolia, and Parthenocissus quinquefolia, important in the forest interior-north, were replaced by Smilax bona- nox (catbrier) in the transition zone-north. The opening was dominated by prairie forbs such as Brickellia eupatoriodes (false boneset) and Silphium terebinthinaceum (prairie-dock) interspersed with Schizachyrium scoparium, Bouteloua curtipendula, and Carex sp. The transition zone-south contained a mixture of woodland (e. g., Parthenocissus quinquefolia) and prairie species (e.g., Ratibida pinnata). Finally, the forest interior-south was dominated by woodland species such as Verbesina virginica (tickweed) and T oxicodendron radicans. Just as managed sites exceeded unmanaged sites in the mean number of herb species (per plot) in both 1988 and 1993 (Table 11), managed sites exceeded umnanaged sites in herb species richness (the number of species per m2) in all subhabitats in 1994 (except the transition zone-north) (Table 31). While there was no difference in the number of herb species among subhabitats for managed sites, the opening subhabitat (and transition zone-south) was lowest in species richness for unmanaged sites. This may suggest that management was effective in maintaining (or enhancing) species richness in the opening of managed sites. It may also indicate that herb species richness in the Opening subhabitat of the unmanaged sites (Berryville, Pounds, and Round) was typically lower than other subhabitats because the Openings of these glades were characterized by high insolation, exposed rock “pavement” and a few herb species adapted to tolerate moisture- 149 stress such as spring ephemerals and succulents (e.g., Opuntia sp., Sedum pulchellum and T alinum parviflorum). In accord with the findings for herb cover in 1988 and 1993 (T able 12), in 1994, managed site herb cover exceeded unmanaged site cover, and did so in all subhabitats (Table 32). Managed site herb cover was highest in the opening. Conversely, herb cover did not peak in the Opening of unmanaged sites, suggesting that management was effective in maintaining (if not enhancing) relatively dense herb cover in the opening. Because the Opening of the unmanaged glade sites (all four umnanaged sites were glades) was strongly influenced by the substrate, herb cover might be expected to trough therein. However, herb cover was lower in the transition zone-south than all other subhabitats. This may reflect the importance of the southern exposure (insolation and moisture stress) adjoining the characteristic glade “pavement”. Although the R1 of forest-Opening Poaceae was greater in the Opening (and forest interior- north) of unmanaged sites than of managed sites, inspection of RI on a site by site basis showed peaks in abundance of Poaceae in all subhabitats except the forest interior-south (Table 28). Based on field observations, Poaceae RI seemed to track topographic moisture gradients more than any other factor, increasing in dry areas. For example, high exposure, raised topography, and xerophytic vegetation in the opening and northern subhabitats were visible features at Berryville- UMG and Poaceae RI averaged 36.2% higher in these subhabitats than in the transition zone-south and forest interior-south. Similarly, the opening and southern subhabitats at Brown-MGD appeared to be more xeric than transition zone-north and forest interior-north, and were also zones of highest Poaceae RI, differing by an average of 17.8%. Preliminary soil moisture results, however, did not support this hypothesis (Table 62). Most sites showed moisture trends which did not correspond with Poaceae R1 in a positive or negative way. However, far more extensive testing of soil moisture, especially of seasonal variation across the subhabitats, would be necessary to establish its effect on herb species distribution. 150 Although all forest-opening subhabitats were within one standard error of each other (=16 to 24%) for Asteraceae RI (Table 31), mean Asteraceae R1 was higher in the opening (and transition zone-north) of managed sites than of unmanaged sites. According to a model by Gibson and Hulbert (1987) for the tallgrass prairie, diversity (primarily determined by the number of forb species) could be expected to increase for six or seven years afler fire, after which diversity declines. Since management occurred an average of 3.8 years prior to the 1994 sampling, fire may have maintained species richness in the opening (Table 31) as well as herb species diversity (Table 37) as, in part, reflected in Asteraceae RI values relative to the unmanaged sites. Relative importance for Cyperaceae, as for Poaceae, was greater in the opening subhabitat of unmanaged sites than of managed sites (Table 30). However, unlike the grasses, sedges tended to dominate in subhabitats which were relatively mesic. For example, at Berryville-UMG sedges increased in the transition zone-south and forest interior-south which were lower topographically and more mesic than in the north. At Brown-MGD the transition zone-north and forest interior- north which were canopied, had higher sedge RI values (averaging 10.9%) than the exposed and drier southern site perimeter (5.2%). In general, broadleaf species characteristic of the forest interior (e.g., Parthenocissus quinquefolia, T oxicodendron radicans, and Galium sp.) were absent among the top three RI positions in the Opening, appearing in the transition and forest interior subhabitats (Table 27). At Cave-MGD and Wildcat-MGD (sites which had been burned more than once prior to 1994) the herb species in the first RI position in the opening was not ubiquitous in the oak-hickory forests of southern Illinois and was not prevalent in adjacent subhabitats. At Cave-MGD Brickellia eupatorioides was 25.2% more important in the opening than the species in the second RI position and therefore may serve as an opening indicator species. Furthermore, this species was not found in sample plots outside the opening subhabitat. At Wildcat-MGD Silphium terebinthinaceum did occur in the transition zone-south (1.5%) (Appendix C); however, R1 was 9.9% higher in the 151 opening (11.4%). Therefore, Silphium terebinthinaceum may also serve as an indicator of the opening subhabitat at Wildcat-MOD. Danthom‘a spicata of dry woods and bluffs, Dichanthelium laxiflorum of woodlands and xeric Carex sp. (probably C. artitecta of dry woods and C. umbellata of dry woods and sandstone rocks) accounted for 21.4% of the total opening R1 for herbaceous species, while Schizachyn’um scoparium, a characteristic prairie species, accounted for 8.0% of the total opening RI (Table 27, Appendix C). Therefore, unusual grass and sedge species appeared to be important forest-opening matrix species which occur within the Andropogon-Panicum-Sorghastrum association of the Midwestern tallgrass prairie (Risser et al. 1981). Categorization of Herb Species Raunkaier’s Life Forms. The majority of the life form category values for managed and unmanaged sites (and across subhabitats) did not differ statistically in 1994 (Table 36). The most important life forms were the geophytes (averaging 38.4% of total life form R1 for herbs across subhabitats) and hemicryptophytes (averaging 37.7% across subhabitats). The ratio of forest— opening geophytes to hemicryptophytes was compared to that of a hill prairie in southern Illinois (38 herb species, Voigt and Mohlenbrock 1964) and a mesic mixed hardwood forest in east-central Illinois (21 herb species, Aikman and Ebinger 1991) to better ascertain typical values for each of these habitats in this region. The forest-opening ratio (geophyteszhemicryptophytes) was 1.02, the prairie, 1.15, and the forest, 3.76. Therefore, the forest-opening herb life form ratio was proportionately more similar to a prairie habitat than to a mesic forest. In this study, the proportion of geophytes in the opening of managed sites was 34.1% greater than that of unmanaged sites with a ratio of 1.20, comparable to the estimate for the hill prairie. Herb Species Diversity Although mean forest-opening herb diversity using the Simpson (D5) and Shannon (H’) indexes did not differ among subhabitats for all sites, consistent differences were observed between 152 managed and unmanaged sites (Table 37). Managed site D5 and H’ diversity exceeded that of unmanaged sites in the opening and southern subhabitats (and in the transition zone-north for H’). Species richness was also greater in the opening and southern subhabitats (and forest interior- north) for managed sites than for unmanaged sites (Table 31). As H’ is influenced more by species richness than Ds, it followed expectation that managed sites exceeded unmanaged sites in diversity in the opening and southern subhabitats. D3 is more sensitive to relative abundance of species (dominance and evenness) and was also greater in the opening and southern subhabitats of managed sites, providing further information about the diversity in these subhabitats. A study of diversity in mixed-grass prairie by Collins and Barber (1985) concurs with Denslow’s (1980) model, more specific than the intermediate-disturbance hypothesis, in which diversity is maximized under a natural disturbance regime and minimized under undisturbed or severely disturbed conditions. The managed sites in this study did have higher diversity in the opening than unmanaged sites. However, a study by Collins and Adams (1983) reported no trends in diversity, evenness, or species richness in the course of succession of mature tallgrass prairie in Oklahoma to shrub-grassland. Woody Species Composition Shade Tolerance. In all three woody strata (seedling, shrub/sapling and tree) the proportion of shade-intolerant species in the opening of unmanaged sites exceeded that of managed sites (Tables 42, 46 and 51). The ratio of glades to barrens was greater for unmanaged (4:0) than for managed sites (2:2). This may indicate that the opening substrate of unmanaged sites was more important in determining woody species composition (shade tolerance) than management. Counter to expectation, however, on average, the shade-intolerant RI values for the opening of all of the glades were comparable to barrens (shrubs and saplings) or lower (seedlings and trees). However, the sandstone and shale glades which comprised the unmanaged site comparison had more exposed rock (22.9%) (Table 33) and a more shallow soil depth (19.7%) (Table 60) in the opening than the 153 managed limestone glades and averaged 91.8% higher in the proportion of shade-intolerant woody species (seedlings, shrubs/saplings and trees) in the opening. Therefore, the xeric nature of the opening of the unmanaged glades may account for the high proportion of shade-intolerant species relative to the managed sites. Alternatively, Anderson and Schwegman (1991) suggested the preclusion of mesophytic woody vegetation (shade-tolerant) in early successional habitats by prairie indigens. Regarding a prairie inclusion in northeastern Nebraska, Hanson (1922) noted that the highest growing-season (May-September) evaporation occurred in the prairie (air near the surface was two to four times drier than that in the surrounding wooded area) and maintained that this, not fire, precluded woody invasion. Percent cover of rock was not given. Hanson (1922) further described grassland succession through three stages as Quercus-Quercus (shade- intolerants) to Quercus~Carya to T ilia-Osnya (shade-tolerants). Hardin (1988) described the seedling stratum in a prairie inclusion in southeastern Ohio as originally consisting of Catya sp. and Quercus alba (shade-intolerants) which succeeded after 22 years by Ostrya virginiana, Crataegus sp., Prunus seron'na and Sassafras albidum (both shade-tolerant and shade-intolerant). Therefore, managed sites may also have incorporated shade-tolerant species due to succession in all of the woody layers since the previous burn (averaging 3.8 years at the 1994 sampling). Also, vigorous resprouting of mesophytic shade-tolerant species such as Ostrya virginiana, Ulmus rubra, Amelanchier arborea and Acer saccharum as well as extensive thickets of Rhus aromanca (shade tolerant) were observed at all of the managed sites. This may account for the decrease in the proportion of shade-intolerant species in the opening. The proportion of shade-intolerant woody species in the opening (of managed and unmanaged sites) increased from the understory to the overstory. In forests, shade-tolerant understory layers generally subtend predominately shade-intolerant overstory (Oliver and Larson 1990). Relative importance of sapling Quercus spp. in the opening (26.5%) was comparable to that of seedlings (28.4%) (Tables 43 and 38, respectively). However, Quercus spp. were more 154 important in the tree stratum (54.6%) of the opening, especially at managed sites (69.7%) (versus 39.4% at unmanaged sites). Selection of fire- and drought-resistant and/or shade-intolerant species would be expected in the opening at both managed and unmanaged sites. Density. The length of time since management (averaging 3.8 years for the 1994 sampling) may have obscured