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The structure and composition of vegetation in the lake-fill Peatlands of Indiana.

Anthony L. Swinehart (1)

ABSTRACT. The vegetation of 16 lake-fill peatlands in northern Indiana was systematically sampled. Peatland types included fens, tall shrub bogs, leatherleaf bogs and forested peatlands. No significant difference in species richness among the four peatland types was identified from the systematic sampling. Vegetation composition and structure, along with water chemistry variables, was analyzed using multivariate statistical analysis. Alkalinity and woody plant cover accounted for much of the variability in the herbaceous and ground layers of the peatlands, and a successional gradient separating the peatlands was evident. A multivariate statistical comparison of leatherleaf bogs from Indiana, Michigan, Ohio, New York, New Jersey and New Hampshire was made on the basis of vegetation composition and frequency and five climatic variables. The vascular vegetation communities of Indiana peatlands and other peatlands in the southern Great Lakes region are distinct from those in the northeastern U.S., Ohio and the no rthern Great Lakes. Some of these distinctions are attributed to climatic factors, while others are related to biogeographic history of the respective regions.

Keywords: Peatlands, leatherleaf bogs, fens, ecological succession, phytogeography

Within midwestern North America, the northern counties of Illinois, Indiana and Ohio represent the southern extent of peatland communities containing characteristic plant species of northern or boreal affinity. Here, peatlands are restricted to favorable microclimates such as small, water-filled kettles; and they never occur in great expanses as they do in the northern Lake States, the northeast and Canada, where regional topography and climate are more favorable for peat production. Peatland plant communities that occur at their range limits are important because they enhance the biological richness of these areas and potentially increase the genetic diversity of the plant species (Lesica & Allendorf 1995) due to their responses to heightened environmental stress. The occurrence of peatland communities with boreal affinities in marginal environments may amplify factors that favor their formation, and therefore provide excellent opportunities to study phytogeography and glacial relics.

While characteristically northern species such as Chamaedaphne calyculata, Andromeda glaucophylla, and Carex oligosperma often make "southern outlier peatlands" conspicuous to botanists, studies of such peatlands in New York, New Jersey and southern Michigan have shown a flora exhibiting mixed geographic affinity (Crow 1969; Lynn & Karlin 1985; Karlin & Lynn 1988). Although many studies have characterized the peatland flora of Canada (Jeglum et al. 1974; Slack et al. 1980; Sims et al. 1982; Glaser 1992), the northern Great Lakes region (Gates 1942; Vitt & Slack 1975; Glaser 1987; Glaser et al. 1992), and the northeastern United States (Worley & Sullivan 1980; Worley 1981; Lynn & Karlin 1985; Sorenson 1986; Damman & French 1987; Dunlop 1987; Karlin & Lynn 1988; Fahey & Crow 1995), relatively few have characterized the peatland plant communities in the southern Great Lakes region.

While extra-local and regional accounts of peatland plant communities in the region have been presented by Transeau (1905; 1906) (southern Michigan) and Waterman (1926) (northern Illinois), Ohio is the only state in the area that is represented by thorough treatments of its peatland plant communities (see Jones 1941; Aldrich 1943; Andreas 1989; Andreas & Bryan 1990; Andreas & Knoop 1992). Local or individual accounts of peatlands in the region include sites in southern Michigan (Crow 1969a,b; Sytsma & Pippen 1981, 1982; Glime et al. 1980, 1982; Keough & Pippen 1981, 1984), northern Illinois (Reichie & Doyle 1965; Sheviak & Haney 1973), Ohio (Aughanbaugh & Diller 1968; McQueen & Giesy 1975; Collins et al. 1982; Knoop 1987; Lauschman 1991), and Indiana (Hessler 1896; Markle 1916; Cain 1928; Lindsey 1932; Potzger 1934; Hull 1937; Friesner & Potzger 1946; Starcs 1961; Welch 1963; Lindsey et al. 1969; Wilcox 1982; Wilcox et al. 1986; Swinehart 1994; and Swinehart & Starks 1994). Few of the investigations of Indian a peatlands were quantitative in nature, and many were conducted on the same peatlands, such as Cabin Creek Raised Bog, due to greater public familiarity with their location.

The relative dearth of scientific attention to the peatlands in Indiana is not a function of insufficient study areas. Over 90 notable sites are known (Swinehart 1997) and, doubtless, many others remain to be found by scientists. Sphagnum-dominated communities in the state have been documented as far south as the city of Indianapolis (Cain 1928), and brown-moss-dominated peatlands reach well into the southern half of Indiana (Friesner & Potzger 1946; Welch 1962). A conservative estimate of historic peatland coverage in the state (based on true peat soils rather than mucks) is 62,000 ha, the main concentration being in northcentral Indiana (see Swinehart 1997). Considering the average surface area of individual peatlands in Indiana (approximately 20 ha (Swinehart 1997)), it becomes apparent that peatlands were extremely frequent in the northern part of the state during pre-settlement times. Unfortunately, as much as 87% of Indiana's wetlands has been destroyed (Dahl 1990), mostly as a result of agricultural d evelopment of the landscape.

The objectives of the present paper are to: 1) characterize lake-fill peatlands in Indiana based on the composition and structure of the vegetation of 16 bogs, fens and forested peatlands, 2) investigate the floral, hydrological and seral relationships of the peatlands using canonical correspondence analysis, and 3) compare the peatland flora of Indiana to other peatlands in the southern Great Lakes region as well as to peatlands in the North and Northeast.

STUDY AREA

The 16 peatlands described in this paper are located within the northern three tiers of counties in Indiana (Fig. 1). The entire area is blanketed with glacial deposits, the most recent being of Wisconsin age (Fig. 1). The glacial drifts cover bedrock composed chiefly of Devonian and Mississippian shales. In some areas of northeast Indiana, this bedrock is separated from the surface by over 150 m of glacial deposits. The elevations of the peatlands range from 155-320 m, with a mean of 262 m (SD = 35) (Swinehart 1997). The glacial drifts are rich in limestone originating from the Michigan basin that, along with the presence of lime-rich bedrock, imparts a strongly alkaline reaction in the ground and surface waters of the region.

Northeast and north-central Indiana is characterized by hundreds of lakes, and a distinct lake district stretches from the Huron River Valley of Michigan (see Transeau 1905-6) through the northeast corner of Indiana to a point nearly 160 km southwest. These lake basins consist mostly of kettles, created in the interlobate area of the former Saginaw and Erie Lobes of Wisconsin age. The lakes occurring west of the interlobate area (north-central Indiana) are a result of the stagnation and downmelting of the Saginaw Lobe circa 15,000 years BP. All of the peatlands in the present paper are the result of in-filling of lake basins (Swinehart 1997; Swinehart & Parker 2000).

The climate of northern Indiana is summarized in Table 1. In relation to the more northern latitudes of Canada, the northern Great Lakes region, and the northeastern United States (with the exception of New Jersey), the mean annual temperature in northern Indiana is warmer on average (Table 1). Mean annual precipitation is similar to other regions. However, mean annual snowfall in northern Indiana is generally less, and the number of frost-free days is generally more with the exception of northern New Jersey which varies from 130-180 frost-free days (Lynn & Karlin 1985) (Table 1). Climatic conditions in Indiana most closely resemble those governing the peatland communities of northern Ohio (Table 1).

Most of the peatlands in the present study are located within the Quercus-Carya (oakhickory) forest type (see Lindsey et al. 1965). Thompson Bog, Mt. Pleasant Bog, Dutch Street Bog, and Svoboda Fen are within the Fagus-Acer (beech-maple) association. Shoemaker Bog occurs within the dry-prairie region of northwest Indiana.

All of the peatlands are presently surrounded by agricultural fields. While Mt. Pleasant Bog (18 ha), Ropchan Memorial Bog (14 ha), Tamarack Bog (14 ha), Thompson Bog (6 ha), Blueberry Bog (10 ha), Hickory Bog (0.6 ha) and Kiser Lake Fen (2 ha) have a limited terrestrial forest buffer, Little Chapman Bog (9 ha), Burket Bog (25 ha), Dutch Street Bog (16 ha), Leatherleaf Bog (4 ha), Shoemaker Bog (15 ha), Yost Bog (8 ha), Little Arethusa Bog (15 ha), Svoboda Fen (25 ha) and Binkley Fen (31 ha) are buffered only by a narrow margin of wet or mesic forest associated with the lagg.

The individual basins range in maximum depth from 4.5-17 m with a mean maximum depth of 14.7 m (Swinehart 1997). The surface peat is classified as Houghton Muck and ranges from 1-7 m in depth (Swinehart & Parker 2000). Only two leatherleaf bogs and one fen exhibit a central pond of open water. The remainder have their entire surface area covered with peat, although below the surface, the peat is mostly unconsolidated and often contains large, open pockets (Swinehart & Parker 2000).

METHODS

Definitions.--In some parts of Europe and in Canada, the term bog is usually confined to any peatland where nutrient input is strictly ombrotrophic, designating all other peatlands, acid or alkaline, strongly or weakly minerotrophic, as fens. Authors studying New England bogs (Worley (1981), Johnson (1985), and Davis & Anderson (1991)) agree that a "true bog" refers to ombrotrophic peatlands. Crum (1988), though sparingly, also uses this definition. The problem with using nutrient source as a defining character arises in the difficulty encountered when trying to separate a bog from a fen, because the outward characteristics (i.e., vegetation) of a minerotrophic Sphagnum-dominated peatland and an "ombrotrophic" Sphagnum-dominated peatland are sometimes patchy or even indistinguishable.

Several authors continue to use a more traditional definition of bog where domination by Sphagnum mosses is a delimiting criterion. This is because the differences between a Sphagnum-dominated peatland and a peatland not dominated by Sphagnum are quite obvious, and the relative rapidity by which a non-Sphagnum-dominated peatland becomes Sphagnum-dominated (see Vitt & Kuhry 1992; Kuhry et al. 1993) denotes drastic ecological changes during and subsequent to the transition. This abrupt, ecological transition should be reflected more definitively in the nomenclature that is used to describe these systems. Andreas (1985), Dunlop (1987), Karlin & Lynn (1988) and Andreas & Bryan (1990) have all used the term bog to refer to peatlands dominated by Sphagnum regardless of the source of nutrient inputs.

In the present paper, peatlands with a bryoflora dominated by Sphagnum are termed bogs, peatlands with a bryoflora dominated by mosses of the family Amblystegiaceae are termed fens, and peatlands that have developed a canopy of forest vegetation are termed forested peatlands--regardless of source of water/nutrients. Bogs, fens and forested peatlands are distinguished from marshes, swamps and sloughs by the presence of significant deposits of peat (partially decayed organic material) rather than muck, and the absence of long periods of standing water (water table is at or slightly below the surface). Differences in mineral richness among bogs are qualified with the terms "mineral-rich" or "mineral-poor."

Selection of sites.--Two main criteria were used in the selection of the peatlands for this investigation: 1) peatlands occurring in defined basins (rather than those associated with rivers and streams), and 2) peatlands exhibiting a characteristic flora that distinguished them from typical marshes, swamps and sloughs. Characteristic plants included Andromeda glaucophylla, Betula pumila, Carex oligosperma, Carex trisperma, Chamaedaphne calyculata, Coptis trifolia, Drosera spp., Larix laricina, Menyanthes trifoliata, Potentilla fruticosa, Sarracenia purpurea, Sphagnum spp., Vaccinium macrocarpon, and V. oxycoccos. Selection of individual sites was contingent upon authorization by landowners.

Vegetation sampling.--The 16 peatlands selected included three fens (Binkley, Kiser and Svoboda), four tall-shrub bogs (Blueberry, Hickory, Little Chapman and Thompson), five leatherleaf bogs (Burket, Dutch Street, Leatherleaf, Shoemaker and Yost), and four forested peatlands (Little Arethusa, Mt. Pleasant, Ropchan Memorial and Tamarack). Vegetation in open peatlands was sampled by establishing a baseline along the long axis of each peatland. Transects spaced at 25 m or 50 m intervals (depending on the size of the peatland), and stretching across the predominant vegetation of each peatland [i.e., exclusive of lagg (the nutrient-rich, swampy margins of a peatland)], were established perpendicular to the baseline. Nested quadrats were placed at 25 m or 50 m intervals along each transect. In tall-shrub bogs and some forested peatlands, terrain hindered systematic spacing of sampling plots. In such cases, a single line transect was established through the predominant vegetation in the respective peatland.

Vegetation was stratified on the basis of height or stem diameter at breast height (DBH): The tree layer included living stems > 4 cm DBH, the shrub layer included living stems [less than or equal to] 4 cm DBH and greater than 1 m in height, the herbaceous layer was vegetation less than 1 m in height, the ground layer was restricted to bryophytes, and the aquatic layer comprised submerged or floating plants (e.g., Potamogeton (pondweeds) and Lemna spp. (duckweeds)). The tree layer was sampled with 400 [m.sup.2] quadrats, the shrub layer with 100 [m.sup.2] quadrats, and all other layers with 1 [m.sup.2] quadrats.

Total percent frequency, total percent cover (density and basal area for trees), and importance values were calculated for each species in each peatland. Percent frequency was calculated by dividing the number of quadrats in which a species was found in a given peatland by the total number of quadrats sampled in each peatland. Percent cover was calculated by dividing the sum of observed cover values (determined visually) for each species in each peatland by the total number of quadrats in each peatland. Importance values for herbs and bryophytes represent the sum of the relative frequency and relative cover, divided by two and multiplied by 100.

Species that were not recorded in the quadrats, but were noted elsewhere in the peatland, were included in the species lists (Tables 3-7) as "present." However, no attempt was made to locate rare plants or to compile a complete flora for the sites. The Indiana Department of Natural Resources, Division of Nature Preserves maintains qualitative lists for many of the peatlands.

Vouchers for all bryophytes were placed in the private herbarium of the first author at Hillsdale College and in the Kriebel Herbarium of Purdue University (PUL). In addition, duplicate voucher specimens of Sphagnum spp. from Hickory, Leatherleaf and Tamarack Bogs were placed in the University of Michigan Herbarium (MICH). All other duplicate Sphagna were placed in the Herbarium of Binghamton University (BING). Nomenclature of Sphagnum follows Andrus (1980, 1987). Nomenclature of all other mosses follows Crum & Anderson (1981). Nomenclature of hepatics follows Crum (1991), and nomenclature of vascular plants follows Swink & Wilhelm (1994), with the exception of plants of the genus Toxicodendron.

