Identifying Reference Conditions for Riparian Areas of

T · H · E OHIO srA1E October 2003 UNIVERSITY Special Circular 192 OARilr Ohio Agricultural Research and Development Center T · H · E OHIO SfA1E UNIVERSITY OARilr

Steven A. Slack Director

Ohio Agricultural Research and Development Center 1680 Madison Avenue Wooster, Ohio 44691-4096 330-263-3700 Identifying Reference Conditions for Riparian Areas of Ohio

P. Charles Goebel School of Natural Resources Ohio Agricultural Research and Development Center

David M. Hix School of Natural Resources The Ohio State University

Marie E. Semko-Duncan School of Natural Resources Ohio Agricultural Research and Development Center

T · H · E OHIO SfA1E October 2003 UNIVERSITY Special Circular 192 OARil[ Ohio Agricultural Research and Development Center Salaries and research support were provided by state and federal funds appropriated to the Ohio Agricultural Research and Development Center and Ohio State University Extension of The Ohio State University's College of Food, Agricultural, and Environmental Sciences.

The use of trade, firm, or corporation names in this publication is for the information and con­ venience of the reader. Such use does not constitute an official endorsement or approval by the Department of Agriculture, the Agricultural Research Service, The Ohio State Uni­ versity, or the Ohio Agricultural Research and Development Center of any product or service to the exclusion of others that may be suitable. Abstract

Using pre-European settlement vegetation broad forest types that once occurred across surveys and studies of the few remaining the state, they lack the site-specific informa­ old-growth remnant forests, we: tion on forest stand structure and vegetation­ environment relationships needed to • Determined the utility of using these adequately predict reference vegetative different sources of information to states. develop reference conditions for riparian forests of Ohio. The studies of the few remaining old-growth remnants do elucidate many of the finer­ • Identified gaps in our understanding of the ecology of Ohio's riparian forests scale vegetation-environment relationships, that need to be addressed to help providing considerable data on the overstory develop reference conditions for these composition and, in some cases, ground­ important landscape components. flora composition of these reference riparian systems. More information, however, is Based on an extensive review of the avail­ needed on the structure of these riparian able literature, we surmised that while forests in order to develop suites of ecologi­ pre-European settlement maps and surveys cally based reference conditions for Ohio's provide generalized information on the riparian forests.

Introduction

For a resource manager interested in tion (Wyant et al., 1995). This is particularly restoring plant communities in disturbed true for riparian areas, where variation in ecosystems, the identification of reference physiography, disturbance regimes, and soil vegetation conditions is an important step characteristics can result in a diverse array in the process of forest ecosystem restora­ of plant communities at a variety of spatial tion (Aronson et al., 1995; Pickett and Par­ scales (e.g., Gregory et al., 1991; Bendix and ker, 1994). Identifying reference conditions Hupp, 2000). (e.g., composition, structure, and function of woody and herbaceous species) for a specific A common source of information on histori­ ecosystem, however, is often a very conten­ cal plant communities that has often been tious issue. used to identify reference conditions for forest ecosystem restoration is the original Successional pathways and plant commu­ surveyor notes of witness trees, such as those nity composition and structure can be highly developed by the General Land Office (GLO) variable, making it difficult to use historical in the early 1800s. From these surveyors' plant communities as templates for restora notes, the pre-European settlement distribu

1 tion of forest ecosystem types has been Other less utilized sources of informa- developed, and relatively detailed infor­ tion on reference vegetation conditions mation on the historical composition and for forest ecosystem restoration are the structure of these forests has been deter­ remaining relatively undisturbed old­ mined. growth forest ecosystems. Although less than 1% of the pre-European settlement For example, in Ohio, Gordon (1966) forests remain in Ohio (Davis, 1996), there developed a pre-European settlement map are examples of minimally disturbed of forest types, and many resource mana­ old-growth(> 150 years old) and mature gers and conservation organizations use second-growth (120 to 150 years old) forest this classification to guide their selection of ecosystems in Ohio, many of which are reference vegetation conditions (Figure 1). located on public lands (e.g., Goebel and Hix, 1996).

s

CJ Ohio Pre-European Settlement Vegetation - Beech Forests CJ Mixed Oak Forests D Oak-Sugar Maple Forests - Elm-Ash Swamp Forests Mixed Mesophytic Forests CJ Prairie Grasslands CJ Oak Savannas CJ Freshwater Marshes and Fens - Sphagnum Peat Bogs CJ Bottomland Hardwood Forests 100 O 100 Kilometers D Mixed Mesophytic Forests (Hemlock-Beech-Chestnut-Oak) D White Pine-Red Maple Swamps ~~----~~~~ D Beaches

Figure I. Presett/ement forest types of Ohio (based on Gordon, 1966).

2 Although they often represent a small and the few remaining old-growth forest area, these forests provide an opportunity ecosystems in each of Ohio's ecoregions to study vegetation-environment relation­ (McNab and Avers 1994; Figure 2) to: ships and develop predictive models of forest ecosystem development that can be • Determine the utility of using each used to develop templates for the restora­ of these sources to develop reference tion of forest composition and structure. conditions for riparian forests of As many of these old-growth forest eco­ small and large streams and rivers in systems also have small streams and rivers each of Ohio's ecoregions. flowing through them, they may provide • Identify gaps in our understand- useful information to develop reference ing of the ecology of Ohio's riparian conditions for riparian forests. forests that need to be addressed to The objective of our report is to review the help develop reference conditions for relevant literature related to the pre-Euro­ these important landscape compo­ pean settlement forest-type surveys nents.

Riparian Areas and Reference Conditions

Riparian areas are functional ecotones or (Ilhardt et al., 2000; O'Laughlin and Belt, transitional areas located between ter­ 1995). Unfortunately, we have a poor restrial and riverine ecosystems. Despite understanding of the patterns of variation their limited areal extent, riparian areas in riparian areas within and among water­ promote many ecosystem functions vital sheds, or the specific landscape features to the health and productivity of forested that control riparian vegetation develop­ watersheds. Not only do riparian areas ment. regulate the flow of water, sediments, and nutrients across system boundaries, Increasingly, riparian areas across Ohio they also contribute organic matter to the are experiencing pressure from a variety aquatic system, increase bank stability, of sources, including developers, farmers, reduce erosion, and provide key wildlife and recreationists. Detailed assessments of habitat (Gregory et al., 1991; llhardt et al., relatively undisturbed riparian areas are 2000). critical elements needed for a variety of ecological purposes, especially for provid­ Additionally, because of their functional ing a benchmark of reference ecological importance, riparian areas serve important conditions necessary for evaluating forest roles in mitigating many of the negative ecosystem restoration programs. impacts of land use on aquatic systems as well as protecting species diversity, pro­ Voluntary best management practices viding potential dispersal corridors for (BMPs) promoted by various state agen­ wildlife, and mitigating flood waters cies (e.g., the Ohio Department of Natu

3 •Drew r-----..1 Woods

•Hueston Woods

Ecoregions D Central Tiii Plains, Beech-Maple Section D Erie and Ontario Lake Plain Section - Interior Low Plateau, Bluegrass Section - South Central Great Lakes Section 100 0 100 Kilometers • Southern Unglaciated Section ~~--~~~~ D Western Glaciated Allegheny Plateau Section Figure 2. Ecoregions of Ohio (McNab and Avers, 1994) and location of remaining old-growth and mature second-growth forests with information on riparian forests published in a refereed journal article. ral Resources, ODNR) may be available lineation of riparian areas is the need for and utilized in riparian areas (Blinn and an explicit quantification of reference, or Kilgore, 2001 ). However, the effective­ benchmark, riparian conditions (Gregory, ness of these practices to help restore or 1999). Although current management sys­ even maintain critical ecosystem linkages tems incorporate the best available science, is untested. These practices are certainly our understanding of specific riparian not site-specific in their application (e.g., areas and the extent of the functions asso­ the width of riparian management zones ciated with these land-water interfaces is is often fixed arbitrarily rather than being often incomplete (Blinn and Kilgore, 2001 ). based on a functional delineation of ripari­ an extent; deMaynadier and Hunter, 1995). Understanding the range of variability in these reference riparian areas is important One important consequence of a shift as they represent the goal of riparian resto­ toward a functional definition and de ration and management efforts (i.e.,