differences in the opening for woody seedling density (per 25 m2) between managed and unmanaged sites, allowing time for seedling establishment at the managed sites (Table 41). Although there was no difference in seedling density between managed and unmanaged site openings, managed sites had more seedlings in the transition zone subhabitats. Seedling density is known to increase in response to management (Anderson and Schwegman 1991). Anderson and Schwegman (1991) found a doubling in seedling density (seedlings were defined by height and therefore may have included resprouts) one year after fire in a mesic southern Illinois barrens site. Seedling density subsequently declined, reaching pre-burn levels about ten years postfire. Conversely, shrub and sapling density decreased one year afier the fire then rapidly increased. Hardin (1988) in a prairie inclusion in southeastern Ohio reported that seedling density was originally higher in the transition zones than in the opening, though this situation was reversed after 22 years of subsequent succession. The forest openings had a sparse shrub and sapling layer, giving them the characteristic open, park-like aspect of midwestem savanna (Table 45). Tree density values and to a lesser extent, shrub and sapling density values, particularly in the forest interior subhabitats, did not reflect what was visually a marked contrast beween the opening and forest interior subhabitats. Species Richness. Mean tree and tree seedling species richness (although sometimes within one standard error) were lower in the opening than in any other subhabitat for managed and run sites (Tables 40 and 49). Managed and unmanaged sites did not differ among subl...r' species richness in any of the woody strata (with the exception of the nort' ..n seedlings). Mean forest-opening density values for all three canOp} stri- 155 within one standard error of other subhabitats) were also lowest in the opening (Tables 41, 45 and 50). Therefore, peaks in the opening percent rock cover (17.2 i 3.6, excepting the transition zone- south), shallow soil depth and exposure (insolation and moisture stress) may account for the trough in woody species richness in the opening. Alternatively, Garman (1925), Transeau (1935) and Walter (1973) attributed the lack of woody seedlings in the prairie to competitive exclusion by grasses. Approximately four more woody seedling species (per 25 m2) were found in the forest interior subhabitats (9.7) than in the opening (5.9) of the forest-openings (1' able 40). The forest interior-north would provide a favorable environment for mesophytic seedlings with less moisture stress and exposure than the opening and southern subhabitats. Species richness of woody seedlings was also more numerous in the northern subhabitats of the managed sites than of the unmanaged sites. This may also be attributed to the more mesic nature of the barrens than the glades which comprised the unmanaged site controls. In combination with the more moderate environment in the northern subhabitats, management may have had a positive influence on seedling species richness by clearing away litter and providing a more favorable substrate for resprouting and/or germination. Shearin et a1. (1972) found that prescribed bunting increased Lirr‘odendron tulipifera seedling number and height growth significantly three growing-seasons after a burn in pine and hardwood plantation stands in South Carolina. They attributed the change to improved (earlier) germination of seedlings. In the present study, a greater number of tree seedling species, including mesophytic species (e. g., Lirr'odena’ron rulipifera, Fagus grandifolia and Liquidambar styracrflua), were counted in 1993 than in 1988. Canopy Measurements. Mean tree crown diameter, height and dbh for all forest-openings did not differ among subhabitats (Tables 54, 53 and 52, respectively). However, mean tree height and crown diameter of managed sites exceeded that of unmanaged sites in the Opening and forest interior subhabitats. This most likely reflects the more mesic nature of the barrens and limestone 156 glades which comprised the managed sites versus the xeric unmanaged sandstone and shale glades. The xeric growing conditions and shallow soil of the unmanaged glade sites are also known to produce a stunted, gnarled growth of trees (Reich and Hinckley 1980). However, height differences were difficult to discern given the lack of habitat qualifying as opening in the subsample transect at the unmanaged sites. Although it is generally assumed that open-grown trees (especially in upper Midwest savanna) are larger (taller, greater dbh) than forest-grown trees, Anderson and Anderson (1975) found that dbh values for savanna trees in Illinois did not differ from those based upon governmental land office records of 52.8 .1: 4.7 cm versus 44.7 :1: 3.3 cm in surrounding forest. Soil Measurements: 1993 and 1994 The classification key by Heikens (1991) separates glades from barrens based upon soil depth. According to her key, glades have a soil depth _<_10 cm whereas barrens have a depth of 10 to nearly 40 cm. However, in 1988 three barrens averaged 9.4 cm depth (the six glades averaged 6.3 cm) and two of three barrens (Brown-MGD and Gibbons-MOD) had soil depth values less than 10 cm (Heikens 1991). In 1993 glades averaged 7.0 i 2.9 cm soil depth (Table 60). Still, the Cave-MGD and Cedar-UMG glades had values of 13.1 i 2.9 cm and 18.4 :t 2.9 cm, respectively. In 1993, barrens averaged 10.9 i 0.9 cm, with Gibbons (9.1 cm) below the lO-cm criterion of Heikens. Findings by Jeffries (1987) concur with the key by Heikens (1991). The three sandstone glades studied by Jeffries (1987) had a mean soil depth of 5.2 cm. Although boundaries for soil depth can be used to distinguish glades and barrens, these results suggest that individual sites may not be representative of their respective forest-opening type. SUMMARY AND CONCLUSIONS The effect of management on Poaceae and Asteraceae RI followed trends described by others in the literature. Averaging 4.8 years since fire management at the 1993 sampling, Poaceae RI declined at both managed (-11.8%) and unmanaged sites (-6.3%) while Asteraceae RI declined at managed sites (-2.0%) but increased at unmanaged sites (2.6%). Gibson and Hulbert (1987) and Piper (1995) reported that as dominance of graminoid species declines, the number of forbs (primarily composed of Asteraceae species) and diversity increase for approximately six to seven years, after which diversity declines. The Fabaceae also showed an increase in RI with management (30.5% for three of four managed sites), even after an extended postfire period whereas RI decreased for four of five unmanaged sites (-46.6%). This complements the findings of Martin et a1. (1975), Anderson and VanValkenburg (1977) and Towne and Knapp (1996) who found increases in legumes (e.g., density, biomass) after fire management. The findings for the 1994 transect data involved the comparison of relatively mesic barrens and limestone glades (the managed sites) with more xeric, exposed sandstone and shale glades (the unmanaged sites) (Gyp, an unmanaged, comparatively mesic barrens, had a closed canopy and therefore zonation into subhabitats was not possible). The ratio of glades to barrens was greater for unmanaged (4:0) than for managed sites (2:2). These site-specific differences confounded the efl‘ects of management. The managed sites were more species-rich (herbs) (excepting the transition zone-south) and had greater herb cover than the unmanaged sites in all subhabitats. In the opening managed sites showed a peak in herb cover while unmanaged sites troughed in herb species richness. These differences are believed to reflect differences imposed by exposure, moisture regime and exposed rock cover (e. g., barrens averaged 0.8% while glades averaged 21.3%). unmanaged sites. Sites managed (prescribed burn) more than once contained characteristic prairie species which were in the first RI position and were unique to the opening (excepting the transition 157 158 zone-south at Wildcat-MGD). Management was also believed to maintain zonable subhabitats and However, it is also believed that management was responsible for maintaining, if not enhancing, herb species richness and cover in the managed forest-openings, especially in the opening. Asteraceae RI and herb species diversity were also higher in the opening of managed than of distinct distributions of herbs at the more mesic barrens and limestone glade sites (versus Gyp- UMG which had succeeded to a closed-canopy). Herb species composition showed some striking similarities among the sampled forest- openings, regardless of previous management history (if any), substrate or forest-opening type. There was little variability among sites in the proportion of grasses, forbs, legumes and exotics, and in categories of life history (annual, biennial, perennial), life form (Chamaephyte, geophyte, hernicryptophyte, therophyte) and percent association with a particular habitat type (e.g., bluff, open woods). However, when subdivided into subhabitats and compared the sampled forest- opening values did differ from values for similar habitats in other studies. The forest-openings had a lower proportion of perennials (by 220%) and a higher proportion of annuals (by ==20%) than the tallgrass prairie (Risser et al. 1981). This is attributable to their xeric characteristics, i.e., shallow soil, southern to western exposure, elevated topographic position and relatively high percentage of exposed rock. These features were also largely responsible for their longevity as openings (especially the unmanaged glades), causing them to be unattractive for cultivation and slower to succeed to closed forest. The findings of this study supported others (White and Madany 1978, Nuzzo 1986, Packard 1990, Hutchison 1994) who contend that the forest-opening herb flora is transitional between a prairie and a forest, with values intermediate between east-central Illinois mixed hardwood forest and southern Illinois hill prairie habitats in the percentage of typical woodland and prairie species inhabitants. Still, the forest-openings had a composite 58.5% woodland/open woods type herb flora and only an 8.2% association with a prairie habitat. However, the opening 159 subhabitat of managed sites more closely approximated a southern Illinois hill prairie in the ratio of geophytes to hemicryptophytes than an east-central mixed hardwood forest. Also, the high percentage of geophytes (often rhizomatous species) found in the opening of managed sites (34.1% higher in the opening of managed sites than unmanaged sites) is underscored by many authors as an important feature of grasslands (Anderson 1982, Anderson 1990, Seastedt and Ramundo 1990). Forest-opening data were influenced by drought and succession and results attributable to both of these factors supported others in the literature. Both managed and unmanaged sites increased in herbaceous species richness since the 1988 drought-year sampling (by an average of 4.5 species per plot; the average of up to 30, four l-m2 plots in 1993 was compared to that of 50 m2 plots in 1988). The percent increase for unmanaged sites, 58.4%, exceeded that for managed sites, 23.5%. Although no pre-drought data were available, species richness (especially the number of annual species; Tilman and El Haddi 1992) has been reported to be lower during the severe drought and higher before and afterward (Piper 1995). Likewise, the proportion of annuals was 5.6% to 18.1% higher in 1993 for two of the sampled forest-openings. Succession was responsible for the increase in herb cover which occurred at several of the unmanaged sites as evinced by the concommitant increase in relative importance of broadleaf woodland herbs (e. g., Parthenocissus quinquefolr‘a, T oxicodendron radicans) and successional, open woods species (e.g., Helianthus divaricatus, Solidago sp.) and the presence of solitary prairie species under dense,even-aged forest (sapling to small tree in size) bordering the glade rock outcrops. Barbour et al. (1980) state that in general, vegetative cover increases during succession. Woody succession appeared to be taking place at both managed and unmanaged sites. Woody species richness increased at both unmanaged sites (40.4%) and managed sites (75.0%) (averaging 4.8 years since fire management at the 1993 sampling). Similar results are reported by Hardin (1988) and Anderson and Schwegman (1991). Also, in several instances, permanent plots installed in 1988 occurred in area defined as transition in 1994. Although the number of exotic species also 160 increased (11.7) for all sites since 1988, this appeared to correlate with proximity and/or frequency of human visitation, rather than with forest-opening type or management history (if any). Transect data yielded unexpected descriptive information regarding herb species distribution. For example, the openings were composed of unusual matrix species. Although Midwestern savanna are typically dominated by prairie grasses such