Water chemistry.--Water was sampled at three locations in the area containing the predominant vegetation (e.g., open mat rather than lagg). All water samples were collected within a one-week period in late July (1995), after the rainy season had passed. Water was collected from pools of standing water or interstitial waters of the peat depending on conditions. Interstitial waters were obtained by pushing a glass jar into the peat and allowing water to enter the mouth of the jar. Titration of samples for total alkalinity, calcium hardness, and magnesium hardness was conducted within two hours of collection. Values for pH and specific conductance were taken on site with a Cole-Panner digital pH meter, and an Orion Conductivity/TDS meter, respectively.

Statistical analyses.--Canonical Correspondence Analysis (CCA) of the vegetation and environmental variables was conducted with PC-ORD Version 2.0 (McCune & Mefford 1995). Data used in the vegetation matrix include the percent cover for each herbaceous and aquatic layer species in each peatland. Data used in the environmental variable matrix include values for total alkalinity, pH, total percent shrub cover, and total tree basal area in each peatland.

In the regional comparison of vegetation in leatherleaf bogs, data were taken from several published studies. A data-type that was common to all of the papers was percent frequency for each species. To reduce bias caused by differences in sampling design, frequencies were converted to a scale of 1 to 5, where 1 = 1-19%, 2 = 20-39%, 3 = 40-59%, 4 = 60-79%, and 5 = 80-100%. Detrended Correspondence Analysis (DCA) was used to determine the variability in the vegetation. A second matrix that included mean annual precipitation, mean annual temperature, mean July temperature, mean January temperature, and mean number of frost-free days was incorporated with the vegetation data in a CCA ordination.

RESULTS AND DISCUSSION

Species richness.--A total of 131 vascular plants, 49 bryophytes, and one alga was recorded from the 16 peatlands (Tables 3-7). The mean number of vascular plants, calculated from species found in quadrats only, was determined for leatherleaf bogs, fens, tall-shrub bogs and forested peatlands. Although the mean number of species from the leatherleaf bogs appeared comparatively small, no significant differences in species richness among the four peatland types were identified. An analysis of species richness as a function of alkalinity (indicative of the degree of minerotrophy) showed no significant relationship (p = 0.107, [R.sup.2] = 0.175). Although peatlands at extremes of the gradient harbor characteristic, dominant plants that are adapted for specific pH and mineral conditions, data suggest that both ends of the gradient may reduce species richness because of common factors such as substrate inundation and anoxia. While mineral-rich peatlands and fens are generally characterized as having a richer flora , most of the species that contribute to the greater richness are probably infrequent or rare members of the community, and may result from greater habitat heterogeneity rather than an overall more optimal growth environment.

Structure and composition of the plant communities.-Extremely mineral--rich fen: Extremely mineral-rich fens, represented in this study by Kiser Lake Fen, have strong groundwater influence that is often artesian in nature. Because the water is under pressure, these "raised fens" exhibit peat that accumulates far above the average water table of the surroundings. In Kiser Lake Fen, peat deposits are slowly sprawling up nearby hillsides. The substrate, though quaking, is not treacherous. A similar raised peatland occurs in Randolph County (Friesner & Potzger 1946). Peat that accumulates in this type of wetland is often underlain by significant deposits of marl (Friesner & Potzger 1946; Swinehart & Parker 2000) or a peaty tufa (Stewart et al. 1994). Extremely mineral-rich fens in Indiana are characterized by a dense, low growth of Potentilla fruticosa, the fen counterpart of Chamaedaphne calyculata. The herbaceous flora is dominated by grasses, sedges, and plants common to prairies, such as Solidago and Rudbecki a. Parnassia glauca and Selaginella are frequently present and indicate extremely calcium-rich conditions. Common shrubs include Betula pumila, Cornus foemina, C. stolonifera, Salix pedicellaris, and Toxicodendron vernix. The bryophyte community is characterized by calcicolous species such as Amblystegium riparium and Fissidens adianthoides. A few isolated hummocks support Sphagnum fimbriatum and S. magellanicum. Chara (an alga) is common in pools of standing water.

Mineral-rich fen: This type of peatland, like the previous, has a strong groundwater influence and a low shrub component occupied by Potentilla fruticasa. It differs in that its substrate is more treacherous (usually floating), the groundwater is not under pressure, no marl is present in the sediment profiles, and extremely calcicolous species such as Parnassia glauca and Selaginella are absent. Scirpus acutus is a significant component of the herbaceous flora, and Typha is a common associate. Other typical herbaceous species include Drosera rotundifolia, Eleocharis spp., Osmunda regalis, Thelypteris palustris and Viola blanda. Menyanthes trifoliata, Sarracenia purpurea and Vaccinium macrocarpon are often present locally. Shrubs are occasional on these open fens. Toxicodendron vernix is by far the most abundant. Betula pumila, Cornus foemina, C. stolonifera, Rosa palustris, Salix pedicellaris and Spiraea tomentosa are also common. Larix laricina is present locally and usually is found around the periphery of the mat where the substrate is more stable. Closer to the upland, where the peat is grounded, trees of Acer rubrum, Betula allegheniensis and Populus tremuloides are common. The bryophyte community is characterized by Calliergonella cuspidata, Campylium polygamum and C. stellatum. Other common alkaliphilic bryophytes include Bryum pseudotriquetrum, Drepanocladus fluitans, Fissidens adianthoides, Isoptergyium elegans, and Mnium affine var. rugicum. Sphagnum forms isolated hummocks, and includes S. fimbriatum, S. fuscum, S. palustre, S. teres and S. warnstorfi. Sphagnum warnstorfi, recovered from Binkley Fen, is a new record for Indiana (Swinehart & Andrus 1998). Central ponds of open water are occupied by Nuphar advena and Nymphaea tuberosa. Submergent macrophytes include Ceratophyllum demersum and Potamogeton spp. Water-filled pools within the mat are frequented by Chara sp., Lemna minor, L. trisulca, Riccia fluitans, Utricularia intermedia and U. vulgaris.

Mineral-rich tall-shrub bog: Tall-shrub bogs of this type are relatively mineral-rich, and the substrate is extremely treacherous. They are distinguished from fens in that their surface has become dominated by Sphagnum. Where brown mosses once thrived, Sphagnum fimbriatum (characteristic), S. palustre, S. teres, and Aulacomnium pa lustre dominate. Many of the Sphagna form level carpets, while S. fimbriatum often forms hummocks around the stems of shrubs. Toxicodendron vernix forms dense stands and is nearly unavoidable when traversing these peatlands on foot, as it offers the only reliable support on the treacherous mat. Other common, characteristic shrubs include Acer rubrum, Cephalanthus occidentalis, Rosa palustris and Salix pedicellaris. Cornus foemina, C. stolonifera, Ilex verticillata, Salix candida, Spiraea tomentosa, and Vaccinium corymbosum are infrequent locally. The few trees that occur near the periphery of these bogs include Acer rubrum, Populus tremuloides and Quercus palustris. Herbaceous spec ies are quite variable, but usually include Dulichium arundinaceum, Hypericum virginicum and Thelypterus palustris. Calopogon pulchellus, Drosera rotundefolia, Menyanthes trifoliata, Osmunda regalis, Potentilla palustris, Sarracenia purpurea, Triglochin maritima and Vaccinium macrocarpon are present locally. Pools of water in the mat are common. They are occupied by Lemna minor, Potamogeton spp., Riccia fluitans, Utricularia intermedia and U. vulgaris.

Leatherleaf bog: The vegetation assemblages of leatherleaf bogs vary relatively little among different peatlands. All leatherleaf bogs are characterized by a low shrub cover of Chamaedaphne calyculata. Woodwardia virginica is also characteristic of these peatlands, often forming a distinct zone in wetter areas. In most leatherleaf bogs, cranberries (Vaccinium macrocarpon and V. oxycoccos) are associated with both Woodwardia and Chamaedaphne. Other herbaceous layer species common to nearly all leatherleaf bogs in Indiana include Andromeda glaucophylla, Carex oligosperma, C. rostrata, Decodon verticillatus, Drosera spp. (D. rotundifolia and/or D. intermedia), Dulichium arundinaceum, Hypericum virginicum, Sarracenia purpurea, Scirpus cyperinus and Vaccinium corymbosum. Shrubs are sparse on the mat, except for a few sites where dense "islands" of Vaccinium corymbosum are forming in dryer areas. Other characteristic shrubs found scattered on the mat include Acer rubrum, Aronia melanocarpa, Nemopanthus mucronatus and Toxicodendron vein ix. Cephalanthus occidentalis always dominates the standing waters of the lagg. Acer rubrum and Quercus palustris are infrequent as trees on the mat. Curiously, Larix laricina, a common component of leatherleaf bogs in Michigan and other Great Lakes and northern localities, was absent from all of the leatherleaf bogs studied. It does, however, occur in abundance at Pinhook Bog, a leatherleaf bog in LaPorte County, Indiana (Wilcox 1982). The surface of the leatherleaf bogs is characteristically level; and complexes of clearly defined hummocks and hollows, common in northern peatlands (see Vitt et al. 1975), are essentially absent. The Sphagna, rather than occurring as a gradient of species according to the hydrology and chemistry of hummocks and hollows, are more often represented by only a few species common to the extreme conditions. In animal trails and other low, wet depressions and pools, Sphagnum cuspidatum dominates. In drier areas and raised "hummocks," more drought-tolerant spec ies such as S. magellanicum and S. papillosum (and infrequently, S. capillifolium) dominate. These relatively dry areas usually harbor the xerophyte Polytrichum strictum. In more or less level "carpets," S. recurvum and S. bartlettiamum are most common; and S. fallax and S. isoviitae are occasional. Near the more mineral-rich waters of the lagg, S. centrale, S. fimbriatum, S. palustre and S. squarrosum may be present. Aulacomnium palustre is ubiquitous in leatherleaf bogs, and it was also found in all of the other peatlands with the exception of Binkley Fen.

Mineral-poor tall-shrub bog: These peatlands are characterized by a nearly impenetrable cover of shrubs. Vaccinium corymbosum always dominates, and Aronia melanocarpa and Nemopanthus mucronatus are subdominant. The vegetation indicates that these tall-shrub communities developed from leatherleaf bogs. Both representatives in this study (Blueberry and Thompson Bogs), have remnant, depauperate specimens of Chamaedaphne calyculata surviving in the few patches where shrubs have not crowded them into extinction. Woodwardia virginica, another characteristic leatherleaf bog species, is also remnant in these tall-shrub bogs. Other understory flora is variable and relatively scarce due to the dense shrub layer. The lagg waters, like leatherleaf bogs, are occupied almost exclusively by Cephalanthus occidentalis. The bryophyte flora, like some of the vascular plants, suggests that these peatlands are old leatherleaf bogs. Not only is the underlying peat composed of a thick stratum of Sphagnum (Swinehart & Parker 2000), but the present Sphagna are the same species that characterize leatherleaf bogs. In wet areas, S. cuspidatum is common. In moist areas, S. recurvum forms carpets, and low hummocks harbor S. magellanicum. A few other species, namely S. affine and S. palustre, indicate more shaded and mineral-rich conditions. Leucobryum albidum, common in moist, shaded areas, was found in Blueberry Bog and represents a new Indiana record (Swinehart & Andrus 1998).

Forested peatland: Forested peatlands are characterized by a canopy dominated by Acer rubrum. Larix laricina was present in all of the sites included in the present study, but many individuals were dead and most of the living specimens were etiolated and depauperate as a result of shading by A. rubrum. Other members common in the tree layer include Acer saccharinum, Prunus serotina, Quercus palustris and Ulmus americana. Betula allegheniensis, Fagus grandifolia, Fraxinus pennsylvanica, Liriodendron tulipifera, Nyssa sylvatica and Sassafras albidum were infrequent locally. Pinus strobus was a dominant member of the canopy at Mount Pleasant Bog, and the presence of individuals in both the shrub and herbaceous layers suggests that the species will persist well beyond the demise of Larix, which never reproduces in the understory of these forested peatlands. A single individual of P. strobus was also found at Ropchan Memorial Bog. The shrub canopy in forested peatlands is dominated by Acer rubrum, Aronia melanocar pa, Ilex verticillata, Nemopan thus mucronatus, Toxicodendron vernix and Vaccinium corymbosum. Amelanchier laevis, Lindera benzoin and Rosa palustris are more local. Other shrubs include Gaylussacia baccata and Nyssa sylvatica. The herbaceous flora is characterized by Bidens cernua, Maianthemum canadense, Osmunda cinnamomea and Trientalis borealis. Other notable but more local herbaceous species include Carex trisperma, Coptis trifolia, Cypripedium acaule, Dryopterus spinulosa, Isotria verticillata, Lycopodium lucidulum, L. obscurum, Mitchella repens and Rubus hispidus. Acer rubrum seedlings are always abundant, indicating the importance of this species in the long-term composition of the tree canopy. The most characteristic and abundant bryophyte of forested peatlands in Indiana is Pallavicinia lyellii. Although it often forms on decaying tamarack root-stocks, it also forms hummocks on exposed peat. The hummocks are composed almost entirely of the undecomposed tissue of dead individuals, while the surface is colonized by living individuals. The species is considered an acidophile (H.A. Crum, pers. commun.); and, consequently, it is usually found on peat having a pH between 3.5-4.5. Other common bryophytes include Mnium affine var. ciliare, Rhynchostegium serrulatum, Sphagnum affine, Tetraphis pellucida and Thuidium delicatulum. Additional species are more local in distribution (see Table 6).

Variability of the vascular plant communities.--Alkalinity (indicative of the magnitude of groundwater influence) and woody plant cover (indicative of seral stage in terms of substrate density and moisture saturation) explained 21.5% of the variability in the plant communities of the herbaceous layer.

