Figure 4. Maple and elm riparian forest along the Sandusky River flowing through the Sears-Carmean State Nature Preserve.
Twelve studies were found that describe the riparian forests of the Central Till Plains, Beech-Maple ecoregion (Figure 2), with 10 detailing riparian vegetationenvironment relationships, making it the most intensively studied of any ecoregion in the state (Table 1). These studies included riparian areas associated with intermittent stream channels, riparian areas along small streams and ravines, and riparian areas associated with large rivers, broad flats or floodplains, and bottomland swamps.
Crall Woods in north-central Ohio and Cabin Run Forest in southwestern Ohio are both examples of old-growth forests traversed by intermittent streams (Figure 2). However, Aughanbaugh (1964), in his study of Crall Woods, does not differentiate the undulating terrain into riparian and upland areas. The major tree species encountered were sugar maple (Acer saccharum), American basswood (Tilia americana), American beech (Fagus grandifolia), tuliptree (Liriodendron tulipifera), American elm (Ulmus americana), northern red oak (Quercus rubra), shagbark hickory (Carya ovata (P. Mill.) K. Koch), and hophornbeam (Ostrya virginiana (P. Mill.) K. Koch), with more than 90 herbaceous species recorded in the wooded area.
At Cabin Run Forest along intermittent streams that dissect the upland areas dominated by mixed-mesophytic forests, Cobbe (1943) found that sugar maple (Acer saccharum), American beech (Fagus grandifolia), tuliptree (Liriodendron tulipifera), American basswood (Tilia americana), black walnut (Juglans nigra), and white oak (Quercus alba) were nearly always present in the riparian area, with their abundance dependant on local site and microclimatic conditions.
Cobbe (1943) also documents that a small stream bordering the forest at Cabin Run supported a streamside community where American sycamore (Platanus occidentalis) was dominant, along with several other species including sugar maple (Acer saccharum), black walnut (Juglans nigra), tuliptree (Liriodendron tulipifera), and pignut hickory (Carya glabra).
The herbaceous layer was composed of rattlesnake fern (Botrychium virginianum (L.) Sw.), brittle bladderfern (Cystopteris fragilis (L.) Bernh.), Canadian wildginger (Asarum canadense L.), Virginia springbeauty (Claytonia virginica L.), cutleaf toothwort (Cardamine concatenata (Michx.) Sw.), white fawnlily (Erythronium albidum Nutt.), fragrant bedstraw (Galium triflorum Michx.), jewelweed (Impatiens capensis), twinleaf (Jeffersonia diphylla (L.) Pers.), great blue lobelia (Lobelia siphilitica L.), common yellow oxalis (Oxalis stricta L.), wild blue phlox (Phlox divaricata L.), mayapple (Podophyllum peltatum L.), toadshade (Trillium sessile L.), downy yellow violet (Viola pubescens Ait.), and striped cream violet (V. striata Ait).
The old-growth forest of Emery Woods in Hamilton County (Figure 2) includes an area where ravine slopes, associated with small and often times sluggish streams, are prominent. At Emery Woods, Swanson and Vankat (2000) found that the forests of these ravine bottoms tend to support mixed mesophytic species including sugar maple (Acer saccharum), hickory (Carya spp.), oak (Quercus spp.), American beech (Fagus grandifolia), and tuliptree (Liriodendron tulipifera).
These finding are similar to those reported by Braun (1936) in her descriptive comparison of ravine slope forests and flats within the Illinoian till plains region of southwestern Ohio. Similar relationships were observed along a small stream draining a large ravine and two main branches with swampy, flat-bottomed floors at the Hazelwood Botanical Preserve (HBP), also in Hamilton County (Figure 2).
A variety of habitats from extremely wet swamp communities to mesophytic associations were observed at HBP. Specifically, Segelken (1929) found that herbaceous vegetation far outnumbered the woody plants on the wetter sites. Those species reported to occur in the wetter areas include yellow marsh marigold (Caltha palustris L.), rice cutgrass (Leersia oryzoides), arrowleaf tearthumb (Polygonum sagittatum L.), jewelweed (Impatiens capensis), and broadleaf cattail (Typha latifolia L.). Prairie ironweed (Vernonia fasciculata Michx.) was common along the stream, while sweetscented joepyeweed (Eupatorium purpureum L.) and common elderberry (Sambucus canadensis L.) were found on the ravine floor in drier places.