as Schizachyn’um scopan'um, Andropogon gerardii, Sorghastrum nutans and Panicum virgatum, 21.4% of the opening subhabitat R1 was comprised of Danthom'a spicata, xeric Carex sp. (probably C. artitecra and C. umbellata), Dichanthelium laxiflorum (woodland species) and 8.0% Schizachyrium scoparium. The Poaceae appeared to track topographic moisture gradients, increasing in RI in dry subhabitats while the Cyperaceae favored more mesic areas. The opening subhabitat of managed and unmanaged sites was similar in that woody species richness and density for all canopy strata were lower than in the transition or forest interior subhabitats (although sometimes within one standard error). As noted by Heikens (1991) and Jeffries (1987) the openings of different forest-opening types are characterized by particular woody species. For example, Juniperus virginiana was most important in the opening at the sandstone glades. At both managed and unmanaged sites, however, shrub and sapling and tree density values did not reflect what was visually a marked decrease between the opening and forest interior. Differences in forest-opening woody species composition, like herb composition, appeared to be confounded by differences in managed and unmanaged site substrate and moisture regime. For example, in the seedling, shrub and sapling and tree strata of the opening subhabitat, shade- intolerant species were more important at the unmanaged than at the managed sites. Additionally, vigorous resprouting of mesophytic, shade-tolerant species (e.g., Ostrya virginiana, Amelanchier arborea) and extensive thickets of Rhus aromatica (shade-tolerant) often occurred at the managed sites. Tree height and crown diameter in the opening and forest interior of managed sites exceeded that of unmanaged sites. The xeric growing conditions and shallow soil of glade sites are known to 161 produce a stunted, gnarled growth of trees (Reich and Hinckley 1980). The forest-openings, like forests, increased in R1 of shade-intolerant species from the understory to the overstory of the opening regardless of management history (if any) (Oliver and Larson 1990). This was primarily due to an increase in Quercus sp. Selective forces for Quercus include periodic and seasonal drought, fire and exposure in the opening subhabitat. The sampled forest-openings had an aspect of south or west. The perimeter of the opening subhabitats were undulating, and the east-west axis was longer than the north-south axis for five of eight sites. The openings ranged from 450 to 7825 m2. Although the openings of managed sites were larger in area (175.9%) and more species-rich (34.0% for opening plots; plot size was larger in 1988 sampling) than unmanaged sites, there was no correlation between site opening area and total herb number (0.0005oo ooccwocoocooocoocc UINON— LAO DJ -— \ON 00 SS—oowooowoooou‘N—N—owhowo—oo-oooo~n~monoomooooqo peooocooocooo more A 169 T riodanis perfoliata Verbena urticifolia Vernom'a gigantea Viola raphanesquii Vulpia octoflora Woodsia obtusa wooooo pooooo cooooo BROWN SHALE BARRENS Number of circular 50 m2 plots for woody taxa: 30 Number of nested 1 m2 plots for herbaceous taxa: 120 *indicates plant was sampled with the herbaceous taxa and, although bearing a woody stem, was located within the herb layer, i.e., within approximately 1 m of ground level. *.*indicates value less than one tenth of a percent. "*indicates value less than one-one hundredth of a percent. No. of plots Woody taxa of occurrence Dominang Relative Importan_ce_ Acer rubrum 4 0.01 1.2 Acer saccharum 2 0.03 0.7 Amelanchter arborea 0 0 0 Aralia spinosa 0 0 0 Campsis radicans 0 0 0 Carya cordiformis 0 0 0 Carya glabra l 0.02 0.4 Carya ovalis 2 0.02 0.7 Carya ovata 17 0.27 6.2 Carjya texana 0 0 0 Carya tomentosa 0 0 0 Celtis tenurfolia l 0.02 0.4 Cercis canadensis l *** 0.3 Cornusflorida 0 0 0 Coo/[us americana 2 0.03 0.7 Crataegus engelmannii 0 0 0 Crataegus monogyna 1 0.02 0.4 Crataegus sp. 1 0.02 0.4 Diospyros virginiana 3 0.05 1.1 Euonymus atropurpurea 1 0.02 0.4 Fagus grandrfolia 0 0 0 Fraxinus americana 9 0.78 6.7 Gleditsia triacanthos 0 0 0 Hypericum stragulum 0 0 0 Juglans nigra 1 0.13 0.9 Juniperus virginiana 0 O 0 Ligustrum vulgare 1 0.02 14.8 Liriodendron tulipifera 3 0.01 0.9 Lonicera japonica‘ 9 0.2 l 0. 7 Lonicera sp. (shrub) l 0.02 0.4 Moms rubra Ostrya virginiana Parthenocissus quinquefolia“ Prunus serotina Quercus alba Quercus coccinea Quercus imbricaria Quercus prinoides acuminata Quercus rubra Quercus stellata Quercus velutina Rhus copallina Rosa carolina Rubus allegheniensis Rubusflagellaris Sassafras albidum Smilax bona-nox Smilax glauca Symphoricarpos orbiculatus Toxicodendron radicans* Ulmus alata Ulmus americana Ulmus rubra Vaccinium arboreum Viburnum prunifolium Vitis aestivalis“ Vitis vulpina* Herbaceous ta_x_a Acalypha gracilens Acalypha virginica Achillea millefolium Agalinis tenuifolia Agrimonia rostellata Agrostis perennans Ambrosia artemiisifolia Amphicarpa bracteata Anemone virginiana Antennaria plantaginifolia Apocynum cannabinum Arabis laevigata Arisaema dracontium Aristida sp. Aristolochia serpentaria Asclepias tuberosa Asclepias variegata Asparagus ofl‘icinalis Asplenium platyneuron A LII NNNaNo—‘UIOOOHOOOOANNr—Qo—‘OWNO No. of plots of occurrence 0 N ~OO—Or—HOOMOONOOOO O Domin_ance 0.58 O oopoopoooo 0;.) --o UJUJ o o 0.13 0.13 0.7 0.2 0.3 2.6 0.4 0.7 36.5 1.5 ooopoo DJ 0.2 8.9 0.7 10.5 0.7 0.1 0.1 Relative Importance 4.7 O A 4> iv... coopoopoor—oof—oooo N O\ Aster patens Aster sp. Bidens bipinnata Bromus pubescens Cacalia atriplicifolia Cardamine sp. Carex artitecta Carex bushii Carex cephalophora Carex digitalis Carex glaucodea Carex hirsutella Carex muhlenbergii C arex retroflexa Carex rosea Carex sp. Carex umbellata Cassia fasciculata Cassia nictitans Chasmanthium latifolium Cirsium discolor Claytonia virginica Conyza canadensis Coronilla varia C roton monanthogynus Cunila origanoides C ynoglossum virginianum C yperus ovularis C ystopteris protrusa Danthonia spicata Desmodium canescens Desmodium nudiflorum Desmodium paniculatum Desmodium rotundifolium Desmodium sp. Dichanthelium acuminatum Dichanthelium boscii Dichanthelium depauperatum Dichanthelium dichotomum Dichanthelium Iaxiflorum Dichanthelium linearifolium Dichanthelium malacophyllum Dichanthelium polyanthes Dichanthelium sphaerocarpon Dichanthelium villosissimum Diodia teres Dioscorea quaternata Dodecatheon meadia Elymus sp. OO’sN DOA v—OOWOOO"OU—ONOQQOOOOMOAOr-‘NOOOOOVOD—OANONh—OOONOOOO 11.3 Elymus villosus Elymus virginicus Eragrostis spectabilis Erechtites hieracifolium Erigeron annuus Erigeron strigosus Eupatorium rugosum Eupatorium serotinum Euphorbia corollata Galactia regularis Galium aparine Galium circaezans Galium pilosum Galium triflorum Geranium carolinianum Geranium maculatum Geum canadense Gnaphalium purpureum Hackelia virginiana Hedeoma pulegioides Hedyotis purpurea Hedyotis pusilla Helianthus divaricatus Heuchera americana Hieracium gronovii Hypericum drummondi i Hypericum punctatum Juncus secundus Juncus sp. Juncus tenuis Krigia dandelion Lactuca serriola Lechea tenuifolia Lespedeza procumbens Lespedeza repens Lespedeza violacea Lespedeza virginica Liparis lilifolia Lobelia inflata Luzula multiflora Manfreda virginica Monarda bradburiana Muhlenbergia capillaris Muhlenbergia sobolifera Myosotis verna Nothoscordum bivalve Oxalis stricta Oxalis violacea Parietaria pensylvanica A M b) H \00\ owgowooooo—oowoo—-—oowu—oooooc'noooowoo—n—o—ooooo—Nooooo A *N \II' it» _ cpooooopoooopoopyc>fioopoay~ooooo 00 o'er—o'oooopaooooof—Qoo u—nm —- 0* N #0 I—‘L’I Paronychia pensylvanica 0 O 0 Passiflora lutea l *** “ * Penstemon pallidus 5 0.17 * * Penstemon sp. 16 0.49 1 3 Physalis pruinosa 0 0 0 Physalis virginiana 4 0.13 0 3 Phytolacca americana 0 0 0 Plantago lanceolata 0 0 0 Plantago virginica 16 0.09 0.8 Poinsettia dentata 0 0 0 Polygala verticillata 25 0.16 1 3 Polygonum tenue l *** * * Polystichum acrostichoides 0 0 0 Potentilla simplex 0 0 0 Prenanthes altissima 0 0 0 Psoralea psoralioides 0 0 0 Pycnanthemum tenuifolium 28 0.40 1 7 Ranunculus hispidus 4 0.01 0 2 Rudbeckia hirta 0 0 0 Ruellia caroliniensis 0 0 0 Ruellia humilis 8 0.46 0.9 Sabatia angularis 0 0 0 Sanicula canadensis 0 0 0 Schizachyrium scoparium 103 9.04 16.2 Senecio glabellus 0 0 0 Solidago caesia 0 0 0 Solidago juncea O 0 0 Solidago missouriensis 0 0 0 Solidago nemoralis l 0.13 0.2 Solidago ulmifolia 0 0 0 Sphenopholis obtusata 3 0.01 0 l Stylosanthes biflora 55 2.08 5 0 T radescantia virginiana 0 0 0 Triodanis perfoliata 84 0.38 4 0 Vallerianella radiata 4 0.01 0.2 Veronicastrum virginicum 0 0 0 Viola raphanesquii 3 0.01 0.1 Viola sororia 0 0 0 Viola triloba O 0 0 Vulpia octoflora 3 0.01 0 1 Woodsia obtusa 5 0.1 1 0 4 CAVE CREEK LIMESTONE GLADE Number of circular 50 m2 plots for woody taxa: 27 Number of nested 1 m2 plots for herbaceous taxa: 108 I"indicates plant was sampled with the herbaceous taxa and, although bearing a woody stem, was located within the herb layer, i.e., within approximately 1 m of ground level. 174 “indicates plant was sampled with the woody taxa. *.*indicates value less than one tenth of a percent. *"indicates value less than one-one hundredth of a percent. No. of plots Woody taxa of occurrence Dominant; Relative Importance Acer negundo 5 0.06 1.6 Acer rubrum 6 0.02 1.7 Acer saccharum 7 2.43 9.9 Aesculus glabra 0 0 0 Amelachier arborea 0 0 0 Aralia spinosa 0 0 0 Betula nigra 0 0 0 Bignonia capreolata“ 27 1.13 2.4 Campsis radicans“ 17 0.85 1.7 Carya cordiformis 0 0 0 Carya ovalis 0 0 0 Carya ovata 3 0.06 1.0 Carya texana 0 0 0 Ceanothus americanus 3 0.06 1.0 Celastrus scandens* 4 0.23 0.4 Celtis occidentalis 2 0.04 0.7 Celtis tenuifolia 12 0.21 3.9 Cercis canadensis 17 0.96 7.9 Camus drummondii l 0.02 0.3 Cornusflorida 5 1.50 6.3 Corylus americana 0 O O Crataegus engelmannii 1 0.02 0.3 Crataegus mollis 4 0.07 1.3 Crataegus sp. 1 0.02 0.3 C rataegus viridis 0 0 0 Diospyros virginiana 10 0.31 3.8 Euonymus atropurpurea 4 0.06 1.3 Fraxinus americana 15 0.93 7.2 Gleditsia triacanthos 1 0.02 0.3 Hypericum prolificum 0 0 0 Her decidua 4 0.33 2.2 Juglans nigra 4 0.07 1.9 Juniperus virginiana 8 0.12 2.6 Lindera benzoin 0 0 0 Liquidambar styraciflua O 0 0 Liriodendron tulipifera 0 0 0 Lonicera japonica 0 0 0 Malus ioensis 1 0.02 0.3 Menispermum canadense 0 0 0 Moms rubra 0 0 0 Ostrya virginiana 1 0.02 0.3 Parthenocissus quinquefolia“ 9 0.24 0.7 Prunus americana 3 0.06 1.0 Prunus serotina Quercus marilandi ca Quercus prinoides acuminata Quercus rubra Quercus shumardii Quercus stellata Quercus velutina Rhus aromatica Rhus copallina Rhus glabra Rosa carolina“ Rosa multrflora Rubus allegheniensis* Rubus enslenii (IL-E) Rubusflagellaris Rubus occidentalis Sassafras albidum Smilax bona-n0x* Smilax hispida Smilax rotundifolia Symphoricarpos orbiculatus T oxicodendron radicans“ Ulmus alata Ulmus rubra Viburnum prunifolium Viburnum rufidulum Vitis aestivalis“ Vitis vulpina Herbaceous taxa Abutilon theophrastii Acalypha gracilens Agrimonia rostellata Agrostis alba Allium vineale Ambrosia artemisiifolia Anagallis arvensis Andropogon gerardi i Anemone virginiana Apocynum cannabinum Arabis canadensis Aristida sp. Aristolochia serpentaria Arundinaria gigantea Asarum canadense Asclepias syriaca Asclepias tuberosa Asclepias verticillata fl .5 00 N OWOfl:NOOOOWOOOOQOHN—WOOI—‘t—NOH No. of plots of occurrence 0 23 O F-‘OOOOUJwt-‘OOOOQF—O Dominance O 0.1 l 0.3 17.6 0.3 14.7 cpl-‘3‘93‘o’o .. N—wo cowoooor— ox Rglative Importance 0 1 . l 0 0 t . * 0.4 Asclepias viridiflora Aster oblongifolius Aster patens Aster pilosus Aster sp. Aster turbinellus Aureolariaflava Boehmeria cylindrica Botiychium virginianum Bouteloua curtipendula Brickellia eupatorioides Bromus commutatus Bromus pubescens Bromus racemosus Cacalia atriplicifolia Camassia scilliodes Campanula americana Capsella bursa-pastoris Cardamine sp. C arex artitecta Carex blanda Carex cephalophora Carex meadii Carex muhlenbergii Carex retroflexa Carex sp. Carex umbellata Cassia fasciculata Cassia marilandica“ Cassia nictitans Cerastium arvense Chamaesyce maculata Cheilanthesfeei Cirsium altissimum Cirsium discolor Clematis pitcheri Conyza canadensis Coreopsis tripteris Cosmos bipinnata Croton monanthogynus Crotonopsis elliptica C unila origanoides Cuscuta sp. C ynanchum laeve Danthonia spicata Dentaria laciniata Desmodi um canescens Desmodium paniculata Desmodium rotundifoium 0w \1 _ N O\ \l NOOOF—WOOOSOWOOON—v—‘OI—wv—O-‘OOOOOOOOGOOOHOQHOOOONOr—ON cocooooo _o .— ox O 0.49 0.06 *** 0.14 0.04 0.07 0.11 0.19 0.05 0.04 0.01 #993 moo-— o_oo N O\u—- ON poooooooocooopopg—ooo A cooocpooog—ocooopppoaeg—o —- --N o w Nv—O *ON \I Desmodium sessilifolium Desmodium sp. Dianthus armeria Dichanthelium boscii Dichanthelium laxiflorum Dichanthelium malacophyllum Dichanthelium sp. Dichanthelium villosissimum Dioscorea quaternata Dioscorea villosa Dodecatheon meadia Echinacea pallida Elymus hystrix Elymus virginicus Eragrostis capillaris Erechtites hieracifolia Erigeron annuus Eupatorium altissimum Eupatorium rugosum Euphorbia corollata Festuca arundinacea Fragaria virginiana Frasera caroliniensis Galactia regularis Galium circaezans Galium concinnum Galium pilosum Gaura longiflora Geranium carolinianum Geranium maculatum Hedeoma pulegioides Hedyotis purpurea Helianthus divaricatus Helianthus microcephalus Heliopsis helianthoides Hordeum pusillum Hybanthus concolor H ydrastis canadensis Hypericum denticulatum Ipomoea pandurata Iva annua Kummerowia stipulacea Lactuca canadensis Lactuca floridana Lactuca serriola Lepidium virginicum Lespedeza cuneata Lespedeza procumbens Lespedeza repens H b) 00 MW ~—-—ooo—-o—oaoowoo—$N~0No\io—mom—womo—-—-—-—-woooo—-—ooo\owo .