Alkalinity: Canonical Correspondence Analysis (CCA) of herbaceous-vascular flora (composition and cover) distinctly separated fens (Binkley, Kiser and Svoboda) from all other peatlands on the basis of alkalinity and pH (Fig. 2). Leatherleaf bogs were tightly clustered at the mineral-poor end of the gradient, but the other peatland types were more loosely clustered (Fig. 2). This tight clustering is attributed to the fact that leatherleaf communities have become as acid and mineral-poor as climatic and edaphic conditions will allow, favoring plant assemblages of close compositional affinity. In the case of fens, acidification of the substrate by brown mosses, and continued separation from the effects of groundwater due to sedimentation, will cause some peatlands to progressively migrate to the mineral-poor end of the spectrum. Binkley Fen, already exhibiting large expanses of colonizing Sphagnum mosses, is an example of this process, hence its position on the lower end (relatively) of the alkalinity gradient ( Fig. 2). In contrast, Kiser Lake Fen is far removed from Binkley Fen because it is extremely mineral-rich due to an artesian water supply and the presence of marl under the peat. The tall-shrub bogs and forested peatlands, due to their variable origins (having arisen from either fens or bogs), are loosely clustered, each showing either mineral-rich or mineral-poor affinities according to their respective developmental pathways. This is supported by the stratigraphic and palaeoecological data presented by Swinehart & Parker (2000).

Woody plant cover: In addition to groundwater influence, seral position was a major source of variability in the herbaceous vegetation of the peatlands. Seral position in these lake-fill peatlands is indicated by the amount of shrub and tree cover. Growth of this woody vegetation is favored by build-up of sediment to the point where the surface becomes dryer and firm enough to support the weight of the plants. As a consequence of this drying, decomposition accelerates and increases nutrient availability (Kratz & Dewitt 1986). With the colonization of dense shrubs or a forest canopy, light becomes limiting for heliophilic species.

Forested peatlands were distinctly separated from the other peatlands (Fig. 2). The distinction between the herbaceous vegetation in the forested peatlands and the tall-shrub bogs suggests that either light and/or edaphic conditions were significantly different. The response of the bryophytes to forest and shrub cover, discussed below, would suggest that moisture was the primary factor distinguishing the herbaceous vegetation of these peatlands.

Variability of the bryophyte communities.--Alkalinity: The four types of peatland were not as clearly separated on the basis of the response of the bryophyte community to alkalinity (Fig. 2). The character of individual peatlands along the gradient was more apparent, suggesting that the bryophyte flora is a better indicator of trophic condition than the vascular flora. According to Crum (1988), vascular plants have a higher trophic plasticity than mosses and, therefore, mosses are better indicators of habitat.

Leatherleaf bogs were clustered tightly at the mineral-poor end of the gradient (Fig. 2). Bryophyte communities of tall-shrub peatlands and forested peatlands shared affinities with either mineral-rich or mineral-poor conditions, depending on their seral origin (fen or bog, respectively). Kiser Lake Fen was distinctly separated from the other fens and forested peatlands due to its extreme mineral-rich, artesian water flow.

Woody plant cover: Shrub cover was a relatively insignificant factor in the ordination of the peatland bryophyte communities ([r.sup.2] < 0.2). However, tree basal area distinguished the bryophyte communities of forested peatlands from all other peatland types. While thick tree cover may have influenced the bryophyte communities by limiting light, the presence of a forest infers a dryer substrate, and this may have been a more important factor. Many of the bryophyte communities of tall-shrub bogs, which exhibited nearly impenetrable shrub canopies, are closely associated with leatherleaf bogs in the ordination. Since light in these tall-shrub bogs is significantly reduced (as it is in forested peatlands), it might be concluded that the moisture saturation of the substrate is a more important factor in distinguishing the bryophyte communities of forested peatlands and tall-shrub bogs. The progressively firmer and dryer substrate that favored the establishment of trees also favored a transition from bog and fen bryophytes to lowland forest species.

Development of the hydrosere.--Analysis of the bathymetry (Swinehart 1997), stratigraphy and subfossil composition (Swinehart & Parker 2000) of peatlands in Indiana, coupled with the present floral survey of the existing representatives of the various types and stages of peatlands in the area, supports the following conclusions about their development: 1) basin-type peatlands in the southern Great Lakes region have a "monophyletic" origin as a lake or pond which, for various reasons, develops into a peat-forming fen, 2) subsequent to the open fen stage, the successional pathway becomes bi-directional; and depending on the morphometric and hydrologic characteristics of the basin (Swinehart 1997), some of the fens become mineral-poor leatherleaf bogs, while others do not, and 3) regardless of their inherent mineral richness, all Indiana peatlands are succeeded by a forest dominated by Acer rubrum. Open fens that were favorable to rapid, large-scale colonization by Sphagnum mosses developed into leatherleaf bogs (Swinehart 1997; Swinehart & Parker 2000). Leatherleaf bogs, as their substrate becomes more stable (and consequently less wet), are succeeded by a tall-shrub community characteristic of acid, relatively mineral-poor conditions (mainly Vaccinium corymbosum, Aronia melanocarpa and Nemopanthus mucronatus). Any Larix trees that pioneer the increasingly stable substrate are quickly displaced by Acer rubrum. The latter is better able to withstand the occasional flooding that results from grounding of the mat (Moizuk & Livingston 1966), and it is tolerant of shade in the understory. In contrast, Larix is shade intolerant and fares poorly in flooded conditions (Duncan 1954). Grounding of the mat favors decomposition of the surface of the peatland to the point where it nearly equals productivity, causing muck (rather than peat) to form. Nutrient availability is increased slightly, but the acid conditions, coupled with shade provided by the dense canopy of arborescent trees results in a sparse subcanopy cover of acid and shade tolerant herbs.

If morphometric and hydrologic characteristics of a fen basin are not favorable to rapid, large-scale colonization of the open mat by Sphagnum mosses, alkaliphilic shrubs invade centripetally as the substrate becomes progressively suitable for their growth. Sphagnum may accompany this invasion, but only as a peripheral ring or as isolated hummocks under the canopy of shrubs and small trees. Eventually, after the peat becomes dense enough to sufficiently reduce groundwater influence, Sphagnum may colonize the entire surface of the peatland. However, the substrate is already grounded, or nearly so, and the shrub canopy is well-developed. Shade from the shrubs and nutrients released from decomposition of the surface peat (due to grounding) will limit typical bog species and favor species typical of forested peatlands. Red maple establishes early and eventually forms a dense lowland forest (Swinehart & Parker 2000).

Unlike peatlands of the northern Great Lakes region and other northern localities in North America, conifers do not characterize the later stages of the peatland hydrosere in Indiana. In many cases, they are absent entirely, even in the early stages of peatland development. In Canada and northern Michigan, mineral-poor peatlands become dominated by Picea mariana and mineral-rich peatlands by Thuja occidentalis (Crum 1988). Picea mariana is not known from Indiana during historic times, and Thuja occidental is is extremely rare and found only near Lake Michigan (Deam 1940).

Acer rubrum is the tentative climax species in Indiana peatlands, in southwest Michigan (Crow 1969a), and northeast Ohio (Andreas & Bryan 1990). It is dominant in the canopy and prolific in the understory. A graphic representation of the size class distribution of Acer rubrum trees in forested peatlands shows a classic reverse "J"-shaped curve, typical of species destined to replace themselves in their own shade. The transient nature of pioneer species such as Betula allegheniensis and Larix laricina is indicated by a bell-shaped distribution representing the eventual extinction of the species for lack of seedling and sucker survival due to shade.

The dominance of red maple in the later stages of peatland development in the southern Great Lakes region, unlike that of black spruce in the North, can be attributed to competitive success. Red maple is not an opportunist that exists in these peatlands due to uncontested survival (as in black spruce of the northern Great Lakes region (Crum 1988)), but rather it thrives and out-competes potential competitors such as Quercus palustris. Whether red maple is truly the ultimate edaphic climax species for the peatland hydrosere of the region is uncertain. If an equilibrium of productivity and decomposition exists in grounded peat, as suggested by Kratz & DeWitt (1986), the structure and composition of the vegetation, barring any major hydrological or elevation change or other major disturbance, would likely remain steady--favoring the continued existence of a red maple-dominated lowland forest. If, however, a slight surplus of productivity occurred as indicated by Swinehart & Parker (2000), mucky soil might eventu ally accumulate high enough to become relatively dry or mesic. If this happened, the existence of a complete hydrosere from open water to a forest of terrestrial affinity is not impossible. Mount Pleasant Bog (Laporte County) may be an early example of this. The substrate is firm and dry in most places, and Pinus strobus grows to maturity and reproduces successfully. Large specimens of Fagus grandifolia, the dominant species occupying the surrounding terrestrial forest, are already occasional in the pine forest.

A definitive conclusion about the nature of the steady-state or climax community in peatlands of glaciated regions is, in the final analysis, speculative due to insufficient time for full development of these communities. Perhaps application of the term "old growth" could be employed in much the same way that it is used to describe steady-state/climax upland forests. Parker (1989) defines mesic old growth forests as possessing: 1) an overstory canopy of trees older than 150 years and with little or no anthropogenic understory disturbance during the past 80-100 years, 2) an all-aged structure with multi-layered canopies, 3) horizontal structure with aggregations of canopy trees interspersed with small, all-aged canopy gaps of varying species composition, and 4) significant numbers of standing, dead snags and downed logs. The red maple peatlands meet all of these criteria accept the first. The oxygen profile in the peatland allows only the top 20-30 cm for root growth. Consequently, wind-throws are extremely co mmon, especially large (older) trees, and the maximum age (and size) of trees is limited by the very nature of the substrate.

Regional identity of leatherleaf bog cornmunities.--A comparison of the vegetation of leatherleaf bogs of different regions was facilitated by: 1) relative intraregional homo geneity, 2) by the large amount of published data from different regions, and 3) a common variable in all of the studies that allowed standardization of the data for analysis. Scarcity of similar quantities of appropriate regional data on non-Sphagnum-dominated fens prevented statistical comparison of fens.

Indirect gradient analysis (Detrended Correspondence Analysis (DCA)) of vegetation frequency (ranked) from 27 leatherleaf bogs in the eastern United States (Table 8) showed three distinct clusters: those from southern Michigan and northern Indiana, those from northern Michigan, and those from southern New York, northern New Jersey and northeast Ohio (Fig. 3). Direct gradient analysis (Canonical Correspondence Analysis (CCA)) of the vegetation and five climatic variables showed a similar clustering; however, the Ohio peatlands were more distinctly separated from the northeastern sites, and the New Hampshire bog (Pequawket Bog) associated more closely with the southern Great Lakes sites (Fig. 3).

Much of the variability separating the "southern peatlands" (southern Michigan, northern Indiana, northeast Ohio, southern New York, northern New Jersey and New Hampshire) from the "northern peatlands" (represented by northern Michigan) is explained by mean annual temperature and mean annual precipitation (both having [r.sup.2] values > 0.8). The climate surrounding the northern peatlands is colder and dryer on average. Peatlands in southern Michigan and northern Indiana were further distinguished from the northern sites by having much hotter mean July temperatures. The number of frost-free days was different between bogs from the southern Great Lakes region and bogs from the northeastern U.S.; however, this variable was relatively insignificant with an [r.sup.2] value of < 0.2.

Analysis of the frequency of the most prevalent leatherleaf bog species from all of the regions and sites listed in Table 8 revealed seven geographic groupings: 1) species common to all leatherleaf bogs in the eastern United States, 2) species characteristic of northern peatlands only, 3) species restricted to southern peatlands, 4) species found in most regions except for the Northeast, 5) species found in the north and northeast but absent from the southern great lakes region, 6) species found only in the Northeast, and 7) species exclusive to peatlands of the southern Great Lakes region (Table 9). Species such as Carex canescens and Peltandra virginica demonstrated the northeastern affinity of Ohio peatlands, whereas Betula pumila and Carex oligosperma showed the northern Great Lakes affinity of leatherleaf bogs in Indiana and southern Michigan.

The results of the ordinations and species groupings suggest that not only are the leatherleaf bog assemblages of the North different from those at the southern limit of the community (as concluded by Lynn & Karlin 1985), but the southern populations also have specific regional affinities. The differences between the southern sites cannot be entirely attributed to climatic variation. Some of the differences must be attributable to biogeographic history as it relates to the various lobes of ice of Wisconsin Age glaciation (see Crow 1969b).

CONCLUSIONS

Systematic vegetation sampling of 16 Indiana peatlands, including fens, tall-shrub bogs, leatherleaf bogs and forested peatlands yielded a total of 181 species. The number of species found at each site was variable, and no significant difference in species richness was noted between fens and bogs. A significant portion of the variability of the vascular and bryophyte vegetation was attributed to groundwater influence as indicated by alkalinity. Fens and bogs were distinctly separated on the basis of alkalinity. In addition to abiotic factors, temporal variables such as shrub and tree cover (indirectly indicative of seral stage, soil moisture and substrate stability) explained a significant portion of the variation. The vascular vegetation of tall-shrub bogs and forested peatlands was distinctly separated from the other sites. In contrast, shrub cover was an insignificant factor in bryophyte variability, although tree cover (Basal Area) was significant. It was thus concluded that substrate moisture rather than light accounted for most of the temporal variability in bryophytes.

Analysis of extant vegetation, along with the consideration of the palaeoecological data presented by Swinehart & Parker (2000), revealed an apparent hydrosere that begins with a lake or pond and ends with a forested peatland. All peatlands have a fen stage, but only some develop into bogs before becoming forested. Regardless of the inherent mineral-richness, all Indiana peatlands become dominated by red maple. Red maple may be the "climax" species for the peatland hydrosere in Indiana, but insufficient time has passed for a definitive conclusion. "Old-growth" might be a better descriptive term for these latter stages of peatland development.

The vascular vegetation communities of Indiana peatlands and other peatlands in the southern Great Lakes region are distinct from those in the northeast, Ohio and the northern Great Lakes. Some of this distinction is attributable to climatic factors, while others are related to biogeographic history of the region.

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[Figure 2 omitted]
Table 1

Climate variables for several regions of eastern North America,
including northwest and northeast Indiana.

(*)Data from Schaal (1966).

(**)Data from NOAA (1974) for Charlottburg.