4 the undisturbed riparian area with all areas in North America, are based on ex­ ecosystem linkages intact) and provide a tending our understanding of forest stand metric or standard with which to compare dynamics from upland forests or other current and future riparian restoration and riparian settings in different regions (Blinn management programs. and Kilgore, 2001; Gregory, 1999). The simple fact that there is a diverse array of While we know very little about the com­ environmental gradients distributed across position and structure of the riparian riparian areas (e.g., microclimate, flood­ forests in the state, many of the current ing) not found in upland settings, or other restoration and management systems used riparian systems, suggests that these com­ in riparian areas in Ohio, as well as other parisons may not be valid (Gregory, 1999).

Methods

We conducted a review of the refereed available information on the major spe­ and non-refereed literature associated cies comprising the overstory (stems> 10 with Ohio's forests, searching a variety cm diameter at breast height [dbh]; 1.4 of sources, including the Internet, on-line m), understory (stems < 10 cm dbh but > databases (e.g., AGRICOLA), and the Ohio­ 2.5 cm dbh), and the ground-flora(< 1 m LINK library catalog using the following tall) by ecoregion as defined by McNab keywords: riparian forest, floodplain, and Avers (1994). We also examined these bottomland, streams, rivers, streamside studies for any useful information that forest, streambank, and old-growth forest. could help identify reference conditions We also reviewed more than 20 M.S. theses for riparian forests. and Ph.D. dissertations from Ohio colleges and universities, as well as numerous It is important to note that a variety of Ohio Biological Survey and Ohio Agricul­ terms that may relate to riparian forests tural Experiment Station (now the Ohio were encountered in these different Agricultural Research and Development sources, such as floodplain forests, swamp Center) forest inventory reports dating forests, bottomland forests, and river or back as far as the early 1920s that pertain stream wetland forests. Consequently, for to the pre-European settlement vegetation the purposes of our review, we consider of many of Ohio's counties. riparian forests to include any forest located near any body of water, such as a Pre-European settlement vegetation de­ lake, stream, or river. Where not defined in scriptions at the state- and county-wide the paper, thesis, or report, we considered level were compiled to examine the useful­ floodplain and bottomland forests to ness of this information to develop lists of be associated with larger river systems reference plant communities for riparian that have extensive floodplains (e.g., forests by ecoregion. We also examined Cuyahoga, Scioto, Little Miami Rivers), more recent descriptive studies of exist­ and river and stream wetland forests to be ing old-growth forests and less disturbed associated with smaller rivers and streams areas across the state and summarized the lacking extensive floodplain development.

5 Categorizing the swamp forests was more whether these forests were associated with problematic; as a result we examined the a larger or smaller stream system. Finally, local physiographic conditions using a plant nomenclature follows Kartez (1994); variety of sources (e.g., local soil survey see the Appendix for a list of Latin and maps, topographic maps) to determine common species names.

Summary of Different Survey Types

The first descriptions of the major pre­ cottonwood dominated the floodplains of European settlement forest types for Ohio larger streams and rivers, while smaller were developed by Sears (1925) and Samp­ streams flowing through narrow valleys son (1927), who transcribed the species of throughout the state were dominated by bearing or witness trees from the original hemlock, American beech, and maple. land surveys conducted in the late 1700s and early 1800s. Sears (1925) classified The most detailed description of the pre­ each township in the state as fundamen­ European forests of Ohio (Figure 1) was tally "beech," "oak," or "ash" or various developed by Gordon (1966, 1969), who combinations of these types and classified synthesized the work of Sears (1925), treeless areas into "prairie" types (Sears, Transeau and Sampson (1938), and Chap­ 1926). man (1944), as well as many county-level forest-type classifications published as Transeau and Sampson (1938) also used M.S. theses, Ohio Biological Survey (OBS) these original land surveys in conjunction reports, and state forestry reports pub­ with field studies of remaining undis­ lished by the Works Progress Administra­ turbed forests in an effort to classify major tion and the Ohio Agricultural Experiment forest types, including swamp forests, and Station (OAES). For example, Gordon develop strategies to better utilize Ohio's (1969) described major plant associations forests. Sampson (1927) mapped major for different floodplains and swamp for­ plant associations and suggested that the ests; however, he realized that these forest riparian forests of the state were domi­ types associated with streams and rivers nated by willow, alder, river birch, and were the least understood and most vari­ maple-cottonwood-sycamore forest asso­ able in terms of species composition in the ciations. state. A similar pre-European forest-type classi­ Furthermore, in one of the more detailed fication as Transeau and Sampson (1938) early botanical studies of Ohio's regional was developed by Chapman (1944). How­ flora, Griggs (1914) stated no problem ever, Chapman was the first to provide a was so difficult as the reconstruction of reconstructed picture of the pre-European the vegetation of the bottomlands along settlement riparian forests, as he docu­ the large streams and in his words "there mented that elm, ash, soft maple (e.g., red is not a vestige left to suggest the original maple), sycamore, burr oak, pin oak, and condition of the Hocking bottom between

6 Lancaster and Logan except the swamps Conversely, on silt-laden deposits of and a few large trees standing in the fields floodplains along major river systems of into which it has been converted." the ecoregion (e.g., Cuyahoga and Scioto Rivers and Killbuck Creek), boxelder The amount and details of specific infor­ (Acer negundo L. ), silver maple (Acer sac­ mation on the vegetation-environment charinum), Ohio buckeye (Aesculus glabra relationships of relatively undisturbed Willd. ), green ash (Fraxinus pennsylvanica vegetation of Ohio's riparian forests are Marsh.), black walnut (Juglans nigra L. ), varied and depend largely on the spe- American sycamore (Platanus occidentalis cific ecoregion. Overall, there tends to L. ), eastern cottonwood (Populus deltoi- be specific information on the overstory des Bartr. ex Marsh.), and American elm composition, but very little data on the (Ulmus americana) dominate the overstory. understory and ground-flora plant com­ munities. For example, many of the The understory of these floodplains along county-level M.S. theses and OBS and larger streams and rivers is similar to the OAES reports tend to be distributed across swamp forests described previously, with each ecoregion of the state and describe the addition of several species including the composition of local riparian forests silver false spleenwort (Deparia acrosti­ (e.g., Diller, 1932; Jones, 1936; Norris, choides (Sw.) M. Kato), eastern bottlebrush 1948). grass (Elymus hystrix L. var. hystrix), white­ grass (Leersia virginica Willd. ), ostrich Furthermore, Andreas (1989) suggests fern (Matteuccia struthiopteris (L.) Todaro), that riparian plant communities of the Virginia creeper (Parthenocissus quinquefo­ glaciated Allegheny Plateau ecoregion lia (L.) Planch. ), marshpepper knotweed are related to predictable assemblages of (Polygonum hydropiper L. ), jumpseed (P plant communities that are organized on virginianum L. ), eastern poison ivy (Toxico­ the basis of geology, topography, mois­ dendron radicans (L.) Kuntze), cutleaf cone­ ture availability, and aspect. For instance, flower (Rudbeckia laciniata L. ), wingstem swamp forests dominated by overstories (Verbesina alternifolia (L.) Britt. ex Kearney), of red maple (Acer rubrum L.), silver maple and striped cream violet (Viola striata Ait.) (A. saccharinum L. ), white ash (Fraxinus americana L.), and American elm (Ulmus However, these sources of reference vege­ americana L.) and understories of common tation conditions often provide generali­ elderberry (Sambucus canadensis (L.) R. Bol­ zations of the different overstory and li), beggarticks (Bidens spp.), sedge (Carex understory species, but not the site-specific spp.), sweet woodreed (Cinna arundinacea information needed to develop detailed L. ), Canadian woodnettle (Laportea ca­ restoration templates for riparian areas nadensis (L.) Weddell), rice cutgrass (Leersia across each ecoregion of the state. oryzoides (L.) Sw.), sensitive fern (Onoclea sensibilis L. ), and reed canarygrass (Phalaris While many of the refereed journal articles arundinacea L.) occur across the ecoregion that examine the ecology of individual on flat, poorly-drained floodplains associ­ old-growth forests do provide site-specific ated with small streams and rivers. information on the vegetation-environ­ ment relationships of riparian forests, they often focus only on overstory composition and rarely on the structure of these