— 177 0.08 0.13 n: 0.15 0.41 1.31 **# 0.05 0.04 0.04 1.38 0.40 0.04 0.15 1.27 0.10 0.06 0.01 *** 0.01 1.36 0.94 0.08 0.06 0.22 *** 0.04 0.04 0.04 Lespedeza sp. Leucanthemum vulgare Liatris scabra Lithospermum canescens Lysimachia lanceolata Manfreda virginica Matricaria matricarioides Medicago lupulina Melica mutica Melilotus alba Monarda bradburiana Monarda fistulosa Muhlenbergia sobolifera Nothoscordum bivalve Onosmodium hispidissimum Oxalis dillenii Oxalis stricta Panicum anceps Passiflora lutea Pellaea atropurpurea Penstemon sp. Phleum pratense Phlox pilosa Phryma leptostachya Physalis virginiana Physostegia virginiana Plantago lanceolata Plantago rugellii Poa compressa Poinsettia dentata Polygonum cristatum Ranunculus sp. Ratibida pinnata Rudbeckia hirta Ruellia caroliniensis Ruellia humilis Ruellia strepens fl N H O O\v-- Oz—NOOOOO-fiNOWOMOHOMOv—ONv—‘NOOOONr-‘fi—OOUJ O 40 17 Salvia azurea grandiflora (IL-T)7 Salvia lyrata Sanguinaria canadensis Sanicula canadensis Schizachyri um scoparium Scutellaria leonardii Setariafaberi Setaria glauca Sida spinosa Silene stellata Silphium integrifolium Silphium terebinthinaceum 2 0 13 57 2 WOO—ONO 179 0.05 1.16 0.11 3° C) Sisyrinchium albidum Solidago sp. Solidago ulmifolia Sorghastrum nutans Spigelia marilandica T aenidia integerrima T araxacum oflicinale T eucrium canadense T orilis japonica T radescantia subaspera T ridens flavus T rifolium campestre T rifolium pratense T riosteum angustifolium Uvularia grandifolia Verbascum thapsus Verbena urticifolia Verbesina helianthoides Verbesina virginica Vernonia gigantea Viola raphanesquii Viola sororia Viola sp. Viola triloba Zizia aurea 00 LII .0 \0 U1 NON NW“ C O 11> O\ LAG—‘0OOAOOOOOOOONOOOOONWWO .0 N N 0\ DJ CEDAR BLUFF SANDSTONE GLADE Number of circular 50 m2 plots for woody taxa: 30 Number of nested 1 m2 plots for herbaceous taxa: 120 ‘indicates plant was sampled with the herbaceous taxa and, although bearing a woody stem, was located within the herb layer, i.e., within approximately 1 m of ground level. *.*indicates value less than one tenth of a percent. "*indicates value less than one-one hundredth of a percent. No. of plots Woody taxa of occurrence Dominance Relative Importance Acer rubrum 0 0 0 Acer saccharum 4 0.04 O. 8 Amelanchier arborea 12 0.30 2.5 Carya cordiformis 0 0 O Carya glabra 20 1.8 5.7 Carya ovalis 12 0.95 3.3 Carya ovata 0 0 0 Carya texana 14 0.70 3.4 Carya tomentosa l 0.02 0.2 C elastrus scandens 0 0 0 Celtis laevigata 1 0.02 0.2 Celtis occidentalis Celtis tenuifolia C ercis canadensis Cornusflorida C rataegus englemannii Diospyros virginiana Fagus grandifolia Fraxinus americana Gleditsia triacanthos Hypericum stragulum“ Juglans nigra Juniperus virginiana Ligustrum vulgare Lonicera japonica“ Lonicera sp. (shrub) Malus ioensis Morus rubra Ostrya virginiana Parthenocissus quinquefolia“ Prunus americana Prunus serotina Quercus alba Quercus Xbushii Quercus coccinea Quercus imbricaria Quercus marilandica Quercus prinoides acuminata Quercus rubra Quercus stellata Quercus velutina Rhus aromatica Rhus copallina Rhus glabra Rosa multiflora Rubus allegheniensis Rubus enslenii (IL-E) Rubusflagellaris“ Smilax bona-nox“ Smilax glauca Symphoricarpos orbiculatus T oxicodendron radicans“ Ulmus alata Ulmus rubra Vaccinium arboreum Vaccinium pallidum Vitis aestivalis“ Vitis vulpina“ O A N \l u—nt—nv—n \IO -_O\NNW\OOAt—OO-‘>~ \OLII 180 0.11 0.05 0.02 0.03 0.36 2.49 *** 15.15 0.70 0.02 2.58 3.30 0.02 0.08 0.02 0.73 0.02 0.23 12.48 0.52 0.52 0.02 0.02 0.07 0.79 0.20 0.15 2.67 2.98 0.02 0.45 0.02 0.03 #¥* 1.6 0.8 0.4 0.4 2.2 5.4 0.1 23.1 1.8 0.2 4.8 9.5 0.4 1.0 0.2 1.9 0.2 1.5 19.5 2.4 3.7 0.2 0.2 0.8 2.5 0.6 1.8 0.07 8.8 0.4 1.6 0.2 0.1 *t Herbaceous taxa Acalypha gracilens Achillea millefolium Agrimonia rostellata Agrostis elliotiana A grostis perennans Allium canadense Allium vineale Ambrosia artemisiifolia Apocynum cannabinum Arabis laevigata Arisaema triphyllum Aristida sp. Aristolochia serpentaria Asplenium pinnatifidum Asplenium platyneuron Aster pilosus Aster sp. Bidens bipinnata Bromus commutatus Bromus pubescens Bromus racemosus Cardamine hirsuta Carex artitecta Carex blanda Carex cephalophora Carex glaucodea Carex hirsutella Carex retroflexa Carex sp. Carex umbellata Cerastium arvense Chasmanthium latifolium Cheilanthes lanosa Corydalisflavula Crotonopsis elliptica C unila origanoides C ystopteris protrusa Danthonia spicata Desmodium paniculatum Desmodium rotundifolium Dianthus armeria Dichanthelium acuminatum Dichanthelium boscii Dichanthelium dichotomum Dichanthelium laxiflorum Dichanthelium linearifolium Dichanthelium malacophyllum 181 No. of plots of occurrence Donfim 71 0.29 0 O O 0.01 0 0 O 0.03 0 0.01 *t* 0.12 6 0.52 0.07 0.04 *** 0.02 0.04 0.05 0.19 0.01 0.89 \O LII *** 0.99 0.49 #0 0.17 0.37 LII LII 2.61 *#* 0.16 1.06 LIIO \l 0 2 0 0 0 6 0 4 l 0 O 0 8 0 2 3 3 l 5 0 0 0 0 9 6 l 3 3 0 l 3 l 0 4 l 0 9 0 l 0 l 6 0 2 0.88 0 0 Relative Importance 3 .8 0 0 0.1 0 0 0 0.3 O 0.2 0.1 0 0 0 0.6 0 2.3 0.3 0.2 0.1 0.3 0 O 0 0.5 0.4 1.3 0.2 3.5 0 0.1 3.9 1.7 0 0.5 1.5 0 9.9 0 0.1 0 0.8 5.2 0 3.1 0 O Dichanthelium villosissimum Digitaria sp. Diodia teres Dodecatheon meadia Dryopteris marginalis Elymus virginicus Erigeron annuus Erigeron strigosus Eupatorium rugosum Eupatorium serotinum Euphorbia corollata Festuca arundinacea Galactia regularis Galium pilosum Gnaphalium purpureum Hedyotis nutalliana Hedyotis pusilla Helianthus divaricatus Heuchera americana Hieracium sp. Hordeum pusillum Hypericum gentianoides Hypericum punctatum Juncus secundus Juncus tenuis Krigia virginica Kummerowia striata Lactuca canadensis Lactuca serriola Lechea tenuifolia Leersia virginica Lepidium virginicum Lespedeza cuneata Lespedeza repens Liatris sp. Manfreda virginica Muhlenbergia sobolifera Myosotis verna Nothoscordum bivalve Oenothera linifolia Opuntia humifiisa Oxalis dillenii Oxalis stricta Oxalis violacea Parietaria pensylvanica Paronychiafastigiata Paspalum ciliatifolium Passiflora lutea Penstemon pallidus NOW—'OGOH—b CD ~— N U) N O J; ~oc—Ehw\J-—Nu—AN—omowoooooo~o~oo-ao-o~o~ 182 0.23 0.23 *** tit 1.33 0.08 0.07 0.22 *t* 0.02 0.01 0.05 0.10 0.03 0.03 *** 183 Penstemon sp. 5 0.14 0.5 Phlox pilosa 39 1.12 4.2 Plantago aristata 0 0 0 Plantago virginica 6 0.02 0.3 Poa compressa 0 0 0 Poa pratensis 7 0.09 0.5 Polygonatum biflorum 0 0 0 Polygonatum commutatum 0 0 O Polygonum cristatum 19 0.08 1.0 Polygonum tenue l *” 0.1 Pycnanthemum tenuifolium 3 0.04 0.2 Pyrrhopappus carolinianus 0 0 0 Ranunculus sp. 2 0.04 0.2 Ruellia humilis 0 0 0 Ruellia pedunculata 0 0 0 Rumex acetosella 0 0 0 Sanicula canadensis 30 0.16 1.7 Schizachyrium scoparium 13 0.23 1.1 Scutellaria leonardii 0 0 0 Sedum pulchellum 18 0.08 0.9 Smilacina racemosa 0 0 0 Solidago caesia 19 0.46 1.8 Solidago nemoralis 0 0 0 Solidago sp. 5 0.08 0.4 Solidago ulmifolia 0 0 0 Sphenopholis obtusata 4 0.04 0.3 Stylosanthes biflora 0 0 0 T ephrosia virginiana 0 0 0 T riodanis perfoliata 39 0.16 2. l Verbascum thapsus 0 0 0 Vulpia octoflora 15 0.06 0.8 Woodsia obtusa 25 0.54 2.3 GIBBONS CREEK SANDSTONE BARRENS Number of circular 50 m2 plots for woody taxa: 15 Number of nested 1 m2 plots for herbaceous taxa: 60 ‘indicates plant was sampled with the herbaceous taxa and, although bearing a woody stem, was located within the herb layer, i.e., within approximately 1 m of ground level. *.*indicates value less than one tenth of a percent. "*indicates value less than one-one hundredth of a percent. No. of plots Woody taxa of occurrence Dominance Relative Importance Acer saccharum 0 0 0 Amelanchier arborea 5 1.20 3 .4 Carya cordiformis l 0.27 0.7 Carya glabra 2 0.30 1.2 Carya ovalis Carya ovata Carya texana Ceanothus americanus C elastrus scandens Celtis occidentalis Celtis tenuifolia C ercis canadensis Camus florida Crataegus sp. Crataegus viridis Diaspyros virginiana Euonymus atrapurpurea F raxinus americana Gleditsia triacanthos Hypericum prolificum Hypericum stragulum“ Juglans nigra Juniperus virginiana Malus ioensis Morus alba Ostrya virginiana Parthenacissus quinquefolia" Prunus americana Prunus serotina Quercus alba Quercus coccinea Quercus imbricaria Quercus marilandica Quercus prinaides acuminata Quercus rubra Quercus stellata Quercus velutina Rhamnus caroliniana Rhus copallina Rosa carolina“ Rubus allegheniensis“ Rubusflagellaris* Rubus occidentalis Sassafras albidum Smilax bona-nox Smilax glauca Smilax hispida Symphoricarpas orbiculatus Toxicodendron radicans“ Ulmus alata Ulmus rubra Vaccinium arboreum Vaccinium pallidum \O O N O:— O OO\OGO\O\OOOOOAh—t—OOH—‘NOUJOOv—«DhOr-‘DJOOOOI—I—OI-‘OUIOt-‘UJNOWOOI-‘OO 12.8 24.7 4.1 10.3 3.5 Vitis aestivalis* Vitis vulpina“ Herbaceous taxa Acalypha gracilens Agrimonia rostellata A grostis alba Ambrosia artemisiifolia Amphicarpa bracteata Anemone virginiana Antennaria plantaginifolia Apocynum cannabinum Arabis canadensis Arisaema dracontium Aristida dichotoma Aristolochia serpentaria Asarum canadense Asclepias tuberasa Asclepias variegata Asclepias verticillata Asplenium platyneuron Aster anomalus Aster patens Aster pilosus Aster sp. Aster turbinellus Aster undulatus Botrychium virginianum Brachyelytrum erectum Brickellia eupatorioides Bromus pubescens Cardamine sp. Carex artitecta Carex blanda Carex bushii Carex cephalophara Carex hirsutella Carex muhlenbergii Carex retroflexa Carex sp. Carex sp. (Montanae) Carex umbellata Cassia fasciculata Cassia marilandica Cassia nictitans Cheilanthes lanosa Cirsium altissimum No. of plots of occurrence 2 O OAOONO!-I—NLIIChI—‘OOOOOOOv—OOQOI-‘ONLIIOOOr—O—‘n—ODJONNO .— L») BL» 185 0.07 0.20 Dominance 0.02 0.2 0.6 Relative Importance 0.2 0 0 0.2 5.3 0 0.5 0 0.1 0.1 0 0.1 0 0 0 0.5 2.5 0 0.1 0 0.7 0 0 0.1 0 0 0.7 0 0 0 0 0.1 0.7 0.5 0.2 1.7 1.6 0 0.2 0 UI Cirsium caralinianum (IL-T) 0 0 Clitoria mariana 1 0.07 Conyza canadensis 0 0 Corydalisflavula 0 0 Crotanopsis elliptica 0 0 C unila origanoides l 0.01 C ynanchum laeve 0 0 Cynoglassum virginianum 0 0 C yperus ovularis 0 0 Danthonia spicata 2 2.18 Dentaria laciniata 0 0 Desmodium canescens 1 0.07 Desmodium nudiflarum 0 0 Desmodium paniculatum 0 0 Desmodium rotundifolium 5 0.10 Desmodium sp. 1 0.07 Dichanthelium acuminatum 3 5.83 Dichanthelium boscii 6 0.28 Dichanthelium depauperatum 0 0 Dichanthelium dichotomum 12 1.70 Dichanthelium laxiflorum 0 0 Dichanthelium linearifolium 5 0.36 Dichanthelium malacophyllum 0 0 Dichanthelium oligosanthes 11 0.15 Dichanthelium sp. 9 0.19 Dichanthelium sphaeracarpon 0 0 Dichanthelium villosissimum 0 0 Dioscorea quaternata 2 0.33 Dioscorea villosa 1 0.01 Dodecatheon meadia 0 0 Elymus hystrix 0 0 Elymus virginicus 0 0 Erechtites hieracifolium 0 0 Erigeron annuus 0 0 Erigeron strigosus 0 0 Eupatorium rugasum 0 0 Euphorbia corollata 9 0.07 Festuca obtusa 0 0 Fragaria virginiana 1 0.01 Frasera caroliniensis 0 0 Galactia regularis 3 0.14 Galium aparine 2 0.01 Galium circaezans l 0.09 Galium concinnum 2 0.02 Galium pilosum 0 0 Galium triflorum 0 0 Geranium caralinianum 2 *** Geranium maculatum 0 0 Geum canadense 1 0.01 v— N \O opoyuooopooogoo N opoooooo o coc LII >— cocoocco — N NGN Gnaphalium purpureum Hedeoma pulegioides Hedyatis langifalia Helianthus divaricatus Heuchera americana Hieracium gronovii Hypericum gentianoides Hypericum punctatum Ipomoea pandurata Koeleria macrantha Krigia dandelion Kummerowia striata Lactuca canadensis Lactuca floridana Lactuca hirsuta (IL-E) Lactuca serriola Lechea tenuifolia Lespedeza hirta Lespedeza procumbens Lespedeza repens Lespedeza violacea Lespedeza virginica Liatris squarrosa Liparr‘s lilifolia Lobelia inflata Lobelia spicata Manfreda virginica Monarda bradburiana Monarda fistulosa Muhlenbergia sobolifera Myosotis macrosperma Nothoscordum bivalve Oxalis dillenii Oxalis stricta Oxalis violacea Panicum anceps Panicum capillare Paronychiafastigiata Parthenium integrifolium Passiflora lutea Penstemon hirsutus Phlox pilosa Phryma Ieptastachya Physalis virginiana Padophyllum peltatum Palygonatum biflorum Polygonum cristatum Polytaenia nuttallii Porteranthus stipulatus \OOOOONOF—Ot—OOOOw—‘OOONNOOOOv-‘OOOOAOOOOOOOLIIUJObJr-‘OOWOOU— \O H p—n LII oopoooopoooooo o .— O O\ 188 0.04 0.21 0.01 0.04 _o m Potentilla simplex Prenanthes altissima Pycnanthemum tenuifolium Ranunculus micranthus Rudbeckia hirta Ruellia humilis Sanicula canadensis Schizachyrium scoparium Scleria pauciflara Setaria glauca Silene stellata Sisyrinchium albidum Smilacina racemosa Solanum ptycanthum Solidago caesia Solidago juncea Solidago nemoralis Solidago petiolaris Solidago ulmifolia Sorghastrum nutans Sphenopholis obtusata Stylosanthes biflora T ephrosia virginiana T haspium trifoliatum T radescantia subaspera T radescantia virginiana T richostema dichotama T ridens flavus T riodanis perfoliata T riosteum sp. Verbesina helianthoides Verbesina virginica Vernonia gigantea Viola raphanesquii Viola sp. Viola triloba Woodsia obtusa CO oo¢¢9 LII—Ix.) 0.09 4 1.50 ooo 50.0 * OLJI O? O p—e A OD v—-L»J ~43 p—a LII p—- O oooopoopooppoopooooo —- OO 0 La) t—‘mi—OOOOOn—OOF—OOF—AOOMOOOOOOO—OOON-bOOLII—‘JELII .