                                              Northern      New
                                  Ontario     New York   Hampshire
                                  (Sims et   (Andrus et   (Fahey &
                                 al. 1982)   al. 1983)   Crow 1995)

Mean annual precip. (cm)             75        100         116.9
Mean annual snowfall (cm)            --        190         287.8
Mean annual temp. ([degrees]C)       -l          --          6.3
Mean January temp. ([degrees]C)     -20         -9.3        -8.8
Mean July temp. ([degrees]C)         15         18.4        19.7
Avg. frost-free days             [+ or -]90    105        105-130

                                   Northern       Northern
                                    lower        New Jersey
                                   Michigan        (Lynn &
                                 (Gates 1942)   Karlin 1985)

Mean annual precip. (cm)             70.6           110
Mean annual snowfall (cm)           157              --
Mean annual temp. ([degrees]C)        5.7             9.8 (**)
Mean January temp. ([degrees]C)      -7.2            -2
Mean July temp. ([degrees]C)         20              22
Avg. frost-free days             [+ or -]138   [+ or -]l40-170

                                                Northeast
                                  Southwest       Ohio
                                   Michigan    (Andreas &
                                 (Crow 1969a)  Bryan 1990)

Mean annual precip. (cm)             83.8         103.8
Mean annual snowfall (cm)           101           154.7
Mean annual temp. ([degrees]C)        9.2           9.6
Mean January temp. ([degrees]C)     -11.4          -3.5
Mean July temp. ([degrees]C)         29.5          21.6
Avg. frost-free days                150            165

                                    Northeast        Northeast
                                     Indiana          Indiana
                                    (LaPorte)        (Albion)
                                   (NOAA 1974)      (NOAA 1974)

Mean annual precip. (cm)             127               83
Mean annual snowfall (cm)          75-125 (*)       75-100 (*)
Mean annual temp. ([degrees]C)         9.9              9.6
Mean January temp. ([degrees]C)       -3.4             -3.4
Mean July temp. ([degrees]C)          23.6             22.9
Avg. frost-free days             [+ or -]150 (*)  [+ or -]150 (*)
Table 2

Mean values (n = 3) for pH, conductivity, alkalinity, and hardness from
interstitial waters of 16 Indiana peatlands. Acronyms in parentheses
correspond with Tables 3, 4, 5, and 6.

(*)Exact locations available from the Indiana Department of Natural
Resources, Division of Nature Preserves.



                                       Township/        Conductivity
Name of peatland (*)        County     Range (*)   pH    ([mu]MHOS)

Binkley Fen (BF)            Steuben    T38N, R13E  6.5      238
Kaiser Lake Fen (KLF)       Kosciusko  T33N, R7E   6.9      606
Svoboda Fea (SVF)           Noble      T35N, R11E  6.5      371
Blueberry Bog (BLB)         Elkhart    T38N, R7E   3.5       68
Hickory Bog (HB)            Noble      T33N, R8E   4.7       27
Little Chapman Bog (LCB)    Kosciusko  T33N, R6E   6.0       95
Thompson Bog (THB)          LaPorte    T38N, R1W   3.4       60
Burket Bog (BUR)            Kosciusko  T32N, R5E   3.9       44
Dutch Street Bog (DSB)      Noble      T35N, R10E  3.7       79
Leatherleaf Bog (LB)        Noble      T33N, R8E   4.7       48
Shoemaker Bog (SHB)         LaPorte    T36N, R3W   3.7       51
Yost Bog (YB)               Lagrange   T38N, R8E   3.2       96
Little Arethusa Bog (LAB)   Kosciusko  T31N, R7E   6.2      146
Mount Pleasant Bog (MPB)    LaPorte    T38N, R2W   3.9       75
Ropchan Memorial Bog (RMB)  Steuben    T38N, R12E  4.4       84
Tamarack Bog (TAM)          Noble      T33N, R9E   3.9       63

                              Total     Calcium   Magnesium
                            alkalinity  hardness  hardness
Name of peatland (*)          (ppm)      (ppm)      (ppm)

Binkley Fen (BF)               120       128         12
Kaiser Lake Fen (KLF)          340       280         48
Svoboda Fea (SVF)              180       160         40
Blueberry Bog (BLB)             20         8          0
Hickory Bog (HB)                30         8          0
Little Chapman Bog (LCB)        80        60          0
Thompson Bog (THB)              20         8          0
Burket Bog (BUR)                12         8          0
Dutch Street Bog (DSB)          12        20          0
Leatherleaf Bog (LB)            20         8          0
Shoemaker Bog (SHB)             20         8          0
Yost Bog (YB)                   12         8          0
Little Arethusa Bog (LAB)       64        56          0
Mount Pleasant Bog (MPB)        12         8          0
Ropchan Memorial Bog (RMB)      20         8          0
Tamarack Bog (TAM)              12         8          0
Table 3

Tree layer vegetation in 16 Indiana peatlands. The first number in each
cell is the percent frequency, the second number is the density per
hectare, the third number is the basal area per hectare, and the fourth
number is the importance value for each species in the respective
peatlands. Peatland acronyms correspond with those listed in Table 2. A
"P" means the species was present as trees, but occurred outside of the
sampling quadrats. "ND" means the species was present, but no numerical
data is available. (*) Data from Swinehart (1994).

                                          Fens
Species                     KLF                SVF

Acer rubrum              --                 --
Acer saccharinum         --                 --
Amelanchier laevis       --                 --
Aronia melanocarpa       --                 --
Betula allegheniensis    --                 P
Fagus gradifolia         --                 --
Fraxinus pennsylvanica   --                 --
Juniperus communis       3, 0.83, 0.004, 8  --
Larix laricina           --                 14, 5.5, 0.02, 100
Liriodendron tulipifera  --                 --
Nyssa sylvatica          --                 --
Pinus strobus            --                 --
Populus tremuloides      3, 0.83, 0.002, 8  --
Prumus serotina          6, 1.7, 0.004, 16  --
Quercus palustris        --                 --
Salix pedicellaris       3, 1.7, 0.007, 12  --
Salix sp.                3, 1.7, 0.07, 23   --
Sassafras albidum        --                 --
Toxicodendron vernix     6, 1.7, 0.007, 16  --
Ulmus americana          --                 --

                                Fens                 Tall-Shrub bogs
Species                     BF                 LCB             HB

Acer rubrum              --                 --                 --
Acer saccharinum         --                 --                 --
Amelanchier laevis       --                 --                 --
Aronia melanocarpa       --                 --                 --
Betula allegheniensis    --                 --                 --
Fagus gradifolia         --                 --                 --
Fraxinus pennsylvanica   --                 --                 --
Juniperus communis       --                 --                 --
Larix laricina           4, 0.9, 0.003, 50  --                 --
Liriodendron tulipifera  --                 --                 --
Nyssa sylvatica          --                 --                 --
Pinus strobus            --                 --                 --
Populus tremuloides      4, 0.9, 0.003, 50  26, 13, 0.11, 63   --
Prumus serotina          --                 --                 --
Quercus palustris        --                 17, 6.5, 0.02, 27  --
Salix pedicellaris       --                 --                 --
Salix sp.                --                 --                 --
Sassafras albidum        --                 --                 --
Toxicodendron vernix     --                 --                 --
Ulmus americana          --                 --                 --

                                 Tall-Shrub bogs
Species                     THB               BLB

Acer rubrum              100, 125, 0.41, 33   P
Acer saccharinum         --                   --
Amelanchier laevis       --                   --
Aronia melanocarpa       --                   --
Betula allegheniensis    33, 8.3, 0.08, 6     --
Fagus gradifolia         --                   --
Fraxinus pennsylvanica   --                   --
Juniperus communis       --                   --
Larix laricina           100, 141.7, 1.8, 55  --
Liriodendron tulipifera  --                   --
Nyssa sylvatica          --                   --
Pinus strobus            --                   --
Populus tremuloides      33, 8.3, 0.04, 6     --
Prumus serotina          --                   --
Quercus palustris        --                   --
Salix pedicellaris       --                   --
Salix sp.                --                   --
Sassafras albidum        --                   --
Toxicodendron vernix     --                   --
Ulmus americana          --                   --

                                     Leatherleaf bogs
Species                     DSB                YB

Acer rubrum              64, 122, 1.5, 98    55, 72.5, 0.54, 83
Acer saccharinum         --                  --
Amelanchier laevis       --                  --
Aronia melanocarpa       --                  --
Betula allegheniensis    --                  --
Fagus gradifolia         --                  --
Fraxinus pennsylvanica   --                  9, 1.1, 0.01, 8
Juniperus communis       --                  --
Larix laricina           --                  --
Liriodendron tulipifera  --                  --
Nyssa sylvatica          2, 0.5, 0.002, 1    --
Pinus strobus            --                  --
Populus tremuloides      --                  --
Prumus serotina          --                  --
Quercus palustris        --                  10, 8.7, 0.09, 17
Salix pedicellaris       --                  --
Salix sp.                --                  --
Sassafras albidum        --                  --
Toxicodendron vernix     --                  --
Ulmus americana          --                  --

                                         Leatherleaf bogs
Species                     BUR               SH               LB

Acer rubrum              26, 24, 0.14, 84  3, 0.78, 0.002, 50  --
Acer saccharinum         4, 3.7, 0.04, 16  3, 0.78, 0.002, 50  --
Amelanchier laevis       --                --                  --
Aronia melanocarpa       --                --                  --
Betula allegheniensis    --                --                  --
Fagus gradifolia         --                --                  --
Fraxinus pennsylvanica   --                --                  --
Juniperus communis       --                --                  --
Larix laricina           --                --                  --
Liriodendron tulipifera  --                --                  --
Nyssa sylvatica          --                --                  --
Pinus strobus            --                --                  --
Populus tremuloides      --                --                  --
Prumus serotina          --                --                  --
Quercus palustris        --                --                  --
Salix pedicellaris       --                --                  --
Salix sp.                --                --                  --
Sassafras albidum        --                --                  --
Toxicodendron vernix     --                --                  --
Ulmus americana          --                --                  --

                                        Forested peatlands
Species                     LAB                MPB             TAM (*)

Acer rubrum              100, 245, 7.8, 36  86, 277, 8.3, 50   ND
Acer saccharinum         64, 63.6, 1.7, 11  36, 19.6, 1.1, 12  ND
Amelanchier laevis       27, 6.8, 0.05, 2   --                 ND
Aronia melanocarpa       9, 2.3, 0.004, <1  --                 --
Betula allegheniensis    --                 --                 --
Fagus gradifolia         --                 P                  --
Fraxinus pennsylvanica   27, 6.8, 0.24, 3   --                 --
Juniperus communis       --                 --                 --
Larix laricina           91, 313, 1.4, 22   64, 41.1, 0.4, 11  P
Liriodendron tulipifera  9, 2.3, 0.02, <1   --                 --
Nyssa sylvatica          --                 14, 3.6, 0.1, 2    --
Pinus strobus            --                 57, 76.8, 4.9, 18  --
Populus tremuloides      --                 --                 --
Prumus serotina          9, 2.3, 0.02, <1   --                 ND
Quercus palustris        --                 7, 1.8, 1.1, 1     ND
Salix pedicellaris       --                 --                 --
Salix sp.                18, 9.1, 0.05, 4   --                 --
Sassafras albidum        --                 --                 ND
Toxicodendron vernix     45, 18.2, 0.05, 4  --                 --
Ulmus americana          100, 156, 1.7, 17  14, 3.6, 0.1, 2    ND

                         Forested peatlands
Species                     RMB

Acer rubrum              100, 485, 4, 45
Acer saccharinum         17, 8.3, 0.09, 2
Amelanchier laevis       --
Aronia melanocarpa       --
Betula allegheniensis    100, 171, 1.3, 21
Fagus gradifolia         --
Fraxinus pennsylvanica   --
Juniperus communis       --
Larix laricina           75, 135, 2.3, 22
Liriodendron tulipifera  --
Nyssa sylvatica          --
Pinus strobus            P
Populus tremuloides      --
Prumus serotina          --
Quercus palustris        --
Salix pedicellaris       --
Salix sp.                8, 2.1, 0.14, 1
Sassafras albidum        --
Toxicodendron vernix     33, 10.4, 0.02, 4
Ulmus americana          25, 18.7, 0.12, 4
Table 4

Shrub layer vegetation in 16 peatlands in Indiana. The first number in
each cell is the percent frequency for each species, the second number
is the percent cover, and the third number is the importance value for
each species in the respective peatlands. Peatland acronyms correspond
with Table 2. A "P" means the species was present as shrubs, but
occurred outside of the sampling quadrats.