Several black willow (Salix nigra) communities were found growing on the ravine floor in both wet and drier sites. Along the sloping margins of the stream bank, American elm (Ulmus americana) and American sycamore (Platanus occidentalis) were found distributed where hydromesophytic conditions prevail.
In addition to these species, Segelken (1929) found the following herbaceous plants on the ravine floor: harvestlice (Agrimonia parviflora Ait.), American waterplantain (Alisma plantago-aquatica L.), Indianhemp (Apocynum cannabinum L.), swamp milkweed (Asclepias incarnata L.), shallow sedge (Carex lurida Wahlenb.), Canadian honewort (Cryptotaenia canadensis (L.) DC.), white turtlehead (Chelone glabra L.), ovate spikerush (Eleocharis ovata (Roth) Roemer & J. A. Schultes), field horsetail (Equisetum arvense L.), cream avens (Geum virginianum L.), spotted St. Johnswort (Hypericum punctatum Lam.), beggarslice (Hackelia virginiana (L.) I. M. Johnston), great blue lobelia (Lobelia siphilitica L.), sharpwing monkeyflower (Mimulus alatus Ait.), ground ivy (Glechoma hederacea L.), pellitory (Parietaria pennsylvanica Muhl. ex Willd.), ditch stonecrop (Penthorum sedoides L.), Canadian clearweed (Pilea pumila (L.) Gray), halberdleaf tearthumb (Polygonum arifolium L.), dotted smartweed (Polygonum punctatum Ell.), common selfheal (Prunella vulgaris L.), blue skullcap (Scutellaria lateriflora L.), roundleaf goldenrod (Solidago patula Muhl. ex Willd.), white vervain (Verbena urticifolia L.), and summer grape (Vitis aestivalis Michx.).
At Cedar Cliffs Prairie, located on a bluff of the Little Miami River (Figure 2), a broad floodplain extends unbroken for miles, with prairie openings that occur in exposed locations where evaporation and exposure are great. The bluff is indented by ravines and gullies of various sizes. These isolated and protected parts are very different from the bluffs themselves.
Irwin (1920) reported that the extremely steep bluff was almost bare of vegetation, whereas the ravines supported a mesophytic habitat where northern red oak (Quercus rubra) and sugar maple (Acer saccharum) were dominant. American elm (Ulmus americana), black walnut (Juglans nigra), blue ash (Fraxinus quadrangulata Michx.), eastern redbud (Cercis canadensis L.), and yellow chestnut oak (Quercus muhlenbergii Engelm.) occurred as saplings.
The most important species of the diverse herbaceous layer consisted of muhly grass (Muhlenbergia spp.), hairy sunflower (Helianthus hirsutus Raf.), American bittersweet (Celastrus scandens L.), upland boneset (Eupatorium sessilifolium L.), longsepal beardtongue (Penstemon calycosus Small), and tall rattlesnakeroot (Prenanthes altissima L.).
Anliot (1973) reported on the vascular flora of Glen Helen, Clifton Gorge, and John Bryan State Park in north central Greene County (Figure 2). The topography of the study area is fairly flat except where dissected by the Little Miami River and its tributaries. Anliot (1973) found there was great variability associated with soil characteristics and drainage among the floodplain communities.
Willow-eastern cottonwood-American sycamore (Salix spp.-Populus deltoides-Platanus occidentalis) communities were found mainly along the larger waterways. Where better drainage occurred, silver maple (Acer saccharinum), American elm (Ulmus americana), and ash (Fraxinus spp.) were dominant, while either bur oak (Quercus macrocarpa Michx.), swamp white oak (Q. bicolor Willd.), and shellbark hickory (Carya laciniosa (Michx. f.) G. Don) or northern red oak (Quercus rubra) and American basswood (Tilia americana) forest communities dominated the best drained sites. In addition, bur oak (Quercus macrocarpa), silver maple (Acer saccharinum), black walnut (Juglans nigra), butternut (J. cinerea L.), Ohio buckeye (Aesculus glabra), and black maple (Acer nigrum Michx. f.) also occurred frequently on alluvial deposits along streams.