—n .o. )— \O creoooooeoopooppoopoooooo_*ooo u—uL/Ir—o 0.01 GYP WILLIAMS SANDSTONE BARRENS Number of circular 50 m2 plots for woody taxa: 30 Number of nested l m2 plots for herbaceous taxa: 120 ‘indicates plant was sampled with the herbaceous taxa and, although bearing a woody stem, was located within the herb layer, i.e., within approximately 1 m of ground level. "‘indicates value less than one-one hundredth of a percent. Woody taxa Acer rubrum Acer saccharum Amelanchier arborea Carya cordiformis Carya glabra Carya ovalis Carya ovata Carya texana C elastrus scandens Celtis tenuifolia Cercis canadensis Cornusflorida Crataegus sp. Crataegus mollis Diaspyras virginiana Fagus grandifolia Fraxinus americana Gleditsia triacanthos Hypericum stragulum Juglans nigra Juniperus virginiana Marus rubra Ostrya virginiana Parthenocissus quinquefolia" Prunus americana Prunus serotina Quercus alba Quercus Xbushii Quercus imbricaria Quercus marilandica Quercus prinaides acuminata Quercus rubra Quercus stellata Quercus velutina Rhamnus caroliniana Rhus copallina Rosa carolina“ Rubus allegheniensis Rubusflagellaris“ Rubus occidentalis“ Sassafras albidum Smilax bana-nox Smilax glauca“ Symphoricarpas orbiculatus T oxicodendran radicans“ Ulmus alata Ulmus rubra No. of plots of occurrence 1 0 1 1 0 10 l u—n \l LI) \J udUJ—‘NI—INN—OOLIIO\ON—OOLIIOF—N—r—O-bQO-BON u—O‘tu—n u—-\) 0\ Oh) N 10 189 Dominance t t t 0 0.67 0 0.92 0.02 0.02 11.08 0 0.07 0 0.17 0.07 0 0.22 0.02 2.37 0.02 O 1.23 0.60 0.02 0.15 4.33 0 0.04 0.10 0 0.27 4.30 0.03 0.18 15.30 0.18 0.05 0.02 0.71 0 0.48 0.17 0 0 0.03 0.22 2.69 11.65 0.02 Relative Importance 0.2 0 2.7 0 2.7 0.2 0.2 15.9 0 0.8 0 1.4 0.8 0 2.6 0.2 7.2 0.2 0 2.1 2.0 0.2 0.5 11.3 0 0.9 1.2 0 3.3 8.1 0.4 2.2 19.9 2.2 0.6 0.2 2.6 0 1.2 0.4 0 0 0.1 2.7 5.1 17.0 0.2 190 Vaccinium arboreum 0 0 Viburnum rufidulum 6 0.03 1.1 Vitis aestr'valis“ 5 0.42 No. of plots Herbaceous taxa of occurrence Dominpn__c_e Relative Importance Acalypha gracilens 43 0.18 2.2 Achillea millefolium 0 0 0 Agrimonia rostellata 0 0 0 Agrostis perennans 0 0 0 Ambrosia artemisiifolia 5 0.08 0.4 Amphicarpa bracteata 4 0.08 0.3 Antennaria plantaginifalia 64 4.38 9.9 Arabis canadensis 2 0.01 0.1 Aristolochia serpentaria 4 0.10 0.3 Asclepias verticillata 1 0.03 0.1 Asplenium platyneuron 58 1.46 4.9 Aster patens 3 0.01 0.2 Aster sp. 46 1.42 4.4 Aster turbinellus 0 0 0 Aureolariaflava 0 0 0 Botrychium virginianum 0 0 0 Bromus pubescens 26 0.46 1.9 Campanula americana 0 0 0 Cardamine sp. 0 0 0 Carex artitecta 0 0 0 Carex cephalophara 0 0 0 Carex glaucodea 1 0.03 0.1 Carex hirsutella 0 0 0 Carex muhlenbergii 1 *** 0.1 Carex sp. 42 0.41 2.6 Carex umbellata 0 0 0 Cassia fasciculata 9 0.22 0.8 Cassia nictitans 3 0.01 0.2 Cheilanthes lanosa 0 0 0 Cirsium altissimum 0 0 0 Cirsium discolor 0 0 0 Cunila origanoides 10 0.29 0 9 C ynoglossum virginianum 0 0 0 Danthonia spicata 48 1.13 3.9 Dentaria laciniata 0 0 0 Desmodium canescens 0 0 0 Desmodium glutinosum l *** 0.1 Desmodium nudiflorum 0 0 0 Desmodium paniculatum 0 0 0 Desmodium rotundifolium 0 0 0 Diarrhena americana 0 0 0 Dichanthelium acuminatum 18 0.22 1.2 Dichanthelium boscii Dichanthelium dichotomum Dichanthelium laxiflorum Dichanthelium linearifolium Dichanthelium malacophyllum Dichanthelium polyanthes Dioscorea quaternata Dodecatheon meadia Elymus hystrix Elymus villosus Elymus virginicus Erechtites hieracifolia Erigeron annuus Erigeron strigosus Eupatorium rugosum Euphorbia corollata Festuca obtusa Galactia regularis Galium circaezans Galium pilosum Geum canadense Geum virginianum Hedeama pulegioides Hedyotis longifolia Helianthus divaricatus Helianthus sp. Heuchera americana Hieracium sp. Hypericum punctatum Juncus secundus Juncus tenuis Koeleria macrantha Lactuca canadensis Lactuca hirsuta (IL-E) Lespedeza cuneata Lespedeza procumbens Lespedeza repens Lespedeza violacea Lespedeza virginica Liatris squarrosa Liparis lilifolia Lithaspermum canescens Labelia spicata Manfreda virginica Monarda bradburiana Monardafistulasa Muhlenbergia sobolifera Oxalis dillenii Parietaria pensylvanica 13 28 24 # NOONOSr-‘I-‘NOO v—NO 000 Ch Cub & b.) ©0500~OOOOWOWOOOON-50hUJOOO ooeoooo LII (a) Parthenium integrifolium 0 0 0 Passiflora lutea 0 0 0 Penstemon sp. 3 0.10 0 3 Phlox pilosa 3 0.10 0 3 Phryma leptostachya 0 0 0 Physalis virginiana 3 0.07 0 3 Polygonatum biflorum 0 0 0 Polygonum cristatum 2 0.01 0 l Polygonum virginianum 0 0 0 Polytaenia nuttallii 0 0 0 Porteranthus stipulatus 7 0.09 0.5 Potentilla simplex 0 0 0 Prenanthes altissima 0 0 0 Psoralea psoraliaides 0 0 0 Pycnanthemum tenuifolium 0 0 0 Rudbeckia hirta 14 0.17 0 9 Ruellia humilis l 0.03 0 1 Sanicula canadensis 41 0.89 3 3 Schizachyrium scoparium 53 1.15 4 2 Sedum pulchellum 0 0 0 Sisyrinchium albidum 0 0 0 Solidago caesia 0 0 0 Solidago juncea 4 0.10 0 3 Solidago nemoralis 27 0.97 2 8 Solidago petialaris 0 0 0 Solidago ulmifolia 2 0.07 0 2 Sarghastrum nutans 9 0.25 0 8 Sphenapholis obtusata 3 0.04 0 2 Stylosanthes biflora 4 0.02 0 2 T ephrosia virginiana 0 0 0 T riodanis perfoliata 4 0.01 0 2 Verbesina helianthoides 0 0 0 Viola triloba 0 0 0 Woodsia obtusa 23 0.42 1 7 POUNDS HOLLOW SANDSTONE GLADE Number of circular 50 m2 plots for woody taxa: 30 Number of nested l m2 plots for herbaceous taxa: 120 *indicates plant was sampled with the herbaceous taxa and, although bearing a woody stem, was located within the herb layer, i.e., within approximately 1 m of ground level. "*indicates value less than one-one hundredth of a percent. No. of plots Woody taxa of occurrence Dominance mtive Importance Amelanchier arborea 12 0.94 6.6 Carya glabra 4 0.17 1.8 Carya ovalis 0 0 0 Carya ovata Carya texana Celtis tenuifolia Camus florida Diaspyros virginiana Fagus grandifolia Fraxinus americana Hypericum stragulum Juglans nigra Juniperus virginiana Liriadendran tulipifera Lonicera japonica Parthenocissus quinquefolia Pinus echinata Prunus serotina Quercus alba Quercus coccinea Quercus imbricaria Quercus marilandica Quercus rubra Quercus stellata Quercus velutina Rhamnus carolinianus Rhus copallina Rosa carolina Rosa mulnflora Rubus allegheniensis Sassafras albidum Smilax bona-nax* Smilax glauca T oxicodendron radicans“ Ulmus alata Vaccinium arboreum Vitis aestivalis Herbaceous taxa Acalypha gracilens Agalis tenuifolia Agrostis elliotiana Allium canadense Allium vineale Ambrosia artemisiifolia Amphicarpa bracteata Andropogan virginicus Antennaria plantaginifalia Aristida sp. Asplenium pinnatifidum Asplenium platyneuron ONOOI—NOOWF—bO-bON NNOOOQOOOOGO—HwQNOOO No. of plots of occurrence 2 \ON ONi—OSONOI— 193 0.03 0.05 0.05 *** 0.05 8.11 *#* 0.02 0.09 0.03 0.87 0.05 2.04 0.02 2.32 2.15 Dominance 0.04 0.02 0.04 *** 0 0.17 0 0.25 0 0.08 0.04 0 0.8 1.5 1.5 0.3 1.1 31.4 1.7 13.4 12.9 Relative Importance 0.5 1.2 1.6 0.2 0 1.2 0 2.8 0 2.1 0.5 0 Aster pilosus Aureolariaflava Blephilia ciliata Bromus pubescens Bromus racemosus Carex bushii Carex glaucodea Carex hirsutella Carex sp. Carex umbellata Cassia fasciculata Cerastium arvense Chasmanthium latrfolium Cheilanthes lanosa Claytonia virginica Clitoria mariana Croton mananthogynus Crotonopsis elliptica C unila origanoides Danthonia spicata Dentaria laciniata Dichanthelium acuminatum Dichanthelium depauperatum Dichanthelium dichotomum Dichanthelium Iaxiflorum Dichanthelium linearifolium Dichanthelium malacaphyllum Dichanthelium aligosanthes Dichanthelium sphaerocarpon Dichanthelium villasissimum Digitaria sanguinalis Diodia teres Dodecatheon meadia Elymus virginicus Erechtites hieracifolia Erigeron strigosus Eupatorium serotinum Euphorbia corollata Festuca arundinacea Galactia regularis Galium pilosum Geranium maculatum Hedyotis longifolia Hedyotis purpurea Hedyotis pusilla Helianthus divaricatus Heuchera americana Hypericum denticulatum Hypericum gentianoides \) N \O ~NONOOhio'—OOWOOOOOF—OI—NOAOSOr—t—ONHWOOOQOOOOHOOOU—OOOO 194 O p—n * * ooooco *oopoooo O O A r— 0.07 0.10 N N oomoooopooocoooo so LII N LII opoococoopoooo LII 9’93 -—-\r Juncus secundus Juncus tenuis Krigia brflora Krigia dandelion Kummerawia striata Lactuca canadensis Lechea tenuifolia Lespedeza cuneata Lespedeza repens Lespedeza violacea Liatris squarrosa Linum virginianum Luzula multiflora Manfreda virginica Melica mutica Myosotis verna Nothoscordum bivalve Oenothera linifolia Opuntia humifiisa Oxalis dillenii Oxalis stricta Oxalis violacea Oxypolis rigidior Parietaria pensylvanica Paspalum ciliatifolium Passiflara lutea Penstemon hirsutus Phlox pilosa Plantago pusilla Plantago virginica Paa campressa Polygonatum biflorum Polygonum tenue Psoralea psoraliaides Pycnanthemum tenuifolium Ranunculus sp. Ruellia humilis Schizachyrium scoparium Sedum pulchellum Solidago juncea Solidago nemoralis Solidago ulmifolia Sphenapholis obtusata Sporobalus asper Stylosanthes brflora T ephrosia virginiana T ridens flavus T riadanis perfoliata Viola raphanesquii ONOOWNOOOOANGOHOkOONOOMOOOOON—o—n—Or-‘OOOOOOOAI—‘AOr—OOOO poooo U) Vulpia octoflara 3 *** 0.5 Woodsia obtusa 0 0 0 ROUND BLUFF SANDSTONE GLADE Number of circular 50 m2 plots for woody taxa: 30 Number of nested l m2 plots for herbaceous taxa: 120 *indicates plant was sampled with the herbaceous taxa and, although bearing a woody stem, was located within the herb layer, i.e., within approximately 1 m of ground level. "*indicates value less than one-one hundredth of a percent. No. of plots Woody taxa of occurrence Dominm Mtive Importance Acer rubrum 1 0.03 1.7 Amelanchier arborea 1 0.07 1.8 Campsis radicans 0 0 0 Carya glabra 23 0.09 2.2 Carya ovalis 0 0 0 Carya ovata 4 0.13 2.7 Carya texana 0 0 0 Celtis occidentalis 2 0.02 0.5 Celtis tenuifolia 7 0.15 3.2 C ercis canadensis 0 0 0 Camus florida 0 0 0 Corylus americana 0 0 0 Crataegus sp. 0 0 0 Diaspyros virginiana 1 0.09 1.8 Fagus grandifolia 0 0 0 Fraxinus americana 6 0.35 4.3 Gleditsia triacanthos 0 0 0 Hypericum stragulum“ 3 0.02 0.3 Juglans nigra 0 0 0 Juniperus virginiana 4 20.24 43.3 Liquidambar styraciflua 0 0 0 Liriodendron tulipifera 0 0 0 Lonicera japanica“ 3 0.05 0.5 Marus rubra 0 0 0 Parthenocissus quinquefolia“ 2 0.09 0.5 Prunus serotina 5 "* 0.4 Quercus alba 1 *** 0.4 Quercus marilandica 2 0.02 0.5 Quercus rubra 0 0 0 Quercus stellata 5 3.59 12.2 Quercus velutina 15 0.04 0.9 Rhus aramatica 1 0.48 4.1 Rhus copallina 8 0.03 0.9 Rosa carolina 0 0 0 Rubus allegheniensis“ Rubus enslenii (IL-E) Rubusflagellaris Sassafras albidum Smilax bona-nox* Smilax glauca Smilax rotundifolia Symphoricarpas orbiculatus T oxicodendron radicans* Ulmus alata Vaccinium arboreum Vaccinium pallidum Vitis aestivalis Herbaceous taxa Acalypha gracilens Agrimonia parviflara Agrostis elliotiana A grastis perennans Allium canadense Allium vineale Ambrosia artemisiifolia Arabis canadensis Aristida sp. Asplenium platyneuron Aster sp. Bidens bipinnata Blephilia hirsuta Bromus commutatus Bulbostylis capillaris Cardamine sp. Carex artitecta Carex blanda Carex cephalaphora Carex glaucodea Carex hirsutella Carex muhlenbergii Carex retroflexa Carex sp. Carex umbellata Cerastium arvense Cheilanthes lanosa C laytonia virginica C onyza canadensis Corydalis flavula Crotonapsis elliptica Cunila origanoides Cyperusfiliculmis HID-50019001900019 No. of plots of occurrence 37 0 ON .5 NOWOOOO‘OOF—OHOv—OOOOWI—OA—‘NOI—O’IOH Domina_nc_e 0.23 0 0.24 0 0.01 0 0.11 0.01 0 0.09 0.01 0.06 0 0.01 0.05 Relative Importance 4.2 0 3.1 0 0.1 0 0.9 0.1 0 0.5 0.1 0.6 0 0.1 0.5 0 0 0 0 2.2 0 0.1 0 3.7 0 0 2.7 0 0 0 4.9 O 0.2 C yperus sp. C yperus ovularis Danthonia spicata Daucus carota Dentaria laciniata Desmodium rotundifolium Dichanthelium acuminatum Dichanthelium boscii Dichanthelium laxiflorum Dichanthelium linearifolium Dichanthelium palyanthes Dichanthelium sphaerocarpon Dichanthelium villasissimum Diodia teres Elymus villosus Elymus virginicus Erechtites hieracifolia Erigeron strigosus F estuca arundinacea Galium circaezans Galium pilosum Gnaphalium purpureum Gratiola neglecta Hedyatis longifolia Hedyotis nigricans Hedyotis pusilla Helianthus divaricatus Hypericum drummondii Hypericum gentianoides Hypericum punctatum Juncus interior Juncus secundus Krigia virginica Kummerawia stipulacea Kummerawia striata Lactuca serriola Lepidium virginicum Lespedeza procumbens Lespedeza repens Linum sp. Manfreda virginica Muhlenbergia sobolifera Nathascardum bivalve Oenothera linifolia Opuntia humifiisa Oxalis stricta Parietaria pensylvanica Paronychiafastigiata Penstemon sp. LII r—b LII O WOALIIOOLIILIION—OOOONo—W-BOGN—OOOOOOI—OOLIIOAI—WO'IOF-‘ONNr-‘NOO-hNN 0.2 0.2 10.5 Phlox pilosa 0 0 0 Physalis virginiana 0 0 0 Phytolacca americana 0 0 0 Plantago virginica 9 0.05 0.9 Poa compressa 0 0 0 Polyganum cristatum O 0 0 Polygonum scandens 3 0.02 0 3 Polygonum tenue 6 0.03 0 7 Pycnanthemum tenuifolium 20 0.62 4.3 Pyrrhopappus caralinianus 0 0 0 Ruellia humilis l 0.01 0 1 Sanicula canadensis 9 0.04 0.9 Schizachyrium scoparium 34 1.90 10.8 Sedum pulchellum 25 0.11 2.6 Setaria glauca 0 0 0 Smilacina racemosus 0 0 0 Solidago caesia 0 0 0 Solidago nemoralis 0 0 0 Solidago ulmifolia 13 0.38 2 7 Sphenapholis obtusata 2 0.01 0 2 Talinum parvrflorum 9 0.20 1 6 T richostema dichotoma 0 0 0 T ridens flavus 0 0 0 T riodanis perfoliata 14 0.06 1.5 Verbascum thapsus 0 0 0 Veronica arvensis 0 0 0 Viola raphanesquii 0 0 0 Vulpia octoflora 13 0.1 1 l 6 Woodsia obtusa 16 0.74 4 4 WILDCAT BLUFF LIMESTONE GLADE Number of circular 50 m2 plots for woody taxa: 18 Number of nested l m2 plots for herbaceous taxa: 72 ‘indicates plant was sampled with the herbaceous taxa and, although bearing a woody stern, was located within the herb layer, i.e., within approximately 1 m of ground level. "indicates plant was sampled with the woody taxa. No. of plots Woody taxa of occurrence Dominance Relative Importance Acer negunda 0 0 0 Acer rubrum 4 0.04 1.7 Acer saccharum 3 1. 