                                              Fens

Species                     KLF           SVF          BF

Acer rubrum                 --            7, 0.11, 2   22, 1, 13
Amelanchier leavis          --            --           --
Aronia melanocarpa          --            --           --
Betula alleghenicnsis       3, 0.03, 1    --           --
Bentula pumila              10, 0.23, 3   17, 3.3, 13  4, 0.11, 2
Carpinus Caroliniana        --            --           --
Cephalanthus Occidentalis   --            --           --
Chamaedaphne calyculata     --            --           --
Cornus foemina              20, 4.3, 11   --           4, 0.37, 3
Cornus stolonifera          93, 23.8, 58  34, 3.2, 18  --
Decodon verticillatus       --            --           --
Gaylussacia baccata         --            --           --
Ilex verticillata           --            --           11, 0.85, 8
Larix laricina              --            7,0, 34, 3   48, 3.4, 34
Lindera benzion             --            --           --
Nemopanthus mucronatus      --            --           --
Nyssa sylvatica             --            --           --
Parthenocissu quinquefolia  --            --           --
Pinus strobus               --            --           --
Populus tremuloides         13, 1.2, 5    --           --
Potentilla fruticosa        3, 0.33, 1    --           --
Prunus serotina             --            --           --
Quercus palustris           --            --           --
Rosa palustris              7, 0.33, 2    7, 0.38, 3   4, 0.04, 1
Salix candida               --            --           --
Salix discolor              --            3, 0.17, 1   --
Salix lucida                --            --           4, 0.11, 2
Salix pedicellaris          20, 1.9, 8    3, 0.17, 1   --
Salix sericiea              7, 0.83, 3    --           --
Salix serrissima            --            38, 2.3, 16  --
Salix sp.                   --            --           --
Soirea tomentosa            --            10, 0.55, 4  11, 0.18, 5
Toxicodendron vernix        23, 0.93, 7   58, 8.3, 38  48, 3, 32
Toxicodendron radicans      --            --           --
Ulmus americana             --            --           --
Vaccinium corymbosum        --            --           --
Viburnum lentago            --            --           P
Vitis riparia               3, 0.33, 1    --           --

                                        Tall-shrub bogs

Species                     LCB            HB        THB

Acer rubrum                 17, 0.74, 4    P         100, 15, 17
Amelanchier leavis          --             --        --
Aronia melanocarpa          4.0, 04, 1     --        100, 15, 17
Betula alleghenicnsis       --             --        33, 0.3, 3
Bentula pumila              --             --        --
Carpinus Caroliniana        --             --        --
Cephalanthus Occidentalis   --             29, 3, 4  33, 1.7, 4
Chamaedaphne calyculata     --             --        --
Cornus foemina              9, 0.48, 2     --        --
Cornus stolonifera          4, 0.22, 1     --        --
Decodon verticillatus       --             --        P
Gaylussacia baccata         --             --        --
Ilex verticillata           --             14, 5, 3  --
Larix laricina              --             --        --
Lindera benzion             --             --        --
Nemopanthus mucronatus      --             --        67, 16.7, 15
Nyssa sylvatica             --             --        --
Parthenocissu quinquefolia  --             --        --
Pinus strobus               --             --        --
Populus tremuloides         13, 0.30, 3    --        33, 1.7, 4
Potentilla fruticosa        --             --        --
Prunus serotina             --             --        --
Quercus palustris           26, 2.7, 8     --        --
Rosa palustris              13, 0.69, 3    P         P
Salix candida               43, 3.1, 11    --        --
Salix discolor              --             --        --
Salix lucida                --             --        --
Salix pedicellaris          26, 1.3. 6     --        --
Salix sericiea              --             --        --
Salix serrissima            --             --        --
Salix sp.                   --             P         --
Soirea tomentosa            4, 0.04, 1     --        P
Toxicodendron vernix        100, 38.7, 58  P         33, 3.3, 5
Toxicodendron radicans      --             --        --
Ulmus americana             --             --        --
Vaccinium corymbosum        9, 0.65, 2     --        100, 46.7, 33
Viburnum lentago            --             --        --
Vitis riparia               --             --        --

                             Tall-shrub        Leatherleaf bogs
                                bogs

Species                     BLB          DSB         LB

Acer rubrum                 --           42, 2       P
Amelanchier leavis          --           --          --
Aronia melanocarpa          80, 14, 20   53, 14, 22  --
Betula alleghenicnsis       --           --          --
Bentula pumila              --           --          --
Carpinus Caroliniana        --           --          --
Cephalanthus Occidentalis   40, 19, 16   42, 17, 22  1, 1, <1
Chamaedaphne calyculata     20, 1, 4     --          --
Cornus foemina              --           --          --
Cornus stolonifera          --           --          --
Decodon verticillatus       --           --          --
Gaylussacia baccata         20, 1, 4     --          --
Ilex verticillata           40, 12, 12   --          --
Larix laricina              --           --          --
Lindera benzion             --           --          --
Nemopanthus mucronatus      20, 2, 4     24, 3, 8    --
Nyssa sylvatica             --           --          --
Parthenocissu quinquefolia  --           --          --
Pinus strobus               --           --          --
Populus tremuloides         --           --          --
Potentilla fruticosa        --           --          --
Prunus serotina             --           --          --
Quercus palustris           --           --          --
Rosa palustris              --           --          P
Salix candida               --           --          --
Salix discolor              --           --          --
Salix lucida                --           --          --
Salix pedicellaris          --           --          --
Salix sericiea              --           --          --
Salix serrissima            --           --          --
Salix sp.                   --           --          --
Soirea tomentosa            --           --          3, <1, 1
Toxicodendron vernix        --           11, 0.3, 2  --
Toxicodendron radicans      --           --          --
Ulmus americana             --           --          --
Vaccinium corymbosum        100, 51, 41  80, 21, 34  P
Viburnum lentago            --           --          --
Vitis riparia               --           --          --

                                         Leatherleaf bogs

Species                     YB             BUR          SH

Acer rubrum                 65, 4.5, 15    55, 2.8, 23  3, 0.03, 2
Amelanchier leavis          --             --           --
Aronia melanocarpa          35, 2.9, 8     33, 1.7, 14  3, 0.47, 5
Betula alleghenicnsis       --             --           --
Bentula pumila              --             --           --
Carpinus Caroliniana        --             --           --
Cephalanthus Occidentalis   25, 6.4, 11    37, 1.5, 14  37, 3.3, 42
Chamaedaphne calyculata     5, 0.5, 1      --           --
Cornus foemina              --             --           --
Cornus stolonifera          --             --           --
Decodon verticillatus       --             --           16, 2.9, 27
Gaylussacia baccata         --             4, 0.74, 4   --
Ilex verticillata           45, 4, 11      11, 1.8, 9   9, 0.78, 10
Larix laricina              --             --           --
Lindera benzion             --             --           --
Nemopanthus mucronatus      5, 0.5, 1      22, 1.5, 11  --
Nyssa sylvatica             --             --           --
Parthenocissu quinquefolia  --             --           --
Pinus strobus               --             --           --
Populus tremuloides         --             --           --
Potentilla fruticosa        --             --           --
Prunus serotina             --             --           --
Quercus palustris           10, 0.3, 2     --           --
Rosa palustris              5, 0.5, 1      11, 0.18, 3  --
Salix candida               --             --           --
Salix discolor              --             --           --
Salix lucida                --             --           --
Salix pedicellaris          --             --           --
Salix sericiea              --             --           --
Salix serrissima            --             --           --
Salix sp.                   --             --           --
Soirea tomentosa            5, 0.05, 1     --           --
Toxicodendron vernix        100, 18.2, 36  37, 2.5, 18  16, 0.84, 14
Toxicodendron radicans      --             --           --
Ulmus americana             --             --           --
Vaccinium corymbosum        50, 4.7, 13    7, 0.55, 4   --
Viburnum lentago            --             --           --
Vitis riparia               --             --           --

                                       Forested peatlands

Species                     LAB            MPB           TAM (*)

Acer rubrum                 45, 3.6, 6     36, 3.8, 11   3, 1, 6
Amelanchier leavis          18, 0.91, 2    --            P
Aronia melanocarpa          --             14, 0.43, 3   11, 1, 10
Betula alleghenicnsis       --             --            --
Bentula pumila              --             --            --
Carpinus Caroliniana        9, 0.09, 1     --            --
Cephalanthus Occidentalis   --             7,0.21, 2     --
Chamaedaphne calyculata     --             --            --
Cornus foemina              9, 0.27, 1     --            --
Cornus stolonifera          P              --            --
Decodon verticillatus       --             --            --
Gaylussacia baccata         --             --            --
Ilex verticillata           82, 17.3, 18   7, 0.07, 2    16, 3, 21
Larix laricina              27, 3.6, 5     --            --
Lindera benzion             54, 5.9, 8     --            13, 2, 15
Nemopanthus mucronatus      9, 0.91, 1     --            P
Nyssa sylvatica             --             86, 40.7, 52  --
Parthenocissu quinquefolia  18, 0.73, 2    --            --
Pinus strobus               --             21, 0.64, 5   --
Populus tremuloides         --             --            --
Potentilla fruticosa        --             --            --
Prunus serotina             18, 0.91, 2    --            11, 1, 10
Quercus palustris           --             --            --
Rosa palustris              45, 2.4, 5     P             --
Salix candida               --             --            --
Salix discolor              --             --            --
Salix lucida                --             --            --
Salix pedicellaris          --             --            --
Salix sericiea              --             --            --
Salix serrissima            --             --            --
Salix sp.                   --             --            --
Soirea tomentosa            --             P             --
Toxicodendron vernix        100, 15.9, 19  7, 2.9, 4     5, <1, 6
Toxicodendron radicans      9, 0.45, 1     --            --
Ulmus americana             54, 4.1, 7     --            --
Vaccinium corymbosum        100,18.6, 21   64, 10.4, 22  21, 5, 31
Viburnum lentago            --             --            --
Vitis riparia               9, 0.09, 1     --            --

                               Forested
                               peatlands

Species                     RMB

Acer rubrum                 83, 18.2, 23
Amelanchier leavis          --
Aronia melanocarpa          P
Betula alleghenicnsis       42, 4.6 9
Bentula pumila              --
Carpinus Caroliniana        --
Cephalanthus Occidentalis   --
Chamaedaphne calyculata     --
Cornus foemina              --
Cornus stolonifera          --
Decodon verticillatus       --
Gaylussacia baccata         17, 1.7, 3
Ilex verticillata           100, 28.3, 32
Larix laricina              --
Lindera benzion             --
Nemopanthus mucronatus      33, 5, 8
Nyssa sylvatica             --
Parthenocissu quinquefolia  --
Pinus strobus               --
Populus tremuloides         --
Potentilla fruticosa        --
Prunus serotina             --
Quercus palustris           --
Rosa palustris              --
Salix candida               --
Salix discolor              --
Salix lucida                --
Salix pedicellaris          --
Salix sericiea              --
Salix serrissima            --
Salix sp.                   --
Soirea tomentosa            --
Toxicodendron vernix        33, 4.2, 7
Toxicodendron radicans      --
Ulmus americana             --
Vaccinium corymbosum        75, 12.1, 18
Viburnum lentago            --
Vitis riparia               --

(*)Data from Swinehart (1994).
Table 5

Herbaceous layer vegetation in 16 peatlands in northern Indiana. The
first number in each cell is the percent frequency for each species, the
second number is the percent cover, and the third number is the
importance value for each species in the respective peatlands. Peatland
acronyms correspond with Table 2. A "P" means the species was present in
the herbaceous layer, but occurred outside of the sampling quadrats. (*)
Data from Swinehart (1994).

                                               Fens
      Species                    KLF          SVF           BF

Acer rubrum                  --           3, 0.03, <1  7, 0.07, <1
Alisma subcordata            --           --           --
Andromeda glaucophylla       --           --           P
Aronia melanocarpa           --           --           --
Betula allegheniensis        --           --           --
Betula pumila                3, 0.17, 1   --           7, 0.3, 1
Bidens sp.                   3, 0.17, 1   --           11, 0.11, 1
Calla palustris              --           --           --
Calopogon pulchellus         --           --           --
Caltha palustris             --           P            --
Cardamine bulbosa            P            --           --
Cardamine pratensis          --           31, 0.45, 4  --
Carex crinita                --           --           --
Carex lasiocarpa             --           3, 0.34, 1   --
Carex oligosperma            --           --           --
Carex rostrata               --           --           --
Carex sp.                    80, 53, 45   --           --
Carex trisperma              --           --           --
Cephalanthus occidentalis    --           --           --
Chamaedaphne calyculata      --           --           --
Coptis trifolia              --           --           --
Cornus foemina               7, 0.33, 1   --           4.0.04, <1
Cornus stolonifera           13, 0.57, 3  7, 0.86, 1   7.0.22, I
Cuscuta sp.                  --           --           --
Cypripedium acaule           --           --           --
Decodon verticillatus        --           --           P
Drosera intermedia           --           --           --
Drosera rotundifolia         -            17, 0.17, 2  P
Dryopteris spinulosa         --           --           --
Dulichium arundinaceum       --           --           --
Eleocharis sp.               --           10, 1.2, 2   78, 13.2, 16
Equisetum fluviatile         --           --           --
Equisetum palustris          13, 0.27, 2  --           --
Eriophorum virginicum        --           --           P
Eupatorium maculatum         --           P            P
Eupatorium perfoliatum       --           --           --
Galium sp.                   --           --           --
Gaylussacia baccata          --           --           --
Hypericum virginicum         --           --           33, 0.85, 3
Ilex verticillata            --           --           4, 0.11, <1
Impatiens capensis           17, 1.6, 4   14, 0.59, 2  --
Iris versicolor              --           --           --
Isotria verticillata         --           --           --
Juncus sp.                   --           --           11,0.18, 1
Leersia oryzoides            --           --           --
Lindera benzoin              --           --           --
Liparis loesellii            --           --           --
Liriodendron tulipifera      --           --           --
Lycopodium lucidulum         --           --           --
Lycopodium obscurum          --           --           --
Lysimachia terrestris        --           --           --
Lysimachia thrysiflora       --           --           --
Mainanthemum canadense       --           --           --
Menyanthes trifoliata        --           --           4, 0.92, 1
Mitchella repens             --           --           --
Nuphar advena                --           P            4, 0.37, 1
Nymphaea odorata             --           --           --
Nyssa sylvatica              --           --           --
Onoclea sensibilis           --           --           --
Osmunda cinnamomea           --           --           --
Osmunda regalis              --           7, 1.7, 2    44, 13, 13
Parnassia glauca             P            --           --
Parthenocissus quinquefolia  3, 0.03, 1   --           --
Pinus strobus                --           --           --
Poaceae (unidentified)       10, 1.8, 3   --           4, 0.37, 1
Polygonum sp.                --           --           --
Populus tremuloides          --           --           --
Potentilla anserina          --           --           --
Potentilla fruticosa         60, 20, 22   27, 7.8, 9   48, 4.4, 7
potentilla palustris         --           --           15, 0.59, 2
Prunus serotina              --           --           --
Quereus palustris            --           --           --
Quercus rubra                --           --           --
Ranunculus sp.               3.0, 17, 1   --           --
Rosa palustris               3.0, 33, 1   --           18, 1, 2
Rubus allegheniensis         3.0, 17, 1   --           --
Rubus hispidus               --           10, 0.9, 2   --
Rudheckia sp.                10, 0.10, 2  --           --
Rumex sp.                    --           P            --
Saggitaria latifolia         --           --           4, 0.04, <1
Salixcandida                 --           --           --
Salix pedicellaris           --           7, 0.72, 1   4, 0.04, <1
Salix serrissima             --           7, 0.38, 1   --
Sarracenia purpurea          --           --           18, 1.7, 3
Scirpus acutus               7, 1, 2      34, 19, 17   74, 19.5, 20
Scirpus americanus           3, 0.33, 1   --           --
Scirpus cyperinus            --           --           --
Selaginella sp.              P            --           --
Solanum dulcamara            --           --           --
Solidago sp.                 37, 0.93, 7  --           --
Spiraea tomentosa            --           7, 0.21, 1   7, 0.37, 1
Symplocarpus Foetidus        3.0, 17, 1   --           --
Thelypteris palustris        13, 0.57, 3  45, 6, 10    81, 8, 12
Taxicodendron radicans       --           --           4, 0.04 <1
Toxicodendron vernix         --           --           --
Trientalis borealis          --           --           --
Triglochin maritima          --           --           --
Typha sp(p).                 --           90, 34, 34   26, 1.2, 3
Ulmnus americana             --           --           --
Urtia procera                --           --           7, 0.22, 1
Vaccinium corymbosum         --           --           P
Vaccinium macrocarpon        --           --           52, 2.5, 6
Vaccinium oxycoccos          --           --           --
Viola blanda                 --           24, 0.83, 4  --
Viola cucullata              3, 0.17, 1   P            --
Viola pallens                --           --           --
Viola sp.                    --           --           44, 0.81, 4
Woodwardia virginica         --           --           --
Unidentified sedges          --           --           --