7 riparian forests (Table 1). Additionally, the across the ecoregions of Ohio (Table 1). In information provided by these surveys the sections that follow, we document the and studies is often disparate, resulting in available information of reference riparian considerable gaps in our understanding of conditions for each ecoregion from these the factors that regulate the composition studies of specific old-growth and mature and structure of riparian forests forests.

Table 1. Compositional and Structural Information on Overstory and Ground-Flora Strata of Riparian Forests Provided by Published Studies of Old-Growth Forest Eco- systems.

Composition Structure

Over- Under- Ground Over- Under- Ground Area Citation Soils story story Flora story story Flora

Western Glaciated Allegheny Plateau Johnson Woods Braun, 1950 x x Goebel et al., 2003 x x x

Southern Unglaciated Allegheny Plateau McCarthy et al,. 1987 x x x x x Lake Katherine Runkle & Whitney, 1987 x x x x x Morton's Woods Braun, 1950 x x

Interior Low Plateau Bluegrass Fort Hill State Braun, 1969 x x x x Memorial

Central Till Plains, Beech-Maple Cabin Run Cobbe, 1943 x x Cedar Cliffs Irwin, 1929 x x x Crall Woods Aughanbaugh, 1964 x x x x Drew Woods Boerner & Kooser, 1991 x x x x Emery Woods Swanson & Vankat, 2000 x x x x x Glen Helen Anliot 1973 x x x x Clifton Gorge Anliot 1973 x x x x John Bryan State Anliot 1973 x x x x Park Hazelwood Bot. Segelken, 1929 x x x Preserve Hueston Woods Braun, 1950 x Werth et al., 1984 x x x Sears & Carmean Cho & Boerner, 199la,b x x x x Woods

Erie and Ontario Lake Plain Goll Woods Boerner & Cho, 1987 x x x x North Chagrin Res. Williams, 1936 x x

South Central Great Lakes

8 Riparian Reference Conditions by Ecoregion

(1950) documented that along an inter­ Western Glaciated Allegheny mittent stream in this gently dissected Plateau morainal system classified by Gordon (1969) as an elm-ash swamp forest type, the lower, poorly drained flats were domi­ In the Western Glaciated Allegheny Pla­ nated by red maple (Acer rubrum), Ameri­ teau ecoregion located in northeastern and can elm (Ulmus americana), and swamp central Ohio (Figure 2), data are available white oak (Quercus bicolor), while on the from four different sources, two of which drier swells or morainal ridges, white oak examine the vegetation-environment re­ (Quercus alba), American beech (Fagus lationships of riparian forests that were grandifolia Ehrh.), and sugar maple (Acer conducted at the same location (Graeber saccharum Marsh.) dominated the over­ Woods or Johnson Woods, Figure 3). Braun story.

Figure 3. Intermittent stream flowing through Johnson Woods State Nature Preserve with an understory dominated by jewelweed, Canadian wood nettle, and stinging nettle.

9 Although Braun (1950) makes no men­ (Carya cordiformis (Wangenh.) K. Koch), tion of the herbaceous understory, Goebel and yellow buckeye (Aesculus flava Ait. ), et al. (2003) found that specific ground­ and published stand structural informa­ flora species were associated with the tion including the basal area and density floodplains located along this intermittent of overstory and understory woody spe­ channel, and these were related strongly cies, as well as woody species richness to landform and soil characteristics. For (Table 1). example, jewelweed (Impatiens capensis Meerb. ), Canadian woodnettle (Laportea Braun (1950) documented similar over­ canadensis), whitegrass (Leersia virginica), story composition for the former Morton's and stinging nettle (Urtica dioica L.) were Woods in Gallia County (Figure 2), with all associated with floodplain soils that the addition of American beech (Fagus have finer textures, higher organic matter grandifolia) which was a canopy dominant. content, and higher concentrations of total No information on the riparian ground­ N, nitrate-N, phosphorus, potassium, flora composition or structure is avail­ calcium, and magnesium than adjacent able for either of these old-growth forests upland communities (P. C. Goebel, unpub­ (Table 1). lished data). In the western portion of this ecoregion, Braun (1928) characterized many early successional riparian forests of eastern Adams County as being dominated by Southern Unglaciated Allegheny sweetgum (Liquidambar styraciflua L. ), Plateau which is eventually replaced by more shade-tolerant species such as American sycamore (Platanus occidentalis) and other In the Southern Unglaciated Allegheny mixed mesophytic species. Plateau ecoregion (Figure 2), five pub­ Similar results were reported by Runkle lished studies have examined the vege­ and Whitney (1987) at the Lake Katherine tation-environment relationships of State Nature Preserve (Figure 2) where old-growth or relatively undisturbed floodplains and terraces along smaller forest ecosystems. Both McCarthy et al. creeks are dominated by canopies of tulip­ (1987) and Braun (1950) report informa­ tree (Liriodendron tulipifera), yellow birch tion on riparian forests of narrow ravines (Betula alleghaniensis Britt.), river birch along small intermittent streams flowing (B. nigra L. ), and American sycamore through old-growth forest ecosystems. (Platanus occidentalis) and an understory McCarthy et al. (1987) classified the ri­ of American hornbeam (Carpinus carolini­ parian forests of Hawk Woods in Athens ana Walt.), northern spicebush (Lindera County (Figure 2) as a mixed-mesophytic benzoin), umbrella-tree (Magnolia tripetala type, characterized by white oak (Quercus (L.) L.), American hazelnut (Corylus ameri­ alba), northern red oak (Q . rubra L.), sugar cana Walt.), pawpaw (Asimina triloba (L.) maple (Acer saccharum), tuliptree (Liri­ Dunal), and boxelder (Acer negundo). odendron tulipifera L. ), American basswood (Tilia americana L. ), bitternut hickory