14 5 .8 Amelanchier arborea 0 0 0 Aralia spinosa l 0.03 0.5 Asimina triloba O 0 O Bignonia capriolata“ 1 l 0.37 1.1 Carya cordiformis 0 0 0 Carya glabra Carya ovata Carya texana Ceanothus americanus C elastrus scandens“ Celtis occidentalis Celtis tenuifolia C ercis canadensis Comusflorida Crataegus sp. Diaspyros virginiana Elaeagnus umbellata Euanymus atrapurpurea Fraxinus americana Gleditsia triacanthos Ilex decidua Juglans nigra Liquidambar styraciflua Liriodendron tulipifera Lonicera sp. (shrub) Malus ioensis Menispermum canadensis Morus rubra Ostrya virginiana Parthenocissus quinquefolia“ Prunus americana Prunus serotina Quercus alba Quercus coccinea Quercus macrocarpa Quercus prinaides acuminata Quercus rubra Quercus shumardii Quercus stellata Quercus velutina Rhus aramatica Rosa carolina Rubus flagellaris“ Sassafras albidum Smilax bona-nax* T oxicodendron radicans* Ulmus alata Ulmus rubra Viburnum prunifolium Vitis aestivalis Vitis cinerea" Vitis vulpina" NAOI—ONOx\lr—Ot—Ov—-—N~—wv—I—-NOOOBI—WOOOOv—Lfl—OQOOI—AHOGbNN-‘kov— 0.5 14.9 1.4 4.0 10.2 0.5 7.1 0.1 2.4 1.1 2.9 2.9 0.5 0.7 0.8 Herbaceous taxa Acalypha gracilens Agrimonia rostellata A grostis perennans Ambrosia artemisiifolia Amphicarpa bracteata Andropogan gerardii Anemone virginiana Antennaria plantaginifolia Apocynum cannabinum Arabis laevigata Arisaema dracontium Arisaema triphyllum Aristolochia serpentaria Arundinaria gigantea Asarum canadense Asclepias tuberosa Asclepias verticillata Asclepias viridiflora Aster laevis Aster patens Aster turbinellus Aureolariaflava Boehmeria cylindrica Botrychium virginianum Bouteloua curtipendula Brickellia eupatorioides Bromus commutatus Bromus pubescens Cacalia atriplicifalia Campanula americana Carex artitecta Carex cephalaphora Carex digitalis Carex grisea Carex muhlenbergii Carex sp. Carex umbellata Cassia fasciculata Cassia marilandica" Chamaesyce maculata Cirsium altissimum Cirsium discolor Clitoria mariana Cunila origanoides C ynanchum laeve Danthonia spicata Desmodium canescens 201 No. of plots of occurrence Dominance l8 0. 12 0 0 O O O .0 LII 0‘ p—n .9 .—n N \J _o t» O\ L» .o w t» N o»—cam—oo—N—o—oooooo~o~owooo-wo-oo~ooom-o~w p—o .9 0 LI) LII Relative Imp_ortance l .3 0 0 0.1 1.2 0 0.2 0.1 0.8 0 0 0 0.1 0 0 0.2 0.8 O 0.2 1.5 0.1 0 0 0 0.3 0 1.2 0 0.3 O NI— I--\O-- \O or: p—n ¥ Desmodium rotundifolium Dichanthelium boscii Dichanthelium dichotomum Dichanthelium laxiflorum Dichanthelium polyanthes Dichanthelium villasissimum Dioscorea quaternata Dioscorea villosa Dodecatheon meadia Echinacea pallida Elymus hystrix Elymus virginicus Erechtites hieracifolia Eupatorium altissimum Eupatorium purpureum Eupatorium rugosum Euphorbia corollata Galactia regularis Galium circaezans Galium pilosum Geranium maculatum Geum canadense Helianthus divaricatus Heliopsis helianthoides Heuchera americana Hybanthus concolor H ydrastis canadensis Hypericum denticulatum Ipomoea pandurata Kummerawia striata Lactuca canadensis Lespedeza procumbens Lespedeza repens Lespedeza violacea Lespedeza virginica Liatris scabra Lithaspermum canescens Lysimachia lanceolata Manfreda vi rginica Monarda bradburiana Monardafistulosa Muhlenbergia sobolifera Muhlenbergia sp. Nathascardum bivalve Onasmadium hispidissimum Oxalis stricta Oxalis violacea Passiflora lutea Pellaea atrapurpurea La) OHOAOOONONOOt—‘I—O \Jw OOONUJOF-IF—I—I—NOO‘OI-‘NOOONOOOI—t p—n 202 0.33 0.01 0.11 0.11 0.08 0.59 1.15 1.77 0.39 4.45 0.54 0.06 0.30 0.06 0.11 0.96 0.25 0.11 0.63 0.79 0.01 0.08 0.17 0.01 Phryma leptostachya Physalis virginiana Physostegia virginiana Podophyllum peltatum Polygonum cristatum Pycnanthemum tenuifolium Ratibida pinnata Ruellia caroliniensis Ruellia humilis Sanicula canadensis Schizachyrium scoparium Silphium terebinthinaceum Sisyrinchium albidum Smilacina racemosa Smilax herbacea Smilax pulverulenta Solidago caesia Solidago petialaris Solidago sp. Solidago ulmifolia Sorghastrum nutans Spigelia marilandica T aenidia integerri ma T ephrosia virginiana T haspium trifoliatum T radescantia subaspera Uvularia grandiflara Verbesina helianthoides Verbesina virginica Vernonia gigantea Viola sororia Viola triloba LII 4:. \OQOOI-‘OOOOOI—ONOLIIOOOOOO ...-t 203 0.06 1.36 0.46 0.01 0.01 1.35 0.01 0.62 1.90 16.34 oeowoooooo 0.2 3.6 1.1 0.1 0.2 2.5 0.1 1.9 3.5 20.1 eooooporvogxoooooo LII ON '-‘ 00 00 ¢coo \ILII APPENDIX B Maps for each of the studied forest-openings. 204 205 Figure 8a. Map of Berryville Shale Glade (unmanaged). 206 of Opening .4 207 Figure 8b. Map of Brown Shale Barrens (managed). 208 ~44447BelLJransect (14:14 444444.444-4-44. 7 '. I l "3 1 . :Permonent Plot (soImZ) (lnstalled1989 by A. Heikens) . I ; 5 - 3 ‘ 1 ‘ ' ‘ T - o ‘ ' ~ ‘ . i . ‘ ,4... 4.4.7 - 7 .4 -7 7. . .4 7. 4- ..4 4777;774 771.4 4 447 - .4 4 4 7. 4 - - 4 .4... 4 4-444.444444 1 I . . l n: Not . Preserve . . , I ‘_4__,-_,4_4_4 7 v 7 L— - II 7 I 7 7. ‘I‘rODse’CtéY #371 7 77. -7. 7-77. 7.77- 4 .4 . 7;7_-44|.47.447. eef—BrewnI-ghelle Barrens 1 . ; ‘ ‘ l 1 I TL 3 Y I I 1 ' - i l 1 WmSeoes-Merers I . i 4 * 72" l l bcole 1L I J-lilt II LLECEND i ' . I . : 6 I I 4...-..— ...—7 l | 1 1 ‘ l l 1 ApproxL Lated BqufLEggeL 4.47 1 1/1 jApproximated Foresl Edge ; I 4.4 7.7.7— 7 .7 74 4 4-. l l I I I I l l I l I I .1441 4 . 'Jfll'lfl'lflL. LLocation HofI Measured Bluff Edge 1 ‘ 4444-4444 44444 L ‘ ‘1xxx ‘Location ofI Trees of MeaSured Forest Edge (275% Canopy Cover and 26.6 cm DBH # lMorker Tree with MeasUred DBHi l ' I I l 3 :1 -“‘“" 444 T ‘7 1 Ww‘ l l l 1 V1 %41 lfiEkTéFT‘ “F . I I I 7 I 1 g T. ‘ ‘dercus'h stellata—1 44 444 44 e - 4 e % I» -' ’4—4.¢' ee—e- Date 4140‘) Prepared August. 1994 ‘ 1 . ‘ ‘ F1 cm)! . , . , ,__4..___.. _4 -+— 44 ee — I - I . ‘ I I 4-4. 4 474 ,47 447—..47-‘ l 77 4474.— 4- 4.7 7 774. 7 .7 . . 7 7-. Q 7 "77 e—~—-7.4_4,.4 747.1 144444-74 4’ ' 7' ”Tro—fisfiecf '4" 7+ _4__-_.4__4 A'fi'FCT'T." " i i "1 "7'" GETS—'77 +F‘7' 1 . l ...—4.4-44-7. 7 .7 77 - 7 . 7 .7 ,. 77 7- .777-770-7— ,__._4.-4 '7-7-“ 4 -7‘ 777 77 .7 4 0 77 .7 7777774777 1 I ‘ l l I l I . 1 1 1 1 I I , I ' ‘ ‘ ' I l I 1 I I ‘ , - 7. -. - 77 . . —_ 4__7._7. 7 44 477.4 4... ‘ I 777777.». . 7.7 7 12ng )stellata J4Qumus s==:-llata(415cm)44_..-14 ‘ cm I ; L444.4.47_7 7 4 .47 4 I 7 . “-747 4 777.777 .777.— 77 .7— 7—777 47. 4-7. 4. . .7 -4. 4 7- 44. 4.47444. 4-- l I . . , l 4.4 7 4 74 4 - 7 4 4 7 447 4 4 4 4 - 74- 4 .4 4.47 7 .7 . .7 7. 7. 77.. 77.4 74444 4.. I I . ' I I 1 1 . . ‘ . I 1 I I I . . 77 47.47 7 7 .7 7 . 7 . 4.77 7.7777 .4777- 7 _ ... 77*— 7777—77— -4.— 7— 77- 7. . .7. . 47 7 47 .44 4.. .744..4.¢4 __.. I I I I . ‘ ‘ Transect 3 I 1 . . 1 1 4. 4,4 ....4. 7 7. .7 4 . . -. . 74 . 7 _7—-7 - . - - I 7 7 . - 7 . I I I I I ‘ I I l 7- -. - 7 7. . ¢ ¢ .7 7. 77. 77777.. .774 - .477 77.. 7- 74—7‘ — 7 . 7 . . - . ..7 74-7—7774774.. ‘ I 44 4 7 1 I 1 I 4 4 4 4 4 44 4.4 4..- 4_-4 44 4._- . - . -7 .- .. 4 4.4.4 4444.. I I I l 4 4 47 7 4 7 44 4. .444 4 -47 7.4. .4 4 4. 7 - - -.. . 7- _ 7. 4 -_4_44 444.4 I I .4 4_ 4 - 4 4 4-, ..4447 -. ... -. _ . . 7 7. 7 . 4?. 44744.... 77 4. I— 7 4. 7 . . .7 ._ o- 7 — .7 -74444.4 4 .744. 7 . - 7. 777- 7. 7 II .7 44.774.44.74- 4-. I . . I ‘ I 744.4.-. 7 . 7 -....7 .7. .. . 7. 4 44.4-44.447.4-4 I | ‘ I | . . I - r o I t ’ O O 74 7 4 4 7777—. 4 4 7 _77. 7 4 7 44 I : 4 4. 4 - 7 77 .4 - 7 7 7 7 7 _77 -— -7—H 47 4 - .4 . 7 .— 7 7 77 4. 4 —— 4 4 ' 7. 77 , 7 7 7-47 44 .44 4 4 4 4- - 4 7 I 7 . 4 7 4 7 4 T' 7 L , 4 4 4.7 - 4 .- 4 I4 4 . 1.7. o 7 7 wen—1 209 Figure 8c. Map of Cave Creek Limestone Glade (managed). 211 "r Figure 8d. Map of Cedar Bluff Sandstone Glade (unmanaged). ~ ‘m 213 Figure 8e. Map of Gibbons Creek Sandstone Barrens (managed). 215 Figure 8f. Map of Gyp Williams Sandstone Barrens (unmanaged). 217 Figure 8g. Map of Pounds Hollow Sandstone Glade (unmanaged). r r fi T l LPWM$H¢UI©W~$®MI©§ III@7@I I r I j >777II7 77g) I 7 Id. — — —@ -II— 1 777,- .7. I777I7__J 7 I § I I I I 747 L I 7 7. 7‘, .1 4. *7 7. I_"‘ ] __J._._ _. l I l _ L I l I Scale I I ‘ I' : 7 I . - L,— I I I L r r r I ' I I I ** I . I; I: ' I I I T - s I I LEGéN9%L i JV 4 I l I I I I L i I [77%. I I 7 ' L L7 I 7‘ 7i I I I I I ‘ l I I l I i I I 2 I \ 2. l i I I I I \ J l i I VT I k I 'T— lo T I I Y r I l I I l j I I f I T I I I V“ I I I .5: I App ‘ I f gfl I 7 ' ' ' I l I . I I I‘ I l I L I V I , I I I I f I fi I I i ‘ 7 I . '77 I I I l Im'igd-lF_QI§_SLE£ig§7 I 7 I I V I ' ; I I I I I I i I I I I l I I I I 3 ‘. I I f ' I I II/ Approximated Paved FoOtpoth ‘ I I ;___I I I I I I I I I I I I J I l I I I— I I I I I t I . I I Q I I I I I F f ”Hf—‘3 . . I Mammml7777777 3 I I I '7 I I ; I2 2 I I 7 7 I #4 LIN?“- Location of; Meoswed Bluff Edge I I 4 I. I I I‘ I j I I I . 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I , 7,77 7,...7._ 777 -, .77.- 7.7 V -7 , I I I I I I I I I I I I I I . , I I - 7 ..L .7 7777.77.77.77. .7777- .. 7777 7... 7.7-. 77777-.. 7.. 7.7 . 77,‘ I I l I I I I I I I I I . V I I . I l I . I 7. 77 _ 777—7.- . -7777 - 7 7.7—77.7 l- .7779— ‘ 7 7 I 7 7. 7 f _._r.._ _ . 7 A I I ‘ I 7.77777 7.77 77 “—7“ ‘*:"Tronsect 3 ' _. 7 7.7.77.7 I . 77 777.7, 77777_. I ' I I I . l I I I +77+—o——+——77__.—_._.._._ .777 . 7. I l V I I, I 747777777777 ..77V 77 .77. 777 7- V77... _ 7 -..7. -77.. . -7 777.777._V 7 . I I I I I I I I i | I I 7.7,_. 777-7--7.+77.__ .77--.- .-.. 7 7. ,7 I ‘ I I I I l I V ._77.7 7- _7¢—7-._77—7.7_7..—.7_.._———.7 .7-_. I I I I | .77-_,7 . 7.777 7 7 .777. I I I I 777 77.- 7 ”7777—7777....— ___.___. - 7 I I I I l l . .7..-— 7- 7.— _.—__._——7 __.___7-7_77.7 +_- I t I I 717777. 7-7-7-7777 ronsec 77777.7 7 7 .7 7- 7.77 77.77 77 . I I I 1 I H77. 77- --7 -77. 7p—.—L77+777-—§--~ 9 ~--~—»- » 7-7 7 .7 7.7 .777..— -—.77_._—— —_.—-—— o—————o— —.——-— ‘— I I . I ‘ ‘ I I 777-777...- .7--.7 +7. ...7 7.77 L7 77 .7 - - - .77-- .7 7.- 7.7. ...—7.7 -7- 7.- __. I _‘ I I I I I i i I I I l I 71,7777?— - - 71.7—77.7..- _ . 7 .77 _ 7 7 7-- +7 .7777..— 7777 4777 I l I V‘ I . I _ I I I I I ———o——.77L77+-7——.————r- 77+—-._7_ 77¢ -o--~~ 7.77 7 7 7 - 7. 77.77 .__... 777‘ o 77.7- . — ‘77—7 o————>—A. 77777—- 7_7—. 7-7- I l ‘ I l I I I I I ‘ ' , I 7.7-77-7.— 7'77 7.— »— 4———‘ —a.—~o - 77 ._ v 7 o .7 7, ..__ ... 77 *7 -77 — I I I I I I I I I 7.7—777.77. r—7. .— o- 47 —. 77- ——> 777. o . 777 .7 - -7 o I l l I I 7$___..r7_.__. 7777 7.777 77777— 7 .7 79- 7 - o— -7 7 7 - 77.7 -7 7 7 .77- I I I I I ‘ I >777;_—77._777._7—777 ‘77.. 7..— 7 7; 77—77- _ 77 77 7 7 . 7 7 . 7 - . l l I ’ I I I ‘ ' 77777.77 7.77.7. 7.7» e —— 9 7. ~ J - .. - —— 7. .—— . o—— 7.77 .— .7 7 I I I..-» 7.-..7..-» 7.7. 7._ .7777, 7 .7 .7- 7 77777 - 7 _ __ _- _ I L777-_7 77 77 7 777— _777- .7 _ 77 7 7 7- -7; 7 77 ~ “its. .- 4‘... 7‘1 219 Figure 8b. Map of Round Bluff Sandstone Glade (unmanaged). I A 7—777.7> 777 o7—o-o I I If I Vfi II I ~—‘oI-—mi> I I I *7 97—-—-~——4— —7 + 077II 1"- EVE—o I V 4 ‘T—I ’T": I77‘I I 7 B:@: | | I _ I ‘I_ I I i ‘ I I I77 I I I 7 ‘ L I I IE I yr f—V II‘II' 'Ih‘r I 1771 ‘IVIIIT I I- I. I 225—Meters 7.777 7 I77 I. I I 7. I——I-:qp- :9; I I. I I I I I I I I l I I I | I l I T- I" I I I I I 71 uw-— I IaW’fiWIo Ied‘FofreWEdqu II 1[I’LBelt Tronsecf (5 m wide) I ""I" ’I 7I 47 I77 7 - ”Huger” ' -—I7 7 7-7I7—I77 Wobm7of7uaosuce¢8w¢£7£ I7’ I ~ 7 I 1 I I 7 :7? I WUOCCVWOH?E"‘VFM§WWWPéféfl’fdge,V if I ' I I I ' ,rfi'rd-I—fifirj III I 7 I) 75VTCon0pIy Cover and 2661chDBH) T . IF: I V IL TI 7+7—7T7‘0: ‘ I‘hMWI‘MHh‘ Meosmr'edI—BBI-l~~—e I- I V 7V I7I I I I 7 E I .I II aneosurea‘RegIon of OpenIInglIL ’ I I i . I - f' j V I f‘T’f—“P "““I'II 7floIe+Mop Prepared: JuIy771994r—+7‘ I I i j V 2 I7 , j I7? F—V I -..—- 7.777. 77 .7 7.77777 77 Y 7' o 777 77.- I | I I I I I . I I I I - I I I I I ‘ I I I ' I 777777777.7 .7 7 _I77 77 7.— 77. 7.77 7 7-77. 7-7-.7777 77.7 77 7 .77 . 7 .77. I I ‘ I I I I I I I I I I V 77747777777777.7777 .7 7 7 7.7 7 7 77 . 7.7 77.7 v—77o—79—7—77—97—o—77— ‘ 7 .- 7.74 I ' 1 V I I I I V . I I I | I I I 77777777... 77 7 7.7 .77- .7 7-7 . 77 77 7 7.77 77.777.777.7— 77-77-77 7 -777 7.77.77 .7 7 7 7 - I I I I I L74— 777 .7.— .7 7.7 7 . - -7 -7 .7 .7 7 - 7 . 77- 7.— .-7 .7777- .7... . 7.74 I I ‘ I I I I 77.77.77.777 .7 77.7 -7 777 .7 -7 - 7 7 .777- 77.7 777.7—7I79—7— 7.7 7777.77.77.— 77-7- 7-7—777*-7 . . 