                                         Tall-shrub bogs
      Species                       LCB        HB         THB

Acer rubrum                  17, 0.17, 2   P           33, 0.3, 5
Alisma subcordata            --            --          --
Andromeda glaucophylla       --            --          --
Aronia melanocarpa           --            --          33, 1, 6
Betula allegheniensis        --            --          --
Betula pumila                --            --          --
Bidens sp.                   --            28, 1, 3    --
Calla palustris              --            --          --
Calopogon pulchellus         P             --          --
Caltha palustris             --            --          --
Cardamine bulbosa            --            --          --
Cardamine pratensis          --            --          --
Carex crinita                --            7, <1, 1    --
Carex lasiocarpa             --            --          --
Carex oligosperma            --            --          --
Carex rostrata               --            --          --
Carex sp.                    17, 3, 4      7, <1, I    --
Carex trisperma              --            --          --
Cephalanthus occidentalis    --            29, 3, 4    --
Chamaedaphne calyculata      --            --          P
Coptis trifolia              --            --          --
Cornus foemina               --            --          --
Cornus stolonifera           --            --          --
Cuscuta sp.                  --            --          --
Cypripedium acaule           --            --          --
Decodon verticillatus        --            --          --
Drosera intermedia           --            --          --
Drosera rotundifolia         69, 0.69, 6   --          33, 0.3, 5
Dryopteris spinulosa         --            --          --
Dulichium arundinaceum       --            64, 34, 18  P
Eleocharis sp.               --            --          --
Equisetum fluviatile         4, 0.04, <1   --          --
Equisetum palustris          --            --          --
Eriophorum virginicum        --            --          --
Eupatorium maculatum         --            --          --
Eupatorium perfoliatum       --            7, <1, 1    --
Galium sp.                   --            P           --
Gaylussacia baccata          --            --          --
Hypericum virginicum         22, 0.87, 3   36, 4, 5    --
Ilex verticillata            17, 0.17, 2   --          --
Impatiens capensis           --            64, 6, 8    --
Iris versicolor              4, 0.22, 1    --          --
Isotria verticillata         --            --          --
Juncus sp.                   --            7, <1, 1    --
Leersia oryzoides            --            50, 5, 6    --
Lindera benzoin              --            --          --
Liparis loesellii            --            --          --
Liriodendron tulipifera      --            --          --
Lycopodium lucidulum         --            --          --
Lycopodium obscurum          --            --          --
Lysimachia terrestris        --            --          --
Lysimachia thrysiflora       17, 0.52, 2   --          --
Mainanthemum canadense       --            --          --
Menyanthes trifoliata        22, 5.6, 7    --          --
Mitchella repens             --            --          --
Nuphar advena                P             --          P
Nymphaea odorata             --            --          --
Nyssa sylvatica              --            --          --
Onoclea sensibilis           P             7, <1, 1    --
Osmunda cinnamomea           --            --          --
Osmunda regalis              26, 7.4, 9    --          33, 30, 29
Parnassia glauca             --            --          --
Parthenocissus quinquefolia  --            --          --
Pinus strobus                --            --          --
Poaceae (unidentified)       --            --          --
Polygonum sp.                --            --          --
Populus tremuloides          4, 0.13, <1   --          --
Potentilla anserina          4, 0.04, <1   --          --
Potentilla fruticosa         --            --          --
potentilla palustris         35, 0.75, 3   --          --
Prunus serotina              --            --          --
Quereus palustris            --            P           --
Quercus rubra                --            --          --
Ranunculus sp.               --            --          --
Rosa palustris               13, 0.3, 1    --          --
Rubus allegheniensis         --            --          --
Rubus hispidus               --            --          --
Rudheckia sp.                --            --          --
Rumex sp.                    --            --          --
Saggitaria latifolia         --            --          --
Salixcandida                 --            --          --
Salix pedicellaris           39, 1.3, 4    --          --
Salix serrissima             --            --          --
Sarracenia purpurea          35, 1.6, 4    --          --
Scirpus acutus               4, 0.22, 1    --          --
Scirpus americanus           --            --          --
Scirpus cyperinus            --            P           --
Selaginella sp.              --            --          --
Solanum dulcamara            --            --          --
Solidago sp.                 --            --          --
Spiraea tomentosa            --            --          --
Symplocarpus Foetidus        --            --          --
Thelypteris palustris        78, 5.3, 11   7, <1, 1    --
Taxicodendron radicans       --            --          --
Toxicodendron vernix         26, 1.7, 4    --          33, 1.7, 7
Trientalis borealis          --            --          --
Triglochin maritima          4, 0.22, 1    --          --
Typha sp(p).                 61, 3.8, 8    P           --
Ulmnus americana             --            --          --
Urtia procera                --            14, <1, 2   --
Vaccinium corymbosum         --            --          67, 2.7, 13
Vaccinium macrocarpon        52, 14.6, 18  --          --
Vaccinium oxycoccos          --            --          --
Viola blanda                 P             --          --
Viola cucullata              35, 0.35, 3   --          --
Viola pallens                --            --          --
Viola sp.                    --            --          --
Woodwardia virginica         --            --          100, 15, 33
Unidentified sedges          34, 0.91, 5   35, 1, 4    --

                               Tall-shrub        Leatherleaf bogs
                                  bogs
      Species                   BLB               DSB       LB

Acer rubrum                  20, 0.2, 4    --           3, <1, <1
Alisma subcordata            --            --           P
Andromeda glaucophylla       --            P            11, 4, 3
Aronia melanocarpa           40, 1.2, 9    15, 0.35, 4  --
Betula allegheniensis        --            --           --
Betula pumila                --            --           --
Bidens sp.                   20, 8, 15     --           --
Calla palustris              --            --           --
Calopogon pulchellus         --            --           --
Caltha palustris             --            --           --
Cardamine bulbosa            --            --           --
Cardamine pratensis          --            --           --
Carex crinita                --            --           P
Carex lasiocarpa             --            --           --
Carex oligosperma            --            --           14, 5, 4
Carex rostrata               --            --           78, 24, 18
Carex sp.                    --            --           --
Carex trisperma              --            --           --
Cephalanthus occidentalis    --            6, 1.8, 3    11, 2, 2
Chamaedaphne calyculata      20, 1, 5      42, 24, 31   71, 19, 15
Coptis trifolia              --            --           --
Cornus foemina               --            --           --
Cornus stolonifera           --            --           --
Cuscuta sp.                  --            --           --
Cypripedium acaule           --            --           --
Decodon verticillatus        --            --           P
Drosera intermedia           --            P            --
Drosera rotundifolia         --            P            --
Dryopteris spinulosa         --            --           --
Dulichium arundinaceum       20, 2, 7      2, 0.02, 1   3, <1, <1
Eleocharis sp.               --            --           --
Equisetum fluviatile         --            --           --
Equisetum palustris          --            --           --
Eriophorum virginicum        --            4, 0.33, 1   --
Eupatorium maculatum         --            --           --
Eupatorium perfoliatum       --            --           --
Galium sp.                   21, 1, 5      --           3, <1, <1
Gaylussacia baccata          --            --           --
Hypericum virginicum         60, 12, 28    --           14, <1, 2
Ilex verticillata            --            --           --
Impatiens capensis           --            --           7, <1, 1
Iris versicolor              --            2, 0.02, 1   P
Isotria verticillata         --            --           --
Juncus sp.                   --            --           P
Leersia oryzoides            --            --           --
Lindera benzoin              --            --           --
Liparis loesellii            --            --           --
Liriodendron tulipifera      --            --           --
Lycopodium lucidulum         --            --           --
Lycopodium obscurum          --            --           --
Lysimachia terrestris        --            --           --
Lysimachia thrysiflora       --            --           3, <1, <1
Mainanthemum canadense       --            --           --
Menyanthes trifoliata        --            --           --
Mitchella repens             --            --           --
Nuphar advena                --            P            --
Nymphaea odorata             --            --           --
Nyssa sylvatica              --            --           --
Onoclea sensibilis           --            --           --
Osmunda cinnamomea           --            --           --
Osmunda regalis              --            --           --
Parnassia glauca             --            --           --
Parthenocissus quinquefolia  --            --           --
Pinus strobus                --            --           --
Poaceae (unidentified)       60, 10.6, 26  4, 2, 3      --
Polygonum sp.                --            --           P
Populus tremuloides          --            --           --
Potentilla anserina          --            --           --
Potentilla fruticosa         --            --           --
potentilla palustris         --            --           P
Prunus serotina              --            --           --
Quereus palustris            --            --           --
Quercus rubra                --            --           --
Ranunculus sp.               --            --           --
Rosa palustris               --            --           18, 4, 3
Rubus allegheniensis         --            --           --
Rubus hispidus               --            44, 3, 14    --
Rudheckia sp.                --            --           --
Rumex sp.                    --            --           --
Saggitaria latifolia         --            --           11,<1, 1
Salixcandida                 --            --           --
Salix pedicellaris           --            --           --
Salix serrissima             --            --           --
Sarracenia purpurea          --            P            P
Scirpus acutus               --            --           --
Scirpus americanus           --            --           --
Scirpus cyperinus            --            27, 11, 16   P
Selaginella sp.              --            --           --
Solanum dulcamara            P             --           --
Solidago sp.                 --            --           --
Spiraea tomentosa            --            --           --
Symplocarpus Foetidus        --            --           --
Thelypteris palustris        --            --           --
Taxicodendron radicans       --            --           --
Toxicodendron vernix         --            --           --
Trientalis borealis          --            --           --
Triglochin maritima          --            --           --
Typha sp(p).                 --            --           29, 5, 4
Ulmnus americana             --            --           --
Urtia procera                --            --           --
Vaccinium corymbosum         --            24, 2.7, 8   --
Vaccinium macrocarpon        --            --           10, 2, 2
Vaccinium oxycoccos          --            9, 3, 5      --
Viola blanda                 --            --           --
Viola cucullata              --            --           --
Viola pallens                --            --           --
Viola sp.                    --            --           --
Woodwardia virginica         P             29, 7,14     36, 15, 10
Unidentified sedges          --            --           14, 1, 1

                                          Leatherleaf bogs
      Species                     YB           BUR           SH

Acer rubrum                  15, 0.25, 2   4, 0.11, 1    --
Alisma subcordata            --            --            --
Andromeda glaucophylla       P             P             --
Aronia melanocarpa           --            7, 0.3, 1     3, 0.94, 2
Betula allegheniensis        --            --            --
Betula pumila                --            --            --
Bidens sp.                   --            --            --
Calla palustris              --            --            --
Calopogon pulchellus         --            --            --
Caltha palustris             --            --            --
Cardamine bulbosa            --            --            --
Cardamine pratensis          --            --            --
Carex crinita                --            --            --
Carex lasiocarpa             --            --            --
Carex oligosperma            20, 1.7, 4    74, 15.8, 22  --
Carex rostrata               15, 1.7, 3    11, 0.41, 2   P
Carex sp.                    --            --            --
Carex trisperma              --            --            --
Cephalanthus occidentalis    30, 4, 7      --            3, 0.47, 1
Chamaedaphne calyculata      55, 14.2, 17  85, 46.5, 43  94, 69, 72
Coptis trifolia              --            --            --
Cornus foemina               --            --            --
Cornus stolonifera           --            --            --
Cuscuta sp.                  --            --            --
Cypripedium acaule           --            --            --
Decodon verticillatus        40, 4.2, 8    7, 0.73, 2    --
Drosera intermedia           --            4, 0.04, 1    --
Drosera rotundifolia         P             --            --
Dryopteris spinulosa         --            --            --
Dulichium arundinaceum       5, 0.05, 1    4, 0.37, 1    --
Eleocharis sp.               --            --            --
Equisetum fluviatile         --            --            --
Equisetum palustris          --            --            --
Eriophorum virginicum        --            --            --
Eupatorium maculatum         --            --            --
Eupatorium perfoliatum       --            --            --
Galium sp.                   --            --            --
Gaylussacia baccata          --            4, 0.55, 1    --
Hypericum virginicum         20, 1.4, 4    11, 0.33, 2   3, 0.31, 1
Ilex verticillata            --            --            --
Impatiens capensis           --            --            --
Iris versicolor              --            P             --
Isotria verticillata         --            --            --
Juncus sp.                   --            --            --
Leersia oryzoides            --            --            --
Lindera benzoin              --            --            --
Liparis loesellii            --            --            --
Liriodendron tulipifera      --            --            --
Lycopodium lucidulum         --            --            --
Lycopodium obscurum          --            --            --
Lysimachia terrestris        --            --            --
Lysimachia thrysiflora       P             --            --
Mainanthemum canadense       --            --            --
Menyanthes trifoliata        --            --            --
Mitchella repens             --            --            --
Nuphar advena                5, 1, 1       4, 1.08, 1    --
Nymphaea odorata             P             --            --
Nyssa sylvatica              --            --            --
Onoclea sensibilis           --            --            --
Osmunda cinnamomea           --            --            --
Osmunda regalis              --            --            --
Parnassia glauca             --            --            --
Parthenocissus quinquefolia  --            --            --
Pinus strobus                --            --            --
Poaceae (unidentified)       5, 1, 1       --            --
Polygonum sp.                --            --            --
Populus tremuloides          --            --            --
Potentilla anserina          --            --            --
Potentilla fruticosa         --            --            --
potentilla palustris         --            P             --
Prunus serotina              --            --            --
Quereus palustris            5, 0.15, 1    --            --
Quercus rubra                --            --            --
Ranunculus sp.               --            --            --
Rosa palustris               --            --            --
Rubus allegheniensis         --            --            --
Rubus hispidus               20, 1.3, 3    --            3, 0.16, 1
Rudheckia sp.                --            --            --
Rumex sp.                    --            --            --
Saggitaria latifolia         --            P             --
Salixcandida                 --            --            --
Salix pedicellaris           --            P             --
Salix serrissima             --            --            --
Sarracenia purpurea          15, 1.4, 3    P             --
Scirpus acutus               --            --            --
Scirpus americanus           --            --            --
Scirpus cyperinus            10, 0.2, 1    P             P
Selaginella sp.              --            --            --
Solanum dulcamara            --            --            --
Solidago sp.                 --            --            --
Spiraea tomentosa            15, 3.7, 5    --            --
Symplocarpus Foetidus        --            --            --
Thelypteris palustris        --            --            --
Taxicodendron radicans       --            --            --
Toxicodendron vernix         30, 0.8, 4    4, 0.18, 1    --
Trientalis borealis          --            --            --
Triglochin maritima          --            --            --
Typha sp(p).                 --            7, 0.3, 1     --
Ulmnus americana             --            --            --
Urtia procera                --            --            --
Vaccinium corymbosum         20, 3.5, 5    --            --
Vaccinium macrocarpon        30, 7.7, 9    7, 1.8, 2     --
Vaccinium oxycoccos          10, 2, 3      22, 3.2, 6    --
Viola blanda                 --            --            --
Viola cucullata              --            --            --
Viola pallens                --            --            --
Viola sp.                    --            --            --
Woodwardia virginica         35, 17.6, 18  48, 8.8, 13   34, 16.7, 22
Unidentified sedges          --            --            --