10 hemlock (Cicuta maculata L.), and field Interior Low Plateau Bluegrass horsetail (E. arvense L.) Finally, although Bryant (1987) describes California Woods Nature Preserve located Along the Ohio River in southwestern in the Cincinnati Metropolitan area (Figure Ohio in the northern extension of the 2) as a ravine community dominated by Interior Low Plateau Bluegrass ecoregion mixed-mesophytic tree species, no specific (Figure 2), we found only one study that information on the composition or struc­ focused on the vegetation-environment ture is provided. relationships of riparian forests. Braun (1969) characterized the mature riparian forests associated with the rolling low­ lands of wide open ravines and gentle slopes located at the Fort Hill State Memo­ Central Till Plains, Beech-Maple rial (Figure 2). Along the banks of small streams, Braun Twelve studies were found that describe (1969) described a diverse ground-flora the riparian forests of the Central Till (> 40 species), perhaps best character­ Plains, Beech-Maple ecoregion (Figure ized by large patches of bristly buttercup 2), with 10 detailing riparian vegetation­ (Ranunculus hispidus Michx. var. nitidus environment relationships, making it (Chapman) T. Duncan) and a sparse over­ the most intensively studied of any eco­ story of American sycamore (Platanus region in the state (Table 1 ). These studies occidentalis), American elm (Ulmus ameri­ included riparian areas associated with cana), boxelder (Acer negundo), and green intermittent stream channels, riparian ash (Fraxinus pennsylvanica) grading into a areas along small streams and ravines, and mixed-mesophytic forest. riparian areas associated with large rivers, broad flats or floodplains, and bottomland Although not associated with a specific swamps. forest, Braun (1928) documented that ri­ parian forests associated with frequently Crall Woods in north-central Ohio and flooded flats across Highland County are Cabin Run Forest in southwestern Ohio characterized by an overstory of Am,erican are both examples of old-growth forests sycamore (Platanus occidentalis), Ameri­ traversed by intermittent streams (Figure can elm (Ulmus americana), and black 2). However, Aughanbaugh (1964), in his willow (Salix nigra Marsh.) and a diverse study of Crall Woods, does not differen­ ground-flora including cutleaf coneflower tiate the undulating terrain into riparian (Rudbeckia laciniata L. ), crookedstem aster and upland areas. The major tree species (Symphyotrichum prenanthoides (Muhl. ex encountered were sugar maple (Acer sac­ Willd.) Nesom), common sneezeweed charum), American basswood (Tilia ameri­ (Helenium autumnale L. ), wingstem (Ver­ cana), American beech (Fagus grandifolia), besina alternifolia (L.) Britt. ex Kearney), tuliptree (Liriodendron tulipifera), American cup plant (Silphium perfoliatum L.), giant elm (Ulmus americana), northern red oak goldenrod (Solidago gigantea Ait.), fall (Quercus rubra), shagbark hickory (Carya phlox (Phlox paniculata L. ), spotted water ovata (P. Mill.) K. Koch), and hophornbeam

11 (Ostrya virginiana (P. Mill.) K. Koch), with The old-growth forest of Emery Woods in more than 90 herbaceous species recorded Hamilton County (Figure 2) includes an in the wooded area. area where ravine slopes, associated with small and often times sluggish streams, At Cabin Run Forest along intermittent are prominent. At Emery Woods, Swanson streams that dissect the upland areas and Vankat (2000) found that the forests dominated by mixed-mesophytic forests, of these ravine bottoms tend to support Cobbe (1943) found that sugar maple (Acer mixed mesophytic species including sugar saccharum), American beech (Fagus grandi­ maple (Acer saccharum), hickory (Carya folia), tuliptree (Liriodendron tulipifera), spp.), oak (Quercus spp.), American beech American basswood (Tilia americana), (Fagus grandifolia), and tuliptree (Lirioden­ black walnut (Juglans nigra), and white oak dron tulipifera). (Quercus alba) were nearly always present in the riparian area, with their abundance These finding are similar to those reported dependant on local site and microclimatic by Braun (1936) in her descriptive com­ conditions. parison of ravine slope forests and flats within the Illinoian till plains region of Cobbe (1943) also documents that a small southwestern Ohio. Similar relationships stream bordering the forest at Cabin Run were observed along a small stream drain­ supported a streamside community where ing a large ravine and two main branches American sycamore (Platanus occidentalis) with swampy, flat-bottomed floors at the was dominant, along with several other Hazelwood Botanical Preserve (HBP), also species including sugar maple (Acer sac­ in Hamilton County (Figure 2). charum), black walnut (Juglans nigra), tulip­ tree (Liriodendron tulipifera), and pignut A variety of habitats from extremely hickory (Carya glabra). wet swamp communities to mesophytic associations were observed at HBP. Spe­ The herbaceous layer was composed of cifically, Segelken (1929) found that her­ rattlesnake fern (Botrychium virginianum baceous vegetation far outnumbered the (L.) Sw.), brittle bladderfern (Cystopteris woody plants on the wetter sites. Those fragilis (L.) Bernh. ), Canadian wildginger species reported to occur in the wetter (Asarum canadense L.), Virginia spring­ areas include yellow marsh marigold beauty (Claytonia virginica L.), cutleaf (Caltha palustris L.), rice cutgrass (Leersia toothwort (Cardamine concatenata (Michx.) oryzoides), arrowleaf tearthumb (Polygonum Sw. ), white fawnlily (Erythronium albidum sagittatum L. ), jewelweed (Impatiens capen­ Nutt.), fragrant bedstraw (Galium triflorum sis), and broadleaf cattail (Typha latifolia Michx.), jewelweed (Impatiens capensis), L. ). Prairie ironweed (Vernonia fasciculata twinleaf Ueffersonia diphylla (L.) Pers.), Michx.) was common along the stream, great blue lobelia (Lobelia siphilitica L. ), while sweetscented joepyeweed (Eupa­ common yellow oxalis (Oxalis stricta L.), torium purpureum L.) and common elder­ wild blue phlox (Phlox divaricata L. ), may­ berry (Sambucus canadensis L.) were found apple (Podophyllum peltatum L. ), toadshade on the ravine floor in drier places. (Trillium sessile L. ), downy yellow violet (Viola pubescens Ait. ), and striped cream Several black willow (Salix nigra) commu­ violet (V striata Ait). nities were found growing on the ravine floor in both wet and drier sites. Along the

12 sloping margins of the stream bank, Ameri­ These isolated and protected parts are very can elm (Ulmus americana) and American different from the bluffs themselves. sycamore (Platanus occidentalis) were found distributed where hydro-mesophytic condi­ Irwin (1920) reported that the extremely tions prevail. steep bluff was almost bare of vegetation, whereas the ravines supported a meso­ In addition to these species, Segelken phytic habitat where northern red oak (1929) found the following herbaceous (Quercus rubra) and sugar maple (Acer plants on the ravine floor: harvestlice saccharum) were dominant. American elm (Agrimonia parviflora Ait. ), American water­ (Ulmus americana), black walnut (Juglans plantain (Alisma plantago-aquatica L.), nigra), blue ash (Fraxinus quadrangulata Indianhemp (Apocynum cannabinum L.), Michx.), eastern redbud (Cercis canaden­ swamp milkweed (Asclepias incarnata L.), sis L.), and yellow chestnut oak (Quercus shallow sedge (Carex lurida Wahlenb.), muhlenbergii Engelm.) occurred as sap­ Canadian honewort (Cryptotaenia canaden­ lings. sis (L.) DC.), white turtlehead (Chelone glabra L. ), ovate spikerush (Eleocharis The most important species of the diverse ovata (Roth) Roemer & J. A. Schultes), herbaceous layer consisted of muhly grass field horsetail (Equisetum arvense L. ), (Muhlenbergia spp. ), hairy sunflower (Heli­ cream avens (Geum virginianum L.), spot­ anthus hirsutus Raf.), American bittersweet ted St. Johnswort (Hypericum punctatum (Celastrus scandens L.), upland boneset Lam.), beggarslice (Hackelia virginiana (L.) (Eupatorium sessilifolium L.), longsepal I. M. Johnston), great blue lobelia (Lobelia beardtongue (Penstemon calycosus Small), siphilitica L. ), sharpwing monkeyflower and tall rattlesnakeroot (Prenanthes altissi­ (Mimulus alatus Ait. ), ground ivy ( Glechoma ma L.). hederacea L. ), pellitory (Parietaria pennsyl­ Anliot (1973) reported on the vascular vanica Muhl. ex Willd. ), ditch stonecrop flora of Glen Helen, Clifton Gorge, and (Penthorum sedoides L. ), Canadian clear­ in north central weed (Pilea pumila (L.) Gray), halberdleaf Greene County (Figure 2). The topography tearthumb (Polygon um arifolium L. ), dotted of the study area is fairly flat except where smartweed (Polygonum punctatum Ell.), dissected by the and its common selfheal (Prunella vulgaris L. ), tributaries. Anliot (1973) found there was blue skullcap (Scutellaria lateriflora L. ), great variability associated with soil char­ roundleaf goldenrod (Solidago patula Muhl. acteristics and drainage among the flood­ ex Willd. ), white vervain (Verbena urticifo­ plain communities. lia L. ), and summer grape (Vi tis aestivalis Michx.). Willow-eastern cottonwood-American sycamore (Salix spp.-Populus deltoides-Pla­ At Cedar Cliffs Prairie, located on a bluff tanus occidentalis) communities were found of the Little Miami River (Figure 2), a mainly along the larger waterways. Where broad floodplain extends unbroken for better drainage occurred, silver maple miles, with prairie openings that occur in (Acer saccharinum), American elm (Ulmus exposed locations where evaporation and americana), and ash (Fraxinus spp.) were exposure are great. The bluff is indented dominant, while either bur oak (Quercus by ravines and gullies of various sizes. macrocarpa Michx. ), swamp white oak (Q. bicolor Willd.), and shellbark hickory