7 . . 7 7‘ I I I I I l = ‘- I 77. 777—-7 7—77- . 7 . J7 .77 . 7 <7 o o- ‘ 7 7-777777777.x—.7 7. -7 7 . .77... I § L77777 .7 77477777 7- 7. 777.7 7.7 7.7 ,7 .7. 777 .7 7.- 77.7. 77 77777.77..- .777.7 .777 77 -777... .7 7.7.77 7 . . .7 7.77. -r... .7 . o .7 . . I V I I I I I . 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M r‘ 3 ‘ { , ' ‘e‘tes J I ‘ . ‘ ‘ \ ‘ ‘ ‘ ‘ ‘ ¢ \ ‘ ‘ , ‘ ‘ 1 x ‘ ‘ ‘ ‘ x —‘—r*7" ‘ M_.._ ‘ WkH—L'—E*~fi++“fi . 1 \ ‘ l Transect 1, ‘ ‘ I l ‘ ‘ I ‘ ‘ ’ 1 I Wm“: 1 ‘ __ s| e a. a \ ; ‘ . w ‘ 3 ‘ ‘ ‘ S j ; ‘x * — w—Po _ __ -...774 “I l l__,__,_4__ ‘ +14% _JI:_L I APPENDIX C Species lists for woody and herbaceous taxa in five subhabitats of the study sites and summary of relative importance information relating to species for the sampling in 1994. Relative importance is the sum of relative dominance and relative frequency divided by two and multiplied by 100 (yielding a percentage between one and 100). Seedlings were defined as stems <2.54—cm diameter at breast height, shrubs and saplings as stems 22.54 to <6.6 cm dbh and trees 26.6 cm dbh. FI- N=Forest Interior-North, TS- =Transition Zone-North, OP=Opening, TS-S=Transition Zone- South, FI-S=Forest Interior-South. BERRYVILLE SHALE GLADE Woodv Seedlings: Acer saccharum Acer rubrum Amelanchier arborea Carya glabra Carya texana Celtis tenuifolia C ercis canadensis Comusflorida Diaspyros virginiana Fagus grandifolia caroliniana F raxinus americana Juniperus virginiana Ostrya virginiana Prunus serotina Quercus alba Quercus imbricaria Quercus marilandica Quercus rubra Quercus stellata Quercus velutina Rhus aromatica Rhus copallina Sassafras albidum Ulmus alata Vaccinium arboreum Vaccinium pallidum Bi! 0 O 24.5 1.7 1.8 0 0 1.7 5.2 1.7 1.7 O O 0 3.5 1.7 5.7 3.3 6.7 8.3 O O O 6.9 3.7 22.1 TS-N 3.6 18.8 2.7 21.9 7.8 4.6 10.9 223 O '1: tool Or—Mowco N!— 14.1 3.1 17.9 6.6 4.3 12.6 1.2 1.3 7.6 26.0 12.3 11.4 3.1 2.5 6.6 0‘99r‘NCF‘ONr‘E‘NOOHP’V‘ON'fi AAN-h— \) \ihxiq :H" \J; 13.8 9.3 12.2 3.1 Shrub/ Saplings: Carya texana Fraxinus americana Quercus marilandica Quercus stellata Sassafras albidum Ulmus alata Vaccinium arboreum Vaccinium pallidum Trees: Carya glabra Carya texana Quercus marilandica Quercus stellata Quercus velutina Herbs: Acalypha gracilens Amphicarpa bracteata Antennaria plantagim'folia Asclepias tuberosa Aster patens Aster sp. Carex hirsutella Carex sp. (Montanae) C unila origanoides Danthonia spicata Dichanthelium linearifolium Euphorbia corollata Galium circaezans Hedyon's longifolia Helianthus divaricatus Lespedeza repens Lespedeza sp. Paronychiafasn'giata Penstemon sp. Psoralea psoraliaides Rosa carolina Sanicula canadensis Schizachyrium scoparium Solidago nemoralis Solidago sp. T ephrosia virginiana T oxicodendron radi cans Vitis aestivalis FI-N 20.5 10.3 45.5 23.7 FI-N 62.5 37.5 2.1 'z .0 \l .‘lf‘ Nb \0 \o .9 \) ofboyoo—ooooogzoooooowwooooo—oo o 224 TS-N 47.2 18.1 34.7 TS-N 50.0 50.0 TS-N 00 AA w .°°.U' w #0 A co;ooooooxoooaxoognoow—ooognoo ko O *6 v-‘\O OOO ONOOv—UIOO 5° N owaooo 9’9" 1: N00 \l ”U N N m 39.9 0\ ooporboooooo N 10.5 12.8 5.3 5.3 2.3 FLS 14.3 14.3 14.3 14.3 42.9 gs 1.3 5.4 3.8 3.8 9.2 1.6 4.7 15.3 4.7 1.6 2.9 1.6 15.3 10.5 3.8 4.5 1.6 1.6 5.4 1.6 Vulpia ocloflora BROWN SHALE BARRENS Woody Seedlings: Acer rubrum Acer saccharum Amelanchier arborea Carya cordiformis Carya glabra Carya ovata Carya texana Cercis canadensis Camus florida Crataegus engelmannii Crataegus sp. Diaspyros virginiana Euonymus atropurpurea Fagus grandifolia caroliniana Fraxinus americana Gleditsia triacanthos Juglans nigra Ligustrum vulgare Morus rubra Ostrya virginiana Prunus americana Prunus serotina Quercus alba Quercus imbricaria Quercus marilandica Quercus prinaides acuminata Quercus rubra Quercus stellata Quercus velutina Sassafras albidum Ulmus alata Ulmus rubra Vaccinium arboreum Shrub/ Saplings: Quercus stellata Ulmus alata Vaccinium arboreum FI-N 225 0.7 13.7 2.3 1.4 1.7 2.4 2.7 14.9 8.1 5.2 0.8 TS-N 000 O 1: cor—o —n 6.5 10.9 12.0 13.2 QB 50.0 50.0 TS-S ¥P¥OONO WNCh '— \J N OMHOfiOpJOf—ObepJOOQOOOOOOO so: \I A 16.6 26.0 2.2 O O\ .mpoosoorvoaxooooo N O Trees: Carya glabra Carya ovalis Quercus stellata Ulmus alata Herbs: Acalypha gracilens A grostis perennans Ambrosia artemisifolia Amphicarpa bracteata Antennaria plantagimfolia Arabis laevigata Asclepias variegata Asplenium platyneuron Asplenium sp. Aster patens Aster sp. Bromus pubescens Carex glaucodea Carex hirsutella Carex muhlenbergii Carex retroflexa Carex umbellata Chasmanthium latifolium Croton monanthogynus Cunila origanoides Danthonia spicata Desmodium nudiflorum Dichanthelium acuminatum Dichanthelium boscii Dichanthelium dichotomum Dichanthelium laxiflorum Dichanthelium linearifolium Dichanthelium sphaerocarpon Elymus villosus Galactia regularis Galium circaezans Galium pilosum Geranium caralinianum Hedyon's purpurea Helianthus divaricatus Lechea tenuifolia Lespedeza procumbens Lespedeza repens Lonicera japom'ca Luzula multiflora Manfreda vi rgim' ca FI-N 25.0 50.0 25.0 FI-N 0.6 10.8 1.2 1.7 0.6 0.6 226 M 15.6 10.6 63.3 10.6 17.9 0.7 0 1.4 O '1: o-—-oo o .0 0 col poooor—IO m M'U ONO & 1.4 3.3 0.5 5.1 1.4 5.2 6.0 11.9 0 Ow—oo—y o o S” 'o m ...] 9” m oaxooooooo 10.6 15.7 1.8 11.3 ’11 5,", 100.0 E 0‘ I (I) ooopop ox Af—OOOO \OUJQ 1.3 O in ooooepoopoowoqooopoooo WA DJ t—iI—i use y—n NOOf—O .5 U) .fim—n i Muhlenbergia sobolifera Oxalis sp. Parthenocissus quinquefolia Passiflora lutea Penstemon sp. Plantago virginica Poinsettia dentata Pycnanthemum tenuifolium Rosa carolina Rudbeckia hirta Ruellia caroliniensis Ruellia humilis Sanicula canadensis Schizachyrium scopari um Solidago caesia Solidago nemoralis Solidago ulmifolia Stylosanthes biflora T oxicodendron radi cans T richostema dichotomum Woodsia obtusa CAVE CREEK LIMESTONE GLADE Woody Seedlings: Aesculus glabra Acer negundo Acer rubrum Acer saccharum Amelanchier arborea Aralia spinosa Betula nigra Carya cordiformis Carya glabra Carya ovalis Carya ovata Carya texana Cassia marilandica C eanothus americanus Celtis tenuifolia C ercts canadensis C ornus drummondi 1' C omus flortda Cornus sp. Corylus americana Crataegus engelmanniz’ 0.6 8.7 1.2 0.6 3.7 1.2 0.6 0.6 1.8 0.6 1.2 9.7 0.6 M 0 0.7 0 0 1.7 2.7 0 1.5 0 0 0.7 3.9 0.7 0 9.4 7.7 O 1.5 0 0.9 0.7 227 TS-N O 1.4 ooopo A \l N ooocwo¢~c~¢¢c~ \Ov—'\O\Ov-‘ QB 0.2 2.8 0.3 1.5 0.4 0.3 0.4 0.9 0.4 2.4 8.4 8.5 0.9 3.4 0.6 A h N O\ 4} ooopopopog—OOOg—Og—ooooo N -i S” m opo LII OOOOO OEOOOOOOOOOOO DJ Of—Of—OQOF‘ w m N .O‘ \0 fi‘“ :___ i ..b Crataegus sp. Crataegus viridis Diaspyros virginiana Euonymus atropurpurea Fraxinus americana Gleditsia triacanthos Ilex decidua Ilex verticillata Juglans nigra Juniperus virginiana Lindera benzoin Malus ioensis Morus rubra Ostrya virginiana Prunus americana Prunus serotina Quercus marilandica Quercus prinaides acuminata Quercus rubra Quercus shumardii Quercus stellata Quercus velutina Rhus aromatica Rhus glabra Sassafras albidum Ulmus alata Ulmus rubra Viburnum prunifolium Shrub/Saplings: Acer saccharum Carya texana Celtis tenuifolia C ercis canadensis Camus drummondii Cornus florida Diaspyros virginiana Fraxinus americana Juniperus virginiana Ostrya virginiana Quercus prinaides acuminata Quercus shumardii Quercus stellata Ulmus rubra Trees: Acer saccharum 228 0.8 0.2 2.7 4.1 4.4 0.7 1.6 4.5 0.7 3.5 2.6 4.0 0.9 2.8 19.9 0.6 0.4 5.0 1.7 6.3 0.2 1.7 0.3 2.2 2.6 0.6 1.3 0.2 1.2 12.7 '-i S” m .o oo ooooogooooqooo 'N CH 9” (A \O LI’IU—lt—l r—n \O \0 OD) opoooowoppooocowopooog—g—oyu LII O '1'] '5; Carya texana C ercis canadensis Camus drummondii Carnusflarida Diaspyros virginiana Fraxinus americana Juglans nigra Ostrya virginiana Quercus prinaides acuminata Quercus shumardii Quercus stellata Ulmus rubra Herbs: Acalypha gracilens A grimania rostellata Ambrosia artemisifalia Amphicarpa bracteata Apocynum cannabinum Aristolochia serpentaria Asclepias sp. Asclepias verticillata Asclepias viridis Aster ablangifalius Aster patens Aster pilosus Aster sp. Bignania capriolata Boehmeria cylindrica Batrychium virginianum Bouteloua curtipendula Brickellia eupatorioides Bromus commutatus Bromus pubescens Bromus racemosus Campanula americana Campsis radicans Carex artitecta Carex blanda Carex cephalaphora Carex meadii Carex muhlenbergii C arex retraflexa Carex sp. Carex umbellata Cassia fasciculata Cassia marilandica C elastrus scandens 00 00 oooooooopopoooc 00 ooooooooeooooog—oocooow N 0 8.3 8.3 16.7 8.3 33.3 25.0 QB 0.1 0.3 0.2 0.5 0.2 0.1 0.3 4.2 1.4 0.2 1.3 1.3 0.1 0.6 33.1 0.1 0.5 0.1 0.1 0.6 0.1 0.1 0.1 0.3 0.1 0.1 0.3 0.2 0.1 OOF‘ONOOO 9° .0‘ b.) \I 36.7 18.3 -l S” m Chm DJLII 00 \O coopooooocowoopoooooooopg—ooor-g—oooo OO OOOOO '11 hr: (I) N r—KII M N DJ Chamaesyce maculata Croton monanthogynus Crotonapsis elliptica Cunila origanoides C ynanchum laeve Danthonia spicata Desmodium canescens Desmodium paniculatum Desmodium sessilifolium Dichanthelium boscii Dichanthelium laxiflorum Dioscorea quaternata Echinacea pallida Elymus virginicus Erigeron annuus Eupatorium altissimum Euphorbia corollata Festuca arundinacea Galactia regularis Galium circaezans Geranium maculatum Helianthus divaricatus Helianthus microcephalus Hybanthus concolor Hypericum denticulatum Ipomaea pandurata Lactuca serriola Lespedeza cuneata Lespedeza procumbens Lespedeza repens Lespedeza sp. Liatris scabra Lithospermum canescens Lonicera japonica Lysimachia lanceolata Manfreda virginica Medicago Iupulina Monarda bradburiana Muhlenbergia sobolifera Onosmodium hispidissimum Oxalis stricta Parthenocissus quinquefolia Passiflora lutea Phlox pilosa Physalis virginiana Physostegia virginiana Polygonum cristatum Ratibida pinnata Rosa carolina 00 \OA 00 00 O\ p.— b.) QNOOOCOOONOOOOOOOt—OMPJOOf—OOOg—ONOOOOOQOOO a CO 000’\ 10.2 230 ON (It ooooopooogopooo KI! -—-4> Nb.) 1.2 0.6 0.6 0.6 2.5 0.4 0.8 0.1 0.1 0.1 0.1 0.1 0.1 0.6 0.1 0.4 1.7 0.4 0.1 2.8 2.9 1.4 3.2 1.4 0.1 6.2 0.1 0.4 0.6 0.3 0.4 0.2 0.4 0.6 0.3 0.4 0.2 1.6 0.4 1.7 0.5 0.1 0.7 0.3 1.6 1.2 3.4 0.3 1.6 0.9 v—OO ocw cow owcoow v-' 0000 CW \IOO opg—opooppoooooow com 00 owowoéoooooh 00$ 0000\1 \O Lit ooooowoor—oooooo occ \J\l Of‘OQF‘f‘f‘J‘Q N \imowmq \O oeoooooooeooopoooooo .--‘O N“ Rosa multiflora Rubus allegheniensis Rubus enslenii Rubusflagellaris Rudbeckia hirta Ruellia caroliniensis Ruellia humilis Salvia azurea grandiflara Sanicula canadensis Schizachyrium scoparium Scutellaria leonardii Setaria glauca Silphium integrifolium Silphium terebinthinaceum Sisyrinchium albidum Smilax bana-nox Solidago sp. Solidago ulmifolia Sarghastrum nutans T aenidia inte gerri ma T oxicadendron radicans T radescantia subaspera T rifolium campestre T riasteum angustzfolium Verbesina helianthoides Verbesina virginica Viola sororia Viola sp. Viola triloba Vitis aestivalis Vitis vulpina Zizia aurea CEDAR BLUFF SANDSTONE GLADE Woody seedlings: Acer saccharum Amelanchier arborea Carya glabra Carya ovalis Carya ovata Celtis tenuifolia C ercis canadensis Diaspyros virginiana Fagus grandifolia caroliniana Fraxinus americana 00 ego—omoooooeNOOg—ooooo mu 3 1» com ~oopig—ooo 00 WOO ooo. ELM 1.4 2.9 9.4 0.9 11.3 4.8 0.7 1.7 0 6.6 231 0.4 0.3 0.1 0.1 0.5 3.5 1.0 0.8 7.9 0.1 0.1 0.7 7.5 0.4 6.3 0.3 2.3 2.5 0.8 1.1 0.1 0.3 0.1 4.3 0.4 0.1 1.2 opooo \IO 0\ H N p—u U!“ mp A owgnoogvv—oooogpowogognoooopaoQg—ogvooo u—N N r3“..- Juglans nigra Juniperus virginiana Ligustrum vulgare Malus ioensis Ostrya virginiana Prunus serotina Quercus alba Quercus marilandica Quercus rubra Quercus stellata Quercus velutina Rhus aromatica Rhus copallina Symphoricarpas orbiculatus Ulmus alata Ulmus rubra Vaccinium arboreum Shrub/Saplings: Amelanchier arborea Carya ovata Celtis tenuifolia Diaspyros virginiana Fraxinus americana Juniperus virginiana Ostrjya virginiana Quercus rubra Ulmus alata Trees: Carya glabra Carya ovata Diaspyros virginiana F raxinus americana Juniperus virginiana Quercus stellata Ulmus alata Herbs: Acalypha gracilens Arabis laevigata Aster sp. Bromus pubescens Carex blanda Carex cephalaphora Carex glaucadea 0.4 6.6 0.7 0.4 3.3 0.4 2.1 1.1 2.4 11.8 4.7 9.9 2.3 2.4 9.4 0.5 1.7 F_I-B 3.4 3.4 3.4 4.9 9.7 33.5 6.8 3.4 31.6 M 2.4 2.4 2.4 10.6 40.2 9.8 32.1 M 2.0 0.5 0.5 3.6 0.5 1.0 0.9 232 00 oxooooopo O O 25.1 4.8 4.1 18.6 32.8 0 OOOOO Carex hirsutella Carex muhlenbergii Carex retroflexa Carex umbellata C elastrus scandens Chasmanthium latifolium Cheilanthes lanosa Cunila origanoides Danthonia spicata Dichanthelium dichotomum Dichanthelium laxiflorum Dichanthelium linearifolium Dichanthelium villasissimum Elymus virginicus Euphorbia corollata Helianthus divaricatus Juncus tenuis Lactuca serriola Lepidium virginicum Manfreda virginica Muhlenbergia sobolifera Myosotis verna Oxalis sp. Parthenocissus quinquefolia Penstemon sp. Phlox pilosa Polygonum cristatum Polygonum tenue Rubus enslenii Sanicula canadensis Schizachyrium scoparium Smilax bona-nox Solidago caesia Solidago nemoralis Toxicodendron radicans Vulpia actoflora Woodsia obtusa 0.9 0.5 0.5 1.8 1.4 8.5 0.5 0.9 11.9 2.3 0.9 6.9 2.9 0.9 0.9 0.5 0.9 1.0 1.9 10.4 0.5 3.2 1.5 5.1 0.5 2.8 1.5 18.9 233 36.5 GIBBONS CREEK SANDSTONE BARRENS Woody Seedlings: Amelanchier arborea Carya cordiformis Carya glabra Carya ovalis Carya ovata .111! 