                                       Forested peatlands
      Species                    LAB          MPB        TAM (*)

Acer rubrum                  82, 2.1, 7   50, 0.78, 7   86, 3, 10
Alisma subcordata            --           --            --
Andromeda glaucophylla       --           --            --
Aronia melanocarpa           --           14, 0.36, 2   3, <1, 1
Betula allegheniensis        --           --            --
Betula pumila                --           --            --
Bidens sp.                   54, 2.3, 6   14, 2.9, 5    5, <1, 1
Calla palustris              --           7, 3.6, 5     --
Calopogon pulchellus         --           --            --
Caltha palustris             18, 2.7, 4   --            --
Cardamine bulbosa            --           --            --
Cardamine pratensis          --           --            --
Carex crinita                --           14, 0.43, 2   --
Carex lasiocarpa             --           --            --
Carex oligosperma            --           --            --
Carex rostrata               --           --            --
Carex sp.                    18, 1.8, 3   --            --
Carex trisperma              --           --            P
Cephalanthus occidentalis    --           --            --
Chamaedaphne calyculata      --           P             --
Coptis trifolia              P            --            --
Cornus foemina               --           --            --
Cornus stolonifera           --           --            --
Cuscuta sp.                  --           14, 0.14, 2   --
Cypripedium acaule           --           7, 0.21, 1    P
Decodon verticillatus        --           --            --
Drosera intermedia           --           --            --
Drosera rotundifolia         --           --            --
Dryopteris spinulosa         45, 1.4, 4   --            27, 1, 3
Dulichium arundinaceum       --           7, 2.1, 4     --
Eleocharis sp.               --           --            --
Equisetum fluviatile         --           --            --
Equisetum palustris          --           --            --
Eriophorum virginicum        --           --            --
Eupatorium maculatum         --           --            --
Eupatorium perfoliatum       --           --            --
Galium sp.                   18, 0.36, 2  --            --
Gaylussacia baccata          P            --            --
Hypericum virginicum         --           7, 1.4, 3     --
Ilex verticillata            9, 0.27, 1   7, 0.21, 1    8, <1, 1
Impatiens capensis           81, 7.1, 13  7, 0.07, 1    --
Iris versicolor              --           --            --
Isotria verticillata         P            --            --
Juncus sp.                   --           --            --
Leersia oryzoides            --           --            --
Lindera benzoin              --           --            11, 1, 2
Liparis loesellii            P            --            --
Liriodendron tulipifera      --           14, 0.14, 2   --
Lycopodium lucidulum         P            --            P
Lycopodium obscurum          --           --            P
Lysimachia terrestris        --           --            --
Lysimachia thrysiflora       P            7, 0.71, 2    --
Mainanthemum canadense       45, 3.5, 7   71, 13.2, 25  97, 24, 29
Menyanthes trifoliata        9, 0.91, 2   --            --
Mitchella repens             P            --            --
Nuphar advena                --           7, 1.4, 3     --
Nymphaea odorata             --           --            --
Nyssa sylvatica              --           57, 6.2, 15   --
Onoclea sensibilis           P            --            --
Osmunda cinnamomea           45, 6.8, 10  14, 1.3, 3    73, 6, 11
Osmunda regalis              P            --            --
Parnassia glauca             --           --            --
Parthenocissus quinquefolia  54, 2.4, 6   --            13, 1, 2
Pinus strobus                --           7, 0.21, 1    --
Poaceae (unidentified)       --           --            --
Polygonum sp.                --           --            --
Populus tremuloides          --           --            --
Potentilla anserina          --           --            --
Potentilla fruticosa         --           --            --
potentilla palustris         --           --            --
Prunus serotina              --           --            22, 1, 3
Quereus palustris            --           --            13, 1, 2
Quercus rubra                --           7, 0.07, 1    --
Ranunculus sp.               --           --            --
Rosa palustris               18, 0.54, 2  --            --
Rubus allegheniensis         --           --            16, 1, 2
Rubus hispidus               18, 0.73, 2  --            57, 6, 10
Rudheckia sp.                --           --            --
Rumex sp.                    --           --            --
Saggitaria latifolia         --           --            --
Salixcandida                 P            --            --
Salix pedicellaris           P            --            --
Salix serrissima             --           --            --
Sarracenia purpurea          9, 0.27, 1   --            --
Scirpus acutus               --           --            --
Scirpus americanus           --           --            --
Scirpus cyperinus            --           --            --
Selaginella sp.              --           --            --
Solanum dulcamara            --           --            --
Solidago sp.                 --           --            --
Spiraea tomentosa            --           --            --
Symplocarpus Foetidus        54, 7.7, 12  --            --
Thelypteris palustris        --           --            --
Taxicodendron radicans       36, 0.73, 3  --            --
Toxicodendron vernix         --           --            --
Trientalis borealis          18, 0.36, 2  43, 1.3, 7    89, 7, 13
Triglochin maritima          --           --            --
Typha sp(p).                 P            --            --
Ulmnus americana             27, 0.27, 2  --            --
Urtia procera                --           21, 0.5, 3    P
Vaccinium corymbosum         36, 2.7, 5   21, 1.5, 4    11, 1, 2
Vaccinium macrocarpon        9, 0.27, 1   --            --
Vaccinium oxycoccos          --           --            --
Viola blanda                 --           --            --
Viola cucullata              --           --            --
Viola pallens                64, 2.2, 6   --            --
Viola sp.                    --           --            --
Woodwardia virginica         --           7, 0.21, 1    --
Unidentified sedges          --           --            --

                               Forested
                               peatlands
      Species                  RMB

Acer rubrum                  83, 1.3, 12
Alisma subcordata            --
Andromeda glaucophylla       --
Aronia melanocarpa           --
Betula allegheniensis        17, 0.17, 2
Betula pumila                --
Bidens sp.                   8, 0.25, 1
Calla palustris              --
Calopogon pulchellus         --
Caltha palustris             --
Cardamine bulbosa            --
Cardamine pratensis          --
Carex crinita                --
Carex lasiocarpa             --
Carex oligosperma            --
Carex rostrata               8, 1.7, 4
Carex sp.                    --
Carex trisperma              33, 2.2, 8
Cephalanthus occidentalis    --
Chamaedaphne calyculata      --
Coptis trifolia              --
Cornus foemina               --
Cornus stolonifera           --
Cuscuta sp.                  --
Cypripedium acaule           --
Decodon verticillatus        --
Drosera intermedia           --
Drosera rotundifolia         --
Dryopteris spinulosa         --
Dulichium arundinaceum       --
Eleocharis sp.               --
Equisetum fluviatile         --
Equisetum palustris          --
Eriophorum virginicum        --
Eupatorium maculatum         --
Eupatorium perfoliatum       --
Galium sp.                   --
Gaylussacia baccata          --
Hypericum virginicum         --
Ilex verticillata            75, 3.8, 16
Impatiens capensis           17, 1.7, 5
Iris versicolor              --
Isotria verticillata         --
Juncus sp.                   --
Leersia oryzoides            --
Lindera benzoin              --
Liparis loesellii            --
Liriodendron tulipifera      --
Lycopodium lucidulum         --
Lycopodium obscurum          --
Lysimachia terrestris        --
Lysimachia thrysiflora       --
Mainanthemum canadense       58, 2.5, 12
Menyanthes trifoliata        --
Mitchella repens             --
Nuphar advena                --
Nymphaea odorata             --
Nyssa sylvatica              --
Onoclea sensibilis           8, 0.25, 1
Osmunda cinnamomea           42, 3.8, 13
Osmunda regalis              --
Parnassia glauca             --
Parthenocissus quinquefolia  --
Pinus strobus                --
Poaceae (unidentified)       8, 0.83, 8
Polygonum sp.                --
Populus tremuloides          --
Potentilla anserina          --
Potentilla fruticosa         --
potentilla palustris         --
Prunus serotina              --
Quereus palustris            --
Quercus rubra                --
Ranunculus sp.               --
Rosa palustris               8, 0.25, 1
Rubus allegheniensis         --
Rubus hispidus               8, 0.25, 1
Rudheckia sp.                --
Rumex sp.                    --
Saggitaria latifolia         --
Salixcandida                 --
Salix pedicellaris           --
Salix serrissima             --
Sarracenia purpurea          --
Scirpus acutus               --
Scirpus americanus           --
Scirpus cyperinus            --
Selaginella sp.              --
Solanum dulcamara            --
Solidago sp.                 --
Spiraea tomentosa            --
Symplocarpus Foetidus        --
Thelypteris palustris        --
Taxicodendron radicans       8, 0.83, 3
Toxicodendron vernix         --
Trientalis borealis          25, 0.75, 4
Triglochin maritima          --
Typha sp(p).                 --
Ulmnus americana             --
Urtia procera                8, 1.7, 4
Vaccinium corymbosum         25, 1.5, 6
Vaccinium macrocarpon        --
Vaccinium oxycoccos          --
Viola blanda                 --
Viola cucullata              --
Viola pallens                --
Viola sp.                    --
Woodwardia virginica         --
Unidentified sedges          --
Table 6

Bryophytes from 16 peatlands in northern Indiana. The first number in
each cell is the percent frequency for each species, the second number
is the percent cover, and the third number is the importance value for
each species in the respective peatlands. Peatland acronyms correspond
with Table 2. A "P" means the species was present in the ground layer,
but occurred outside of the sampling quadrats.

                                              Fens
Species                         KLF          SVF           BF

Amblystegium riparium       4, 11.9, 55  --           --
Aulacomnium palustre        P            7, 1, 2      --
Bryum pseudotriquetrum      --           3, 0.03, 1   --
Callicaldium haldanianum    --           --           --
Calliergon cordifolium      --           --           --
Calliergon giganteum        --           --           --
Calliergonella cuspidata    --           59, 5.8, 35  44, 5.1, 29
Campylium polygamum         --           17, 2.7, 13  7, 0.92, 5
Campylium stellatum         --           7, 0.45, 3   74, 15.7, 65
Climacium americanum        --           --           --
Drepanocladus fluitans      --           14, 1.5, 9   --
Drepanocladus sp.           --           --           --
Eurynchium hyans            23, 6, 30    --           --
Fissidens adianthoides      3, 0.03, 2   21, 1.4, 11  --
Helodium paludosum          --           --           --
Hypnum pratense             --           --           --
Isopterygium elegans        --           P            --
Leucobyrum albidum          --           --           --
Lophocolia heterophylla     --           3, 0.3, 1    --
Mnium affine var. ciliare   3, 2.7, 8    --           --
Mnium affine var. rugicum   --           3, 0.69, 3   --
Mnium cuspidatum            --           --           --
Pallavicinia Iyellii        --           --           --
Philonotis fontana          --           --           --
Plagiothecium denticulatum  --           --           --
Polytrichum strictum        --           --           --
Rhynchostegium serrulatum   --           --           --
Riccia fluitans             --           P            --
Sphagnum affine             --           --           --
Sphagnum bartlettianum      --           --           --
Sphagnum capillifolium      --           --           --
Sphagnum centrale           --           --           --
Sphagnum cuspidatum         --           --           --
Sphagnum fallax             --           --           --
Sphagnum fimbriatum         P            10, 7, 21    --
Sphagnum fuscum             --           --           P
Sphagnum isoviitae          --           --           --
Sphagnum magellanicum       3, 1, 5      --           --
Sphagnum palustre           --           --           P
Sphagnum papillosum         --           --           --
Sphagnum recurvum           --           --           --
Sphagnum russowii           --           --           --
Sphagnum sp.                --           --           --
Sphagnunm squarrosum        --           --           --
Sphagnum teres              --           P            P
Sphagnum warnstorfii        --           --           P
Tetraphis pellucida         --           --           --
Thuidium allenii            --           --           --
Thuidium delicatulum        --           --           --