13 (Carya laciniosa (Michx. f.) G. Don) or Woods lacks the rocky cliffs and hill prai­ northern red oak (Quercus rubra) and ries of the former areas (Werth et al., 1984). American basswood (Tilia americana) forest communities dominated the best drained While the work of Vankat et al. (1975) sites. In addition, bur oak (Quercus macro­ focused on the woody vegetation of the carpa), silver maple (Acer saccharinum), upland portion of Hueston Woods, Werth black walnut (Juglans nigra), butternut et al. (1984) compiled a list of vascular (J. cinerea L. ), Ohio buckeye (Aesculus plant flora collected from areas through­ glabra), and black maple (Acer nigrum out the park and preserve. Included are 40 Michx. f.) also occurred frequently on allu­ streamside species, nine species found in vial deposits along streams. wooded ravines, and 46 from wet woods. Additionally, Braun (1950) lists sugar Anliot (1973) also documented succes­ maple (Acer saccharum) as the dominant sional patterns along the Little Miami canopy species occurring on alluvial ter­ River in old fields abandoned less than 10 races at Hueston Woods with American years. The fields first became reforested elm (Ulmus americana), white oak (Quercus primarily with boxelder (Acer negundo L.), alba), and shagbark hickory (Carya ovata) American sycamore (Platanus occidenta­ present as minor components. lis), and honeylocust (Gleditsia triacanthos L. ), often in pure stands. Before a canopy While the preceding examples are from developed, Canada goldenrod (Solidago southwestern Ohio, the Sears Woods and canadensis L. and S. canadensis var. scabra Carmean Woods complex in the till plains Torr. & Gray), hairy white oldfield aster of north-central Ohio also exhibit good (Symphyotrichum pilosum (Willd.) Nesom.), examples of typical riparian forests of the white panicle aster (Symphyotrichum lan­ Central Till Plains, Beech-Maple ecoregion ceolatum var. lanceolatum (Willd.) Nesom. ), (Figure 2). In the two studies by Cho and wingstem (Verbesina alternifolia (L.) Britt. Boerner (199la,b), Sears Woods is de­ ex Kearney), giant ironweed (Vernonia gi­ scribed as comprising an area of upland gantea (Walt.) Trel.), Jerusalem artichoke moraine dissected by several major stream (Helianthus tuberosus L. ), Canada thistle drainages and a large, flat floodplain ter­ (Cirsium arvense (L.) Scop.), hairy pagoda­ race along the . Both areas plant (Blephilia hirsuta (Pursh) Benth.), have moderately well to well-drained creeping jenny (Lysimachia nummularia soils. In contrast, the topography of Car­ L. ), and stinging nettle (Urtica dioica) were mean Woods is level with very poorly common. drained soils.

Hueston Woods (Figure 2), a nearby state The old-growth bottomland areas of the park and nature preserve of comparable Sears-Carmean Woods complex are domi­ size containing old-growth forest rem­ nated by silver maple (Acer saccharinum), nants, lies near the southern boundary American basswood (Tilia americana), of the Beech-Maple forest region (Braun American elm (Ulmus americana), and 1950) near the transition to western and sugar maple (Acer saccharum), and the total mixed mesophytic forests (Vankat et al., tree species composition corresponds well 1975). However, unlike the Glen Helen­ to the early descriptions of the red oak­ Clifton Gorge-John Bryan complex, the basswood phase of the elm-ash swamp mildly dissected topography of Hueston forest as described by Sampson (1930).

14 Figure 4. Maple and elm riparian forest along the Sandusky River flowing through the Sears-Carmean State Nature Preseve.

The upland forest component, in which can be found at Drew Woods, an isolated white oak (Quercus alba), northern red oak old-growth forest remnant located on the (Q. rubra), and white ash (Fraxinus ameri­ poorly drained till plains of western Ohio cana) were present in the canopy with co­ (Figure 2). Boerner and Kooser (1991) re­ dominants sugar maple (Acer saccharum) ported that "early European settlers found and American beech (Fagus grandifolia), this area covered by a complex of swamp also corresponds well to Gordon's (1969) forest and bottomland forest broken only description of beech-sugar maple forest. by occasional stream corridors." No ground-flora information was reported Species more abundant on lower, wetter in either study of Sears-Carmean Woods sites included sugar maple (Acer saccha­ by Cho and Boerner (1991a,b). rum), swamp white oak (Quercus bicolor), northern red oak (Q . rubra), bur oak (Q. Examples of old-growth bottomland macrocarpa), and shellbark hickory (Carya swamp forest communities in the Central laciniosa). In contrast, sugar maple (Acer Till Plains, Beech-Maple ecoregion saccharum), white oak (Quercus alba), and

15 shagbark hickory (Carya ovata) occupied (Ulmus americana), tuliptree (Liriodendron the higher, drier sites. The community tulipifera), American basswood (Tilia ameri­ structure of the bottomland areas cor­ cana), slippery elm (U. rubra Muhl.), north­ responds well to the description of oak­ ern red oak (Quercus rubra), flowering maple swamp given by Anderson (1982). dogwood (Cornus fiorida L.), black cherry (Prunus serotina Ehrh. ), black walnut (Jug­ lans nigra), bitternut hickory (Carya cordi­ formis), and butternut (J. cinerea). Although Erie and Ontario Lake Plain Williams (1936) lists all herbaceous plants found in the study area, he did not catego­ rize them by habitat type. The Erie and Ontario Lake Plain ecoregion Schlesinger (1971) also found similar (Figure 2) extends from the shores of upland swamp forest communities along Lake Erie in northeastern Ohio where the the sides and bottom of the large ravine topography includes level highlands cut cutting through his study area in eastern by ravines and develops into broader, flat­ Cuyahoga County, but specific informa­ ter areas in the western portions, especial­ tion on the vegetation from this area was ly along the Maumee Drainage, commonly not sampled or reported. described as the Black Swamp. Data on riparian conditions for this ecoregion are Vegetation-environment relationships of available from six sources; however, only Goll Woods, the last known uncut old­ three provide information on riparian growth forest remnant of the Black Swamp forest vegetation-environment relation­ forest of northwestern Ohio, were studied ships. by Boerner and Cho (1987). They found that the distribution of tree species was Williams (1936) described the North regulated by different drainage conditions. Chagrin Reservation of the Cleveland Species occupying the poorly drained Metropolitan Park System in northeast­ flats were silver maple (Acer saccharinum), ern Ohio (Figure 2) as an area of high American elm (Ulmus americana), bur oak bluffs along the valley (Quercus macrocarpa), and ash (Fraxinus characteristically cut by short gullies and spp. ). As drainage improves, these spe­ deep ravines. Williams also noted that no cies were replaced by American basswood perennial streams run through the study (Tilia americana), American hornbeam (Car­ area, but an excess of moisture is always pinus caroliniana), northern red oak (Quer­ present in the deeper ravines. The ravines cus rubra), and red maple (Acer rubrum) might be considered the beginning of a on transitional sites adjacent to the drier transitional phase toward swamp forest or beach ridges. bottomland conditions where trees often represent bottomland species. Sampson (1930), Braun (1950), and Gordon (1969) discerned the major influences of Some of the tree species encountered in the elevation, soil drainage, and aeration on ravines included eastern hemlock (Tsuga forest composition when they deline~ted canadensis (L.) Carr.), sugar maple (Acer the Black Swamp lake plain into several saccharum), red maple (A. rubrum), white similar regions. Kaatz (1955), in his histori­ ash (Fraxinus americana), American elm cal description of the Black Swamp,