4.4 1.7 2.9 o 3.9 18$ QB 1&8. 1.7 6.1 3.8 o 1.8 2.1 1.2 0.8 1.3 1.3 o o o 0.4 1.3 M 2.6 4.6 Carya texana Celtis tenuifolia Cercis canadensis Cornus florida Crataegus sp. Diaspyros virginiana Fraxinus americana Juglans nigra Malus ioensis Morus rubra Ostrya virginiana Prunus americana Prunus serotina Quercus alba Quercus coccinea Quercus imbricaria Quercus marilandica Quercus prinaides acuminata Quercus rubra Quercus stellata Quercus velutina Rhus copallina Symphoricarpas orbiculatus Ulmus alata Ulmus rubra Vaccinium arboreum Vaccinium pallidum Shrub/ Saplings: Amelanchier arborea Carya ovata Carya texana Diaspyros virginiana Fraxinus americana Juniperus virginiana Quercus marilandica Quercus stellata Ulmus alata Vaccinium arboreum Trees: Carya glabra Carya ovalis Carya ovata Carya texana Fraxinus americana Juniperus virginiana 12.3 0.9 1.1 3.8 3.9 0.9 2.6 1.8 0.9 4.7 2.0 3.2 8.6 5.5 23.1 10.3 FI-N 9.5 9.5 19.1 14.1 9.5 234 12.9 1.1 1.1 3.7 1.1 1.7 1.5 1.4 “Sr-98.30 r—O'xr—I h-—-\o O 21.2 25.6 0 000 9.5 0 14.1 0.8 0.4 1.1 2.4 1.5 0.5 0.4 1.7 0.5 0.4 7.5 7.9 1.5 16.3 7.1 7.4 9.2 0.4 0.5 9.5 g 14.8 14.8 8.1 MAOOOOOOOO .U'P' OO '-1 S” (12 oo-‘HOO 9°.“ DJQ 10.6 '11 c7: .0 o ogoooooomo 'o I? \O l U) OOt—‘OOOO 5’ . OO Quercus marilandica Quercus stellata Ulmus alata Herbs: Acalypha gracilens Ambrosia artemisifolia Antennaria plantaginifolia Aristida sp. Asclepias verticillata Asplenium platyneuron Aster patens Aster pilosus Bromus pubescens Carex artitecta Carex cephalaphora Carex hirsutella Carex muhlenbergii C arex retroflexa Carex sp. Carex umbellata Cassia nictitans C litoria mariana Crotonapsis elliptica C unila origanoides Danthonia spicata Desmodium rotundifolium Dichanthelium acuminatum Dichanthelium boscii Dichanthelium laxiflarum Dichanthelium linearifolium Dichanthelium sphaerocarpon Dichanthelium villasissimum Elymus virginicus Euphorbia corollata Galium circaezans Galium pilosum Helianthus divaricatus Hypericum gentianoides Ipomaea pandurata Koeleria macrantha Kummerawia striata Lactuca hirsuta Lechea tenuifolia Lespedeza procumbens Lespedeza repens Manfreda virginica Monarda bradburiana 28.6 9.5 Fl-N 14.9 pooooo 16.9 44.9 10.2 m 7.9 6.1 0.9 0.9 1.9 0.6 1.7 0.3 2.9 1.2 1.2 6.9 0.3 1.2 0.3 5.0 1.3 9.6 1.3 1.6 2.8 4.1 2.2 8.6 0.3 3.3 0.3 0.6 0.6 1.9 0.9 0 43.7 25.4 -1 E” U) M000 Nooooogujngn 12.8 .73 5b): (’3 ooooooooiuos VOHCOfiENNOONWOOO+OWNPO & U) bow-Ab $0 00 N30 16.4 0 OOOOOOOOO Monardafistulosa Oxalis stricta Paronychia fastigiata Parthenocissus quinquefolia Passiflara lutea Phlox pilosa Physalis virginiana Polyganatum biflorum Porteranthus stipulatus Potentilla simplex Rosa carolina Rubusflagellaris Rudbeckia hirta Ruellia humilis Sanicula canadensis Schizachyrium scoparium Scleria pauciflora Smilax glauca Solidago nemoralis Solidago petiolaris Sorghastrum nutans Stylosanthes biflora Verbesina helianthoides Viola triloba Woodsia obtusa 2.6 7.2 1.8 2.4 0.8 2.6 3.6 2.4 0.8 3.4 0.8 1.8 1.8 1.8 0.8 0.8 236 POUNDS HOLLOW SANDSTONE GLADE Woody Seedlings: Amelanchier arborea Carya glabra Carya ovalis Carya ovata Carya texana Celtis tenuifolia Comusflarida Diaspyros virginiana Fagus grandifolia caroliniana Fraxinus americana Juglans nigra Juniperus virginiana Liriodendron tulipifera Pinus echinata Prunus serotina Quercus alba Quercus coccinea ELM 4.2 6.9 1.0 1.0 0 1.6 1.0 0 3.6 1.2 1.0 6.4 0 2.1 O 3.3 2.2 AU.) N O ooooooooowocooonOg—oompoo ox oooo-ooo .0 \o Nowoegro b A \O—‘W 23.4 6.4 . . .A_ - .1). ' Quercus imbricaria Quercus marilandica Quercus rubra Quercus stellata Quercus velutina Rhamnus caroliniana Rhus copallina Sassafras albidum Ulmus alata Vaccinium arboreum Shrub/Saplings: Amelanchier arborea Diaspyros virginiana Juniperus virginiana Pinus echinata Quercus alba Quercus coccinea Quercus marilandica Quercus stellata Quercus velutina Ulmus alata Vaccinium arboreum Trees: Carya glabra Diaspyros virginiana Juglans nigra Juniperus virginiana Quercus alba Quercus coccinea Quercus marilandica Quercus stellata Ulmus alata Herbs: Antennaria plantaginifalia Aristida sp. Asplenium platyneuron Aureolariaflava Carex bushii Carex sp. (Montanae) Cheilanthes lanosa Crotonapsis elliptica C unila origanoides Danthonia spicata 237 1.1 5.9 1.1 19.9 5.4 5.8 6.3 12.6 CD 1.0 O '11 u—- t—- \l \l o—ooowo-— k: NNO HO.) NW 0 ~o Ot—OOQOOO 9° 9° 1)) b») 13.3 N51 0 \lm \ooomoo— '... .O‘ A ON '00 -1 ('1: OWOOOOO 10.0 10.0 ~11 T‘ (I) ~ >0 \o 238 Dichanthelium depauperatum 1.8 0 2.5 0 0 Dichanthelium dichotomum 0 1.7 0 0 4.1 Dichanthelium laxiflorum 5 .4 0 8 .6 10.0 15 .9 Dichanthelium linearifolium 5. l 0 2.5 O 0 Dichanthelium villasissimum 0 1.7 6.1 0 4.1 Diodia teres 0 0 2.5 0 0 Dodecatheon meadia 5.1 12.3 0 0 0 Elymus virginicus 1.8 0 0 0 0 Hedyotis purpurea 0 1.7 0 0 0 Helianthus divaricatus 4.1 17.8 0 0 0 Hypericum gentianoides 0 0 2 5 0 0 Krigia biflora 1 8 0 O 0 0 Krigia dandelion 7 2 5.2 5 0 0 12.2 Krigia sp. 0 1.7 O 0 0 Lespedeza repens 8 9 1.7 0 0 0 Luzula multiflara 0 0 0 30.0 8.1 Manfreda virginica 0 3.4 2.5 0 0 Opuntia humifitsa 0 0 0 0 1 1.8 Oxypolis rigidior ambigua 0 1.7 0 0 O Parthenocissus quinquefolia 5.1 8.9 0 0 0 Phlox pilosa 0 1.7 0 0 0 Palygonatum biflorum 0 0 0 0 l 1.8 Ruellia humilis 1.8 1.7 6.1 0 0 Schizachyrium scoparium 5.4 3.4 5.0 0 0 Smilax bona-nox 0 0 18.5 10.0 0 Smilax glauca 0 0 2.5 0 0 Solidago sp. 1.8 0 0 0 O Stylasanthes biflora 1.8 0 0 0 O T oxicadendran radicans 16.6 10.7 0 0 0 T riodanis perfoliata 0 1.7 0 0 4.1 Vulpia octaflora 0 1.7 0 0 0 Woodsia obtusa 0 1.7 0 O 0 ROUND BLUFF SANDSTONE GLADE Woody Seedlings: FI_—N M Q13 Acer rubrum 2.0 5.2 0 Amelanchier arborea 4.7 4.6 4.9 Carya glabra 4.1 4.9 0 Carya ovata 9.3 10.7 0 Carya texana 0.5 0 0 Celtis tenuifolia 8.7 7.5 0 Cercis canadensis 0.4 0 0 Cornusflaria’a 0.9 0 0 Crataegus sp. 0.4 0 0 Diaspyros virginiana 5.5 1.5 0 Fagus grandifolia caroliniana F raxinus americana Juniperus virginiana Liquidambar styraciflua Liriodendron tulipifera Morus rubra 0sttya virginiana Prunus serotina Quercus alba Quercus marilandica Quercus rubra Quercus stellata Quercus velutina Rhus aromatica Sassafras albidum Symphoricarpas orbiculatus Ulmus alata Vaccinium arboreum Shrub/ Saplings: Carya glabra Carya ovata Juniperus virginiana Ulmus alata Trees: Carya glabra Fraxinus americana Juniperus virginiana Ulmus alata Herbs: Acalypha gracilens Agrostis sp. Aristida sp. Bidens bipinnata Blephilia sp. Bromus pubescens Carex blanda Carex cephalaphora Carex hirsutella Carex muhlenbergii Carex retroflexa Carex sp. Carex umbellata Cheilanthes lanosa 0.5 7.2 11.0 0.4 1.8 0.6 0.4 1.8 1.5 0.4 0.5 8.0 3.1 8.8 0.9 0.5 13.9 1.8 M 50.0 11.3 16.3 22.5 239 5.1 8.6 0.9 1.7 1.8 0.9 8.0 1.1 16.1 3.7 16.6 1.0 TS-N 73.3 26.7 TS-N 19.8 31.3 48.9 r—paoooeoooo o Coo oo oSoo—oooooooopoo io in 12.8 50.2 0 '1: 240 Crotonapsis elliptica 0 8.7 12.1 Danthonia spicata 12.0 8.8 4.7 Dichanthelium acuminatum 0 1.0 9.3 Dichanthelium linearifolium 0.7 2.9 0 Dichanthelium polyanthes 0.7 0 0 Dichanthelium villasissimum 7.5 3.8 0 Diodia teres 0 5.7 16.7 Echinacea pallida 0.7 0 0 Elymus virginicus 3.3 0 O Erigeron strigosus 0 1.0 O Galium circaezans 1.3 0 0 Galium pilosum 0.7 1.0 0 Helianthus divaricatus 2.0 3.9 0 Hypericum gentianoia'es 0 2.9 12.4 Lespedeza procumbens 1.4 0 0 Lespedeza repens 0.7 0 0 Lonicera japonica 4.1 0 0 Manfreda virginica 0 1.9 O Muhlenbergia sobolifera 0.7 0 0 Opuntia humifitsa 0 2.9 2.8 Parietaria pensylvanica 0.7 0 0 Paronychiafastigiata 0 1.0 0 Parthenocissus quinquefolia 9.9 2.1 0 Palygonum cristatum 3.4 0 0 Rosa carolina 1.4 0 0 Rubus enslenii 2.7 0 0 Sanicula canadensis 8.9 2.9 0 Schizachyrium scoparium 0.7 17.9 23 .3 Sedum pulchellum O 1.0 O Smilax bana-nox 1.4 0 0 Smilax glauca O 1.0 0 Solidago caesia 1.4 1.9 0 Solidago nemoralis 0 1.9 O Solidago ulmifolia 1.4 0 O Toxicodendron radicans 4.1 3.8 0 Vitis aestivalis 2.0 0 0 Woodsia obtusa 3.8 1.9 0 WILDCAT BLUFF LIMESTONE GLADE Woody Seedlings: m M QB IS_-_§ F_I-§ Acer negundo 1.9 0.8 0.1 3.5 2.7 Acer rubrum 0 0.8 0.1 0.9 0 Acer saccharum 8.9 7.2 2.2 0.9 0.8 Amelanchier arborea 2. 1 0 0.4 0 0 Aralia spinasa 2.4 3 .4 1.6 0 0 Asimina triloba Carya cordiformis Carya glabra Carya ovalis Carya ovata Carya texana Cassia marilandica Ceanothus americanus Celtis tenuifolia Cercis canadensis Carnusflorida Crataegus sp. Diaspyros virginiana Elaeagnus umbellata Euonymus atropurpurea Fraxinus americana Gleditsia triacanthos Ilex decidua Juglans nigra Juniperus virginiana Liquidambar slyraciflua Lonicera sp. (shrub) Malus ioensis Morus rubra Ostrya virginiana Prunus americana Prunus serotina Quercus alba Quercus prinaides acuminata Quercus rubra Quercus shumardii Quercus stellata Quercus velutina Rhus aromatica Rhus glabra Sassafras albidum Ulmus alata Ulmus rubra Viburnum prunifalium Shrub/Saplings: Acer saccharum Amelanchier arborea Carya ovata Celtis tenuifolia Crataegus sp. Ostrya virginiana Quercus prinaides acuminata oooooooox N L» 19.3 0.9 2.4 1.9 0.9 0.9 2.3 2.8 1.9 2.4 17.6 £121. 21.6 10.8 10.8 46.0 241 11.1 TS-N 25.0 000° 75.0 0 5.6 11.2 Quercus stellata Ulmus rubra Trees: Acer saccharum Carya glabra Carya ovata Carya texana Cornusflorida Crataegus sp. Fraxinus americana Ostrya virginiana Quercus alba Quercus prinaides acuminata Quercus rubra Quercus shumardii Quercus stellata Quercus velutina Sassafras albidum Ulmus rubra Herbs: Acalypha gracilens Agrostis perennans Ambrosia artemisifolia Amphicarpa bracteata Andropogan gerardii Anemone virginiana Apocynum cannabinum Aristolochia serpentaria Arundinaria gigantea Asclepias tuberosa Asclepias verticillata Asclepias viridis Aster laevis Aster patens Bignania capriolata Botrychium virginianum Bouteloua curtipendula Brickellia eupatorioides Bromus pubescens Cacalia atriplicifolia Campanula americana C arex digitalis Carex grisea Carex umbellata Cassia fasciculata E '2 ox oopoooooopg—ooooooo—y O\ O\ Nooopwoo~r~ocooopoooopoow ax axes a» O 242 LAN (.1) Z LII ONONN LI! MW ow \) 6.3 19.0 Q13 10.4 11.7 5.2 3.8 5.2 20.9 2.6 16.9 15.5 2.6 2.6 2.6 Q 1.0 0.1 0.1 0.2 0.7 0.1 3.6 0.3 0.4 0.7 0.3 0.1 0.1 0.5 1.8 0.1 0.1 2.1 0.9 0.2 0.7 0.9 25.0 i-l S” m GOOD—CO 90 DJ Or—u—Iv—IO .°°P°.°° WWW NOOOO -1 S” m —- 00 U’I .— reg—coor—ooooxoooooosuocoooop LII LII 10.1 "1'1 \07‘ m 00 \O pooppoooooooeoooeooo \oxo Or—t—O—O 0000000 \0 1': C hamaesyce maculata Clitoria mariana Cunila origanoides Danthonia spicata Desmodium canescens Desmodium paniculatum Desmodium sp. Dichanthelium boscii Dichanthelium linearifolium Dioscorea quaternata Echinacea pallida Elymus hystrix Elymus virginicus Erechtites hieracifolia Eupatorium altissimum Eupatorium rugosum Euphorbia corollata Galactia regularis Galium circaezans Galium pilosum Helianthus divaricatus Helianthus helianthoides Helianthus microcephalus Hybanthus concolor Hypericum denticulatum Ipomoea pandurata Lespedeza procumbens Lespedeza repens Lespedeza sp. Lespedeza violacea Lespedeza virginica Liatris scabra Lysimachia lanceolata Manfreda virginica Monarda bradburiana Monardafistulosa Muhlenbergia sobolifiara Onosmodium hispidissimum Oxalis stricta Parthenocissus quinquefolia Passiflora lutea Pellaea atropurpurea Phryma leptostachya Physalis virginiana Physostegia virginiana Polyganum cristatum Ranunculus sp. Ratibida pinnata Rubusflagellaris b) LII A \O A N opoowoeooowoeooo~ooo ox over—N \O AON A O’IO’I O\ ooooooeooxoxooguog—ooooooooo A 243 \I ogsooopoOOg—oooysooooooo xo CAN \OLII (Dr—- N 00000 1.2 1.2 2.5 5.0 3.5 2.5 12.6 O OOOOOOOO 0.1 0.2 0.3 0.3 0.3 0.1 0.1 1.9 0.3 0.7 0.1 0.1 0.3 2.4 0.4 2.6 2.9 1.9 0.3 7.3 3.1 1.0 0.9 0.1 0.3 0.1 0.7 1.1 0.6 0.3 1.6 0.2 0.2 0.9 1.3 0.8 0.4 0.2 4.7 0.3 0.1 0.4 1.2 0.7 0.4 1.4 0.1 b.) oooooeowooopooo u: 00 HA UIOI 00 '— OON w o.— LII LII Og—oooofuooswopooooo oxu CI 00 LA) o3—oooooooopaoooooor-g—oMoooooeowooooooo 00 0000 O \O ooeooopoo'xoivo. oo 00 \000 Ruellia caroliniensis Ruellia humilis Sanicula canadensis Schizachyrium scoparium Silphium terebinthinaceum Smilacina racemosa Smilax bona-nax Smilax herbacea Solidago caesia Solidago petiolaris Solidago sp. Solidago ulmifolia Sorghastrum nutans Spigelia marilandica T aenidia integerrima T haspium trifoliatum T oxicodendron radicans Verbesina virginica Vernonia gigantea Viola sororia Viola sp. Viola triloba Vitis aestivalis \O A opoooeoooooooooooowoo 00 N.N UIOI 244 \l ooosogng—ooooowoo LII ON MU) \JN oooiv. LII —-—- GIN 0.3 1.2 0.5 1.1 11.4 1.2 0.9 4.9 0.8 0.2 0.6 0.5 5.7 6.1 0.3 0.3 0.2 1.6 3.1 7.1 3.6 1.8 1.8 1.8 3.6 1.9 0.9 11.3 1.8 1.9 2.8 0 LITERATURE CITED LITERATURE CITED Abbadie, L., A. Mariotti, and J. Menaut. 1992. Independence of savanna grasses from soil organic matter for their nitrogen supply. Ecology 73(2):608-613. Aber, JD. and J.M. Melilo. 1991. Terrestrial ecosystems. Saunders College Publishing, Orlando, Florida. Abrams, MD. 1988. Effect of burning regime on buried seed banks and canopy coverage in a Kansas tallgrass prairie. Southwestern Naturalist 33:65-70. Abrams, MD. and LC. Hulbert. 1987. Effect of topographic position and fire on species composition in a tallgrass prairie in northeast Kansas. American Midland Naturalist ll7(2):442-445. Abrams, M.D., AK. Knapp, and LC. Hulbert. 1986. 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