Anliot (1973) also documented successional patterns along the Little Miami River in old fields abandoned less than 10 years. The fields first became reforested primarily with boxelder (Acer negundo L.), American sycamore (Platanus occidentalis), and honeylocust (Gleditsia triacanthos L.), often in pure stands. Before a canopy developed, Canada goldenrod (Solidago canadensis L. and S. canadensis var. scabra Torr. & Gray), hairy white oldfield aster (Symphyotrichum pilosum (Willd.) Nesom.), white panicle aster (Symphyotrichum lanceolatum var. lanceolatum (Willd.) Nesom.), wingstem (Verbesina alternifolia (L.) Britt. ex Kearney), giant ironweed (Vernonia gigantea (Walt.) Trel.), Jerusalem artichoke (Helianthus tuberosus L.), Canada thistle (Cirsium arvense (L.) Scop.), hairy pagoda-plant (Blephilia hirsuta (Pursh) Benth.), creeping jenny (Lysimachia nummularia L.), and stinging nettle (Urtica dioica) were common.
Hueston Woods (Figure 2), a nearby state park and nature preserve of comparable size containing old-growth forest remnants, lies near the southern boundary of the Beech-Maple forest region (Braun 1950) near the transition to western and mixed mesophytic forests (Vankat et al., 1975). However, unlike the Glen Helen-Clifton Gorge-John Bryan complex, the mildly dissected topography of Hueston Woods lacks the rocky cliffs and hill prairies of the former areas (Werth et al., 1984).
While the work of Vankat et al. (1975) focused on the woody vegetation of the upland portion of Hueston Woods, Werth et al. (1984) compiled a list of vascular plant flora collected from areas throughout the park and preserve. Included are 40 streamside species, nine species found in wooded ravines, and 46 from wet woods. Additionally, Braun (1950) lists sugar maple (Acer saccharum) as the dominant canopy species occurring on alluvial terraces at Hueston Woods with American elm (Ulmus americana), white oak (Quercus alba), and shagbark hickory (Carya ovata) present as minor components.
While the preceding examples are from southwestern Ohio, the Sears Woods and Carmean Woods complex in the till plains of north-central Ohio also exhibit good examples of typical riparian forests of the Central Till Plains, Beech-Maple ecoregion (Figure 2). In the two studies by Cho and Boerner (1991a,b), Sears Woods is described as comprising an area of upland moraine dissected by several major stream drainages and a large, flat floodplain terrace along the Sandusky River. Both areas have moderately well to well-drained soils. In contrast, the topography of Carmean Woods is level with very poorly drained soils.
The old-growth bottomland areas of the Sears-Carmean Woods complex are dominated by silver maple (Acer saccharinum), American basswood (Tilia americana), American elm (Ulmus americana), and sugar maple (Acer saccharum), and the total tree species composition corresponds well to the early descriptions of the red oak-basswood phase of the elm-ash swamp forest as described by Sampson (1930). The upland forest component, in which white oak (Quercus alba), northern red oak (Q. rubra), and white ash (Fraxinus americana) were present in the canopy with co-dominants sugar maple (Acer saccharum) and American beech (Fagus grandifolia), also corresponds well to Gordon's (1969) description of beech-sugar maple forest.
No ground-flora information was reported in either study of Sears-Carmean Woods by Cho and Boerner (1991a,b).
Examples of old-growth bottomland swamp forest communities in the Central Till Plains, Beech-Maple ecoregion can be found at Drew Woods, an isolated old-growth forest remnant located on the poorly drained till plains of western Ohio (Figure 2). Boerner and Kooser (1991) reported that "early European settlers found this area covered by a complex of swamp forest and bottomland forest broken only by occasional stream corridors."
Species more abundant on lower, wetter sites included sugar maple (Acer saccharum), swamp white oak (Quercus bicolor), northern red oak (Q. rubra), bur oak (Q. macrocarpa), and shellbark hickory (Carya laciniosa). In contrast, sugar maple (Acer saccharum), white oak (