16 Figure 5. Beech-maple-hemlock forests typical of riparian forests of the eastern Erie and Ontario Lake Plain .

described an area surrounding the set­ tlers' route that crossed the swamp from South Central Great Lakes southeast to northwest as low and wet, and being traversed by many streams. Here, the most frequently mentioned trees We found no source depicting the South were ash (Fraxinus), elm (Ulmus), oak Central Great Lakes ecoregion in Ohio (Quercus), maple (Acer), basswood (Tilia), (Figure 2). One study from southeastern hickory (Carya), and cottonwood (Popu­ Michigan (Hammitt and Barnes 1989), lus) species. Kaatz (1955) also noted that within this ecoregion, examined the swamp forest species were highly sensi­ vegetation of a 150-year-old upland oak­ tive to small variations in surface drainage hickory forest; however, the study does which resulted in a great variety of species not include any riparian forest ecosystems. occupying the wetter sites, in contrast to the beech-maple or oak-hickory forest as­ sociations on better drained sites.

17 Conclusions

Our review of the potential sources systems. Less information is available of information available to resource on the composition of the herbaceous managers on the composition and ground-flora, or the structure of either the structure of undisturbed riparian forests overstory or ground-flora vegetation. of Ohio suggests that these characteristics can be quite variable, depending largely Consequently, the available information on the influence of surficial geology, on reference conditions for riparian forests stream valley geomorphology, soils, in the state is limited, making it difficult to and predominate disturbance patterns. identify specific reference vegetation states Such relationships are similar to those for riparian areas. Additionally, without documented by many others, including some idea of the reference conditions, it is Goebel et al. (1996), Hupp and Osterkamp nearly impossible to measure the success (1996), Pabst and Spies (1998), and Bendix of riparian forest restoration efforts. and Hupp (2000). It should be noted, however, that various The result is that riparian vegetation is small old-growth remnants exist across the often arrayed in predictable patterns state with small streams or rivers (Figure across stream valleys and, with an 2). In many of these remnants, some understanding of the factors that regulate information is available on the riparian their development, reference vegetative vegetation-environment relationships. states can be determined. For example, there are several remnant forests distributed across the state, and By and large, while statewide pre­ information on these forests has been European settlement maps and regional published (e.g., McCarthy et al., 2001); county-wide surveys provide generalized however, no published information information on the broad forest types that was found detailing the composition once occurred across the state, they lack or structure of the riparian forests, the specific information on forest stand including those at Dysart Woods, Fowler structure and vegetation-environment Woods, Davey Woods, and Lawrence relationships needed to adequately predict Woods (Figure 2). Additionally, there are reference vegetative states. As such, these likely other areas in the state that can be sources of information are too coarse for used to help develop riparian reference developing restoration templates for on­ conditions, including forests owned by the-ground riparian forest restoration. The Nature Conservancy and other private organizations. However, the studies of the few remaining old-growth forests in the state elucidate Based on our review, it is clear that a many of the finer-scale vegetation­ framework is needed to help identify environment relationships, providing reference conditions for riparian forests considerable data on the overstory across the state. Hierarchy theory (O'Neill composition of these reference riparian et al., 1986) provides the conceptual

18 basis for an approach we suggest and Starr, 1982; Baker and Barnes, 1998; to understanding interrelationships Bendix and Hupp, 2000). Thus, we suggest among riparian forests and multi-scale that through detailed characterizations environmental factors that can be used of the remaining old-growth and least­ to develop reference or benchmark disturbed mature second-growth riparian conditions for riparian areas across Ohio. forests, reference vegetative conditions for these systems can be developed. When applied to riparian landscapes, Specifically, by quantifying the landscape­ hierarchy theory predicts that the upper scale and local-scale geomorphic features levels of the hierarchy (e.g., ecoregions, that shape the composition and structure physiographic systems) constrain a of riparian areas, we can better understand complex array of hydrogeomorphic the natural variation in riparian forests processes that in turn mediate the and develop suites of reference conditions dynamics of lower hierarchical levels, within and among Ohio's different including stream valley shape, fluvial ecoregions. landforms, and plant communities (Allen

19 Literature Cited

Allen, T. F. H., and T. B. Starr. 1982. Hier­ Blinn, C. R., and M. A. Kilgore. 2001. Ri­ archy: Perspectives for Ecological Complexity. parian management guidelines: A sum­ University of Chicago Press, Chicago, Ill. mary of state practices. Journal of Forestry 310 pp. 99:11-17.

Anderson, D. M. 1982. Plant communities Boerner, R. E. J., and D.-S. Cho. 1987. of Ohio: A preliminary classification and de­ Structure and composition of Goll Woods, scription. Division of Natural Areas and an old-growth forest remnant in north­ Preserves, Ohio Department of Natural western Ohio. Bulletin of the Torrey Botani­ Resources, Columbus, Ohio. 183 pp. cal Club 114:173-189.

Andreas, B. K. 1989. The vascular flora of the Boerner, R. E. J., and J. G. Kooser. 1991. glaciated Allegheny Plateau region of Ohio. Vegetation of Drew Woods, an old-growth Ohio Biological Survey Bulletin (New remnant in western Ohio, and issues of Series) 8:1-191. preservation. Natural Areas Journal 11:48- 54. Anliot, S. E. 1973. The vascular flora of Glen Helen, Clifton Gorge, and John Bryan State Braun, E. L. 1928. The vegetation of the Park. Ohio Biological Survey Biological Mineral Springs Region of Adams County, Notes. No. 5. The Ohio State University, Ohio. Ohio Biological Survey Bulletin 15. 3: Columbus, Ohio. 162 pp. 377-517.

Aronson, J., S. Dhillion, and E. Le Floc'h. Braun, E. L. 1936. Forests of the Illinoian 1995. On the need to select an ecosystem Till Plains of southwestern Ohio. Ecological of reference, however imperfect: A reply Monographs 6:89-149. to Pickett and Parker. Restoration Ecology 3: 1-3. Braun, E. L. 1950. Deciduous Forests of Eastern North America. Hafner Publishing Aughanbaugh, J. 1964. An ecological study Company, New York. 596 pp. of Crall Woods . Ohio Biological Survey Bio­ logical Notes. No. 2. The Ohio State Uni­ Braun, E. L. 1969. An ecological survey of versity, Columbus, Ohio. 55 pp. the vegetation of Fort Hill State Memorial, Highland County, Ohio, and annotated list of Baker, M. E., and B. V. Barnes. 1998. Land­ vascular plants. Ohio Biological Survey Bul­ scape ecosystem diversity of river flood­ letin (New Series) 3:1-131. plains in northwestern lower Michigan, USA. Canadian Journal of Forest Research 28: Bryant, W. S. 1987. Structure and composi­ 1405-1418. tion of the old-growth forests of Hamilton County, Ohio, and environs. Pp. 317-324, Bendix, J., and C. R. Hupp. 2000. Hydro­ In: R. L. Hay, F. W. Woods, and H. DeSelm logical and geomorphological impacts on (Eds.). Proceedings of the Central Hardwoods riparian plant communities. Hydrological Conference VI. University of Tennessee, Processes 14:2977-2990. Knoxville, Tenn.

20 Chapman, A.G. 1944. Original forests. Pp. water riparian areas of north-central Ohio. 73-84. In: 0. D. Diller (Ed.). Ohio's Forest Proceedings, Thirteenth Central Hardwood Resources. Forestry Publication No. 76. Forest Conference. Champaign-Urbana, Ill. Ohio Agricultural Experiment Station, USDA Forest Service General Technical Wooster, Ohio. In cooperation with the Report (in press). Central States Forest Experiment Station, Columbus, Ohio. Goebel, P. C, B. J. Palik, L. K. Kirkman, and L. West. 1996. Geomorphic influences Cho, D.-S., and R. E. J. Boerner. 1991a. on riparian forest composition and struc­ Canopy disturbance patterns and regen­ ture in a karst landscape of southwestern eration of Quercus species in two Ohio old­ Georgia. Pp. 110-116. In: K. M. Flynn, (Ed.). growth forests. Vegetation 93:9-18. Proceedings of the Southern Forested Wetlands Ecology and Management Conference. Clem­ Cho, D.-S., and R. E. J. Boerner. 1991b. son University, Clemson, S.C. Structure, dynamics, and composition of Sears Woods and Carmean Woods State Gordon, R. B. 1966. Original vegetation of Nature Preserves, North-Central Ohio. Ohio at the time of the earliest land surveys. Castanea 56:77-89. Ohio Biological Survey Informative Circu­ lar No. 1 (map). Cobbe, T. J. 1943. Variations in the Cabin Run Forest, a climax area in southwestern Gordon, R. B. 1969. The natural vegetation Ohio. American Midland Naturalist 29:89- of Ohio in pioneer days. Ohio Biological 105. Survey Bulletin (New Series) 3:1-109. Davis, M. B. (Ed.). 1996. Eastern Old­ Gregory, S. V. 1999. Riparian management Growth Forests: Prospects for Rediscovery and in the 21st century. Pp. 69-85. In: K. A. Recovery. Island Press, Washington, D.C. Kohm, J. F. Franklin, and J. W. Thomas 383 pp. (Eds.). Creating a Forestry for the 21st Cen­ tury. Island Press, New York. 576 pp. deMaynadier, P. G., Jr., and M. L. Hunter, Jr. 1995. The relationship between forest Gregory, S. V., F. J. Swanson, W. A. McKee, management and amphibian ecology: A and K. W. Cummins. 1991. An ecosystem review of the North American literature. perspective of riparian zones. BioScience Environmental Review 3: 230-261. 41:540-551. Diller, 0. D. 1932. The Vegetation of Logan Griggs, R. F. 1914. A botanical survey of the County, Ohio. M.S. Thesis, The Ohio State Sugar Grove Region. Ohio Biological Survey University, Columbus, Ohio. Bulletin 3. 1:245-340. Goebel, P. C., and D. M. Hix. 1996. Hammitt, W. E., and B. V. Barnes. 1989. Development of mixed-oak forests in Composition and structure of an old-growth southeastern Ohio: A comparison of oak-hickory forest in southern Michigan over second-growth and old-growth forests. 20 years. Pp. 247-253. Seventh Central Forest Ecology and Management 84:1-21. Hardwood Forest Conference, Carbondale, Ill. March 5-8, 1989. Goebel, P. C., D. M. Hix, C. E. Dygert, and K. L. Holmes. 2003. Vegetation-environ­ ment relationships of old-growth head

21 Hupp, C. R., and W. R. Osterkamp. 1996. WO-WSA-5. USDA Forest Service, Eco­ Riparian vegetation and fluvial geomor­ system Management, Washington, D.C. phic processes. Geomorphology 14:277-295. 267 pp. Ilhardt, B. L., E. S. Verry, and B. J. Palik. Norris, F. H. 1948. Primary Forest Types of 2000. Defining riparian areas. Pp. 23-42. In: Highland County, Ohio. Ph.D. Dissertation, E. S. Verry, J. W. Hornbeck, and C. A. Doll­ The Ohio State University, Columbus, off (Eds.). Riparian Management in Forests of Ohio. the Continental Eastern United States. Lewis Publishers, New York. 432 pp. O'Neill, R. V., D. L. DeAngelis, J.B. Waide, and T. F. H. Allen. 1986. A Hierarchical Con­ Irwin, N. M. 1920. The cedar cliffs prairie cept of Ecosystems. Princeton University opening of the Cincinnati region. Pp. Press, Princeton, N.J. 264 pp. 201-233. Proceedings of the Ohio Academy of Science. Vol. 8, part 5, special papers No. O'Laughlin, J., and G. H. Belt. 1995. Func­ 21. tional approaches to riparian buffer strip design. Journal of Forestry 93:29-32. Jones, C.H. 1936. The Vegetation of Meigs County. M.S. Thesis, The Ohio State Uni­ Pabst, R. J., and T. A. Spies. 1998. Distri­ versity, Columbus, Ohio. bution of herbs and shrubs in relation to landform and canopy cover in riparian Kaatz, M. R. 1955. The Black Swamp: A forests of coastal Oregon. Canadian Journal study in historical geography. Annals of the of Botany 76:298-315. Association of American Geographers 45:1-35. Pickett, S. T. A., and V. T. Parker. 1994. Kartez, J. T. 1994. A Synonymized Checklist Avoiding the old pitfalls: Opportunities of the Vascular Flora of the United States, in a new discipline. Restoration Ecology 2: Canada, and Greenland. Timber Press, Port­ 75-79. land, Ore. Runkle, J. R., and G. G. Whitney. 1987. McCarthy, B. C., C. A. Hammer, G. L. Vegetation-site relationships in the Lake Kauffman, and P. D. Cantino. 1987. Katherine State Nature Preserve, Ohio: A Vegetation patterns and structure of an northern outlier of the mixed-mesophytic old-growth forest in southeastern Ohio. forest. Ohio Journal of Science 87:36-40. Bulletin of the Torrey Botanical Club 114:33- 45. Sampson, H. C. 1927. The primary plant associations of Ohio: Their distribution McCarthy, B. C., C. J. Small, and D. L. and their significance as habitat indices. Rubino. 2001. Composition, structure, and The Ohio Journal of Science 27:301-309. dynamics of Dysart Woods, an old-growth mixed mesophytic forest of southeastern Sampson, H. C. 1930. Succession in the Ohio. Forest Ecology and Management 140: swamp forest formation in northern Ohio. 193-213. The Ohio Journal of Science 30:340-357. McNab, W. H., and P. E. Avers (Comps). Schlesinger, W. H. 1971. The vegetative 1994. Ecological Subregions of the United composition of a beech-maple climax States: Section Descriptions. Admin. Publ.

22 forest in the glaciated plateau of north­ Bulletin No. 108, June 1938. The Ohio State eastern Ohio. The Ohio Journal of Science 71: University and Ohio Agricultural Experi­ 174-180. ment Station, Columbus, Ohio.

Sears, P. B. 1925. The natural vegetation Vankat, J. L., W. H. Blackwell, and W. E. of Ohio. I. A map of the virgin forest. The Hopkins. 1975. The dynamics of Hueston Ohio Journal of Science 25:139-149. Woods and a review of the question of the successional status of the southern beech­ Sears, P. B. 1926. The natural vegetation of maple forest. Castanea 40:290-308. Ohio. II. The prairies. The Ohio Journal of Science 26:128-146. Werth, C. R., W. P. Pusateri, G. W. Snyder, and W. H. Blackwell. 1984. Vascular plants Segelken, J. 1929. The Plant Ecology of the of and Nature Hazelwood Botanical Preserve. Ohio Biologi­ Preserve. In: G. E. Willeke (Ed.). Hueston cal Survey Bulletin No. 21, 4:219-269. Woods State Park and Nature Preserve Pro­ Swanson, A. M., and J. L. Vankat. 2000. ceedings of Symposium, April 16-18, 1982. Woody vegetation and vascular flora of Williams, A. B. 1936. The composition and an old-growth mixed-mesophytic forest in dynamics of a beech-maple climax com­ southwestern Ohio. Castanea 65:36-55. munity. Ecological Monographs 6:317-408. Transeau, E. N., and H. C. Sampson. 1938. Wyant, J. G., R. A. Meganck, and S. H. Primary vegetation areas in Ohio. Pp. Ham. 1995. A planning and decision­ 18-19. In: J. H. Sitterley, and J. I. Falconer making framework for ecological restora­ (Eds.). Better Land Utilization for Ohio. De­ tion. Environmental Management 19:789-96. partment of Rural Economics Mimeograph

23 Appendix

Latin Name Common Name

Acer negundo boxelder Acernigrum black maple Acerrubrum red maple Acer saccharinum silver maple Acer saccharum sugar maple Aesculus flava yellow buckeye Aesculus glabra Ohio buckeye Agrimonia parviflora harvestlice Alisma plantago-aquatica American waterplantain Apocynum cannabinum Indianhemp Asarum canadense Canadian wildginger Asclepias incarnata swamp milkweed Asimina triloba pawpaw Betula alleghaniensis yellow birch Betula nigra river birch Bidens spp beggarticks Blephilia hirsuta hairy pagoda-plant Botrychium virginianum rattlesnake fern Caltha palustris yellow marsh marigold Cardamine concatenata cutleaf toothwort Carex lurida shallow sedge Carex spp sedge Carpinus caroliniana American hornbeam Carya cordiformis bitternut hickory Carya glabra pignut hickory Carya laciniosa shellbark hickory Carya ovata shagbark hickory Carya spp hickory Celastrus scandens American bittersweet Cercis canadensis eastern redbud Chelone glabra white turtlehead Cicuta maculate spotted water hemlock Cinna arundinacea sweet woodreed Cirsium arvense Canada thistle Claytonia virginica Virginia springbeauty Cornus florida flowering dogwood Corylus americana American hazelnut Cryptotaenia Canadensis Canadian honewort Cystopteris fragilis brittle bladderfern Deparia acrostichoides silver false spleenwort Eleocharis ovata ovate spikerush Elymus hystrix L. var. hystrix eastern bottlebrush grass Equisetum arvense field horsetail Erythronium albidum white fawnlily Eupatorium purpureum sweetscented joepyeweed

24 Latin Name Common Name

Eupatorium sessilifolium upland boneset Fagus grandifolia American beech Fraxinus americana white ash Fraxinus pennsylvanica green ash Fraxinus quadrangulata blue ash Fraxinus spp ash Galium triflorum fragrant bedstraw Geum virginianum cream avens Glechoma hederacea ground ivy Gleditsia triacanthos honeylocust Hackelia virginiana beggarslice Helenium autumnale common sneezeweed Helianthus hirsutus hairy sunflower Helianthus tuberosus Jerusalem artichoke Hypericum punctatum spotted St. Johnswort Impatiens capensis jewelweed Jeffersonia diphylla twinleaf Juglans cinerea butternut Juglans nigra black walnut Laportea Canadensis Canadian woodnettle Leersia oryzoides rice cutgrass Leersia virginica whitegrass Lindera benzoin northern spicebush Liquidambar styraciflua sweetgum Liriodendron tulipifera tuliptree Lobelia siphilitica great blue lobelia Lysimachia nummularia creeping jenny Magnolia tripetala umbrella-tree Matteuccia struthiopteris ostrich fern Mimulus alatus sharpwing monkeyflower Muhlenbergia spp muhly Onoclea sensibilis sensitive fern Ostrya virginiana hophornbeam Oxalis stricta common yellow oxalis Parietaria pennsylvanica pellitory Parthenocissus quinquefolia (L.) Planch. Virginia creeper Penstemon calycosus longsepal beardtongue Penthorum sedoides ditch stonecrop Phalaris arundinacea reed canarygrass Phlox divaricata wild blue phlox Phlox paniculata fall phlox Pilea pumila Canadian clearweed Platanus occidentalis American sycamore Podophyllum peltatum mayapple Polygonum arifolium halberdleaf tearthumb Polygonum hydropiper marshpepper knotweed Polygonum punctatum dotted smartweed Polygonum sagittatum arrowleaf tearthumb

25 Latin Name Common Name

Polygonum virginianum jumpseed Populus deltoides eastern cottonwood Prenanthes altissima tall rattlesnakeroot Prunella vulgaris common selfheal Prunus serotina black cherry Quercus alba white oak Quercus bicolor swamp white oak Quercus macrocarpa bur oak Quercus muhlenbergii yellow chestnut oak Quercus rubra northern red oak Quercus spp oak Ranunculus hispidus Michx. var. nitidus bristly buttercup Rudbeckia laciniata cutleaf coneflower Salix nigra black willow Salix spp willow Sambucus canadensis common elderberry Scutellaria lateriflora blue scullcap Silphium perfoliatum cup plant Solidago canadensis Canada goldenrod Solidago canadensis var. scabra Canada goldenrod Solidago gigantea giant goldenrod Solidago patula roundleaf goldenrod Symphyotrichum lanceolatum var. lanceolatum white panicle aster Symphyotrichum pilosum hairy white oldfield aster Symphyotrichum prenanthoides crookedstem aster Tilia americana American basswood Toxicodendron radicans eastern poison ivy Trillium sessile toadshade Tsuga canadensis eastern hemlock Typha latifolia broadleaf cattail Ulmus americana American elm Ulmus rubra slippery elm Urtica dioica stinging nettle Verbena urticifolia white vervain Verbesina alternifolia wingstem Vernonia fasciculata prairie ironweed Vernonia gigantea giant ironweed Viola pubescens downy yellow violet Viola striata striped cream violet Vitis aestivalis summer grape

26 The information in this publication is supplied with the understanding that no discrimination is intended and no endorsement by The Ohio State University; the College of Food, Agricultural, and Environmental Sciences; the Ohio Agricultural Research and Development Center; or Ohio State University Extension is implied. Due to constantly changing laws and regulations, no liability for the recommendations can be assumed.

All publications of the Ohio Agricultural Research and Development Center are available to clientele on a nondiscriminatory basis without regard to race, color, creed, religion, sexual orientation, national origin, gender, age, disability, or Vietnam-era veteran status. 10-03 Jaf

The Ohio State University Ohio Agricultural Research and Development Center 1680 Madison Avenue Wooster, Ohio 44691-4096 330-263-3700 T · H · E OHIO SfAlE UNIVERSITY OARilL

Ohio Agricultural Research and Development Center

In Partnership With Ohio State University Extension College of Food, Agricultural, and Environmental Sciences