                                         Tall-shrub bogs
Species                        LCB            HB         THB

Amblystegium riparium       --            7, <1, 1    --
Aulacomnium palustre        56, 7.7, 22   7, <1, 1    33, 6.7, 14
Bryum pseudotriquetrum      --            --          --
Callicaldium haldanianum    --            --          --
Calliergon cordifolium      --            --          --
Calliergon giganteum        --            28, 1, 3    --
Calliergonella cuspidata    4, 0.04, 1    --          --
Campylium polygamum         --            --          --
Campylium stellatum         --            --          --
Climacium americanum        --            P           --
Drepanocladus fluitans      --            --          --
Drepanocladus sp.           --            --          --
Eurynchium hyans            --            --          --
Fissidens adianthoides      --            --          --
Helodium paludosum          --            --          --
Hypnum pratense             --            --          --
Isopterygium elegans        --            --          --
Leucobyrum albidum          --            --          --
Lophocolia heterophylla     9, 0.09, 3    --          --
Mnium affine var. ciliare   --            --          --
Mnium affine var. rugicum   --            --          --
Mnium cuspidatum            --            --          --
Pallavicinia Iyellii        --            --          --
Philonotis fontana          --            --          --
Plagiothecium denticulatum  --            --          --
Polytrichum strictum        --            --          --
Rhynchostegium serrulatum   --            --          --
Riccia fluitans             --            P           --
Sphagnum affine             --            --          33, 1.7, 5
Sphagnum bartlettianum      P             --          --
Sphagnum capillifolium      --            --          --
Sphagnum centrale           13, 7, 9      --          --
Sphagnum cuspidatum         --            --          --
Sphagnum fallax             --            --          --
Sphagnum fimbriatum         39, 25.7, 29  28, 25, 11  --
Sphagnum fuscum             --            --          --
Sphagnum isoviitae          --            --          --
Sphagnum magellanicum       --            --          --
Sphagnum palustre           26, 19.1, 21  --          100, 71.7, 75
Sphagnum papillosum         --            --          --
Sphagnum recurvum           --            --          33, 1.7, 5
Sphagnum russowii           --            P           --
Sphagnum sp.                --            --          --
Sphagnunm squarrosum        --            --          --
Sphagnum teres              17, 12.6, 14  --          --
Sphagnum warnstorfii        4, 0.43, 2    --          --
Tetraphis pellucida         --            --          --
Thuidium allenii            --            --          --
Thuidium delicatulum        --            --          --

                              Tall-shrub        Leatherleaf bogs
                                 bogs
Species                        BLB               DSB       LB

Amblystegium riparium       --            --            --
Aulacomnium palustre        20, 0.02, 7   84, 16, 29    12, <1, 1
Bryum pseudotriquetrum      --            --            --
Callicaldium haldanianum    --            --            --
Calliergon cordifolium      --            --            --
Calliergon giganteum        --            --            --
Calliergonella cuspidata    --            --            --
Campylium polygamum         --            --            --
Campylium stellatum         --            --            --
Climacium americanum        --            --            --
Drepanocladus fluitans      --            --            --
Drepanocladus sp.           20, 6, 10     --            --
Eurynchium hyans            --            --            --
Fissidens adianthoides      --            --            --
Helodium paludosum          --            --            --
Hypnum pratense             --            --            --
Isopterygium elegans        --            --            --
Leucobyrum albidum          P             --            --
Lophocolia heterophylla     --            --            --
Mnium affine var. ciliare   --            --            --
Mnium affine var. rugicum   --            --            --
Mnium cuspidatum            --            --            --
Pallavicinia Iyellii        --            --            --
Philonotis fontana          --            --            --
Plagiothecium denticulatum  --            --            --
Polytrichum strictum        --            9, 0.88, 2    --
Rhynchostegium serrulatum   --            --            --
Riccia fluitans             --            --            P
Sphagnum affine             --            --            --
Sphagnum bartlettianum      --            13, 4.7, 6    --
Sphagnum capillifolium      --            6, 4.1, 4     --
Sphagnum centrale           --            --            --
Sphagnum cuspidatum         40, 26, 28    --            25, 12, 8
Sphagnum fallax             --            53, 23, 27    --
Sphagnum fimbriatum         --            2, 0.22, 1    --
Sphagnum fuscum             --            --            --
Sphagnum isoviitae          --            --            --
Sphagnum magellanicum       P             38, 7.8, 13   4, <1, 1
Sphagnum palustre           --            --            4, 3, 2
Sphagnum papillosum         --            --            --
Sphagnum recurvum           60, 59.8, 54  31, 16.6, 18  4, 2, 1
Sphagnum russowii           --            --            --
Sphagnum sp.                --            --            4, <1, 1
Sphagnunm squarrosum        --            --            4, <1, 1
Sphagnum teres              --            --            --
Sphagnum warnstorfii        --            --            --
Tetraphis pellucida         --            --            --
Thuidium allenii            --            --            --
Thuidium delicatulum        --            --            --

                                         Leatherleaf bogs
Species                        YB            BUR            SH

Amblystegium riparium       --            --            --
Aulacomnium palustre        25, 3.2, 7    18, 3.7, 7    6, 0.47, 3
Bryum pseudotriquetrum      --            --            --
Callicaldium haldanianum    --            --            --
Calliergon cordifolium      --            --            --
Calliergon giganteum        --            --            --
Calliergonella cuspidata    --            --            --
Campylium polygamum         --            --            --
Campylium stellatum         --            --            --
Climacium americanum        --            --            --
Drepanocladus fluitans      --            --            --
Drepanocladus sp.           --            --            --
Eurynchium hyans            --            --            --
Fissidens adianthoides      --            --            --
Helodium paludosum          --            --            --
Hypnum pratense             --            --            --
Isopterygium elegans        --            --            --
Leucobyrum albidum          --            --            --
Lophocolia heterophylla     --            --            --
Mnium affine var. ciliare   --            --            --
Mnium affine var. rugicum   --            --            --
Mnium cuspidatum            --            --            --
Pallavicinia Iyellii        --            --            --
Philonotis fontana          --            --            --
Plagiothecium denticulatum  --            --            --
Polytrichum strictum        35, 2.1, 9    7, 1.7, 3     --
Rhynchostegium serrulatum   --            --            --
Riccia fluitans             --            --            --
Sphagnum affine             --            --            --
Sphagnum bartlettianum      10, 2.2, 3    37, 8.5, 15   --
Sphagnum capillifolium      --            --            --
Sphagnum centrale           20, 7.5, 9    --            --
Sphagnum cuspidatum         --            30, 25.2, 23  84, 66.5, 69
Sphagnum fallax             --            --            --
Sphagnum fimbriatum         15, 9, 8      7, 0.48, 2    --
Sphagnum fuscum             --            --            --
Sphagnum isoviitae          --            11, 2.2, 4    --
Sphagnum magellanicum       60, 26.5, 28  4, 0.37, 1    41, 23.9, 29
Sphagnum palustre           --            --            --
Sphagnum papillosum         --            67, 36.7, 40  --
Sphagnum recurvum           60, 37.1, 34  7, 2.2, 3     --
Sphagnum russowii           --            --            --
Sphagnum sp.                --            --            --
Sphagnunm squarrosum        --            --            --
Sphagnum teres              --            --            --
Sphagnum warnstorfii        --            --            --
Tetraphis pellucida         --            --            --
Thuidium allenii            --            --            --
Thuidium delicatulum        --            --            --

                                      Forested peatlands
Species                          LAB          MPB       TAM (*)

Amblystegium riparium       18, 1.4, 5    --           --
Aulacomnium palustre        P             21, 1.7, 13  40, 1, 11
Bryum pseudotriquetrum      --            --           --
Callicaldium haldanianum    --            --           24, 1, 8
Calliergon cordifolium      --            --           --
Calliergon giganteum        --            --           --
Calliergonella cuspidata    9, 0.91, 3    --           --
Campylium polygamum         --            7, 0.71, 5   --
Campylium stellatum         --            --           --
Climacium americanum        --            --           --
Drepanocladus fluitans      --            --           --
Drepanocladus sp.           --            --           --
Eurynchium hyans            --            --           --
Fissidens adianthoides      --            --           --
Helodium paludosum          --            7, 2.1, 10   --
Hypnum pratense             --            21, 2.5, 16  --
Isopterygium elegans        --            --           --
Leucobyrum albidum          9, 3.6, 6     --           --
Lophocolia heterophylla     --            36, 1.3, 15  27, <1, 8
Mnium affine var. ciliare   --            --           --
Mnium affine var. rugicum   --            --           --
Mnium cuspidatum            --            --           13, <1, 5
Pallavicinia Iyellii        36, 5.3, 14   57, 3.8, 31  76, 5, 30
Philonotis fontana          --            --           --
Plagiothecium denticulatum  --            --           13, <1, 5
Polytrichum strictum        --            --           --
Rhynchostegium serrulatum   --            --           5, 1, 4
Riccia fluitans             P             --           --
Sphagnum affine             54, 14.4, 29  7, 0.36, 3   --
Sphagnum bartlettianum      --            --           --
Sphagnum capillifolium      --            --           --
Sphagnum centrale           --            --           --
Sphagnum cuspidatum         --            --           --
Sphagnum fallax             --            --           --
Sphagnum fimbriatum         --            7, 0.36, 3   --
Sphagnum fuscum             --            --           --
Sphagnum isoviitae          --            --           --
Sphagnum magellanicum       --            --           --
Sphagnum palustre           --            --           <1, <1, 3
Sphagnum papillosum         --            --           --
Sphagnum recurvum           --            --           3, 2, 6
Sphagnum russowii           --            --           --
Sphagnum sp.                --            --           --
Sphagnunm squarrosum        --            --           --
Sphagnum teres              --            --           --
Sphagnum warnstorfii        --            --           --
Tetraphis pellucida         18, 0.36, 4   7, 0.36, 3   30, 2, 12
Thuidium allenii            18, 0.91, 5   --           --
Thuidium delicatulum        54, 13, 27    --           5, <1, 4

                               Forested
                              peatlands
Species                        RMB

Amblystegium riparium       17, 2.9, 7
Aulacomnium palustre        67, 12.1, 27
Bryum pseudotriquetrum      --
Callicaldium haldanianum    --
Calliergon cordifolium      8, 0.83, 3
Calliergon giganteum        --
Calliergonella cuspidata    --
Campylium polygamum         --
Campylium stellatum         --
Climacium americanum        --
Drepanocladus fluitans      --
Drepanocladus sp.           --
Eurynchium hyans            --
Fissidens adianthoides      --
Helodium paludosum          --
Hypnum pratense             --
Isopterygium elegans        --
Leucobyrum albidum          --
Lophocolia heterophylla     --
Mnium affine var. ciliare   --
Mnium affine var. rugicum   --
Mnium cuspidatum            --
Pallavicinia Iyellii        42, 3, 12
Philonotis fontana          --
Plagiothecium denticulatum  --
Polytrichum strictum        --
Rhynchostegium serrulatum   8, 0.08, 2
Riccia fluitans             P
Sphagnum affine             42, 11.3, 21
Sphagnum bartlettianum      --
Sphagnum capillifolium      --
Sphagnum centrale           --
Sphagnum cuspidatum         --
Sphagnum fallax             50, 17.9, 29
Sphagnum fimbriatum         --
Sphagnum fuscum             --
Sphagnum isoviitae          --
Sphagnum magellanicum       --
Sphagnum palustre           --
Sphagnum papillosum         --
Sphagnum recurvum           --
Sphagnum russowii           --
Sphagnum sp.                --
Sphagnunm squarrosum        --
Sphagnum teres              --
Sphagnum warnstorfii        --
Tetraphis pellucida         --
Thuidium allenii            --
Thuidium delicatulum        --
Table 7

Aquatic layer vegetation from 16 peatlands in northern Indiana. The
first number in each cell is the percent frequency for each species, the
second number is the percent cover, and the third number is the
importance value for each species in the respective peatlands. Peatland
acronyms correspond with Table 2. A "P" means the species was present in
the ground layer, but occurred outside of the sampling quadrats.

                                         Fens
Species                  KLF       SVF          BF

Chara sp.                P         --           P
Lemna minor              --        3, 0.17, 52  4, 0.04, 11
Lemna trisulca           --        P            --
Potamogeton sp.          --        --           7, 0.07, 22
Ranunculus longirostrus  --        --           --
Riccia fluitans          --        3, 0.03, 24  --
Utricularia intermedia   --        3, 0.03, 24  22, 0.22, 67
Utricularia vulgaris     P         P            --
Utricularia sp.          --        --           --
Wolffia sp.              --        --           --

                                         Tall-shrub bogs
Species                  LCB           HB           THB       BLB

Chara sp.                --            --           --        --
Lemna minor              --            --           --        P
Lemna trisulca           --            --           --        --
Potamogeton sp.          P             --           --        --
Ranunculus longirostrus  --            --           --        --
Riccia fluitans          --            35, <1, 100  --        --
Utricularia intermedia   30, 1.39, 87  --           --        --
Utricularia vulgaris     4, 0.22, 13   --           P         --
Utricularia sp.          --            --           --        P
Wolffia sp.              --            --           --        --

                                      Leatherleaf bogs
Species                  DSB        LB         YB        BUR

Chara sp.                --         --         --        --
Lemna minor              --         <1, <1, 1  --        --
Lemna trisulca           --         --         --        --
Potamogeton sp.          --         P          --        --
Ranunculus longirostrus  --         P          --        --
Riccia fluitans          --         P          --        --
Utricularia intermedia   --         --         P         --
Utricularia vulgaris     --         36, 3, 99  --        P
Utricularia sp.          --         --         --        --
Wolffia sp.              --         P          --        --

                         Leatherlea           Forested peatlands
                           f bogs
Species                  SH        LAB           MPB         TAM (*)

Chara sp.                --        --            --          --
Lemna minor              --        9, 0.09, 100  --          --
Lemna trisulca           --        --            --          --
Potamogeton sp.          --        --            P           --
Ranunculus longirostrus  --        --            --          --
Riccia fluitans          --        --            --          --
Utricularia intermedia   P         --            --          --
Utricularia vulgaris     --        --            --          --
Utricularia sp.          --        --            --          --
Wolffia sp.              --        --            --          --

                            Forested
                           peatlands
Species                  RMB

Chara sp.                --
Lemna minor              8, 0.08, 100
Lemna trisulca           --
Potamogeton sp.          --
Ranunculus longirostrus  --
Riccia fluitans          --
Utricularia intermedia   --
Utricularia vulgaris     --
Utricularia sp.          --
Wolffia sp.              --
Table 8

Names, locations, and sources of vegetation data for 27 leatherleaf bogs
in the eastern United States. Acronyms Correspond with Figure 3.

Peatland name        Acronym  Region

Burket Bog             BUR    NE Indiana
Dutch Street Bog       DSB    NE Indiana
Leatherleaf Bog        LB     NE Indiana
Yost Bog               YB     North-central Indiana
Shoemaker Bog          SHB    NW Indiana
Fern Lake Bog          FLB    NE Ohio
Flatiron Lake Bog      FLT    NE Ohio
Triangle Lake Bog      TLB    NE Ohio
Portage Bog            POR    SW Lower Michigan
North Bog              NOR    SW Lower Michigan
South Bog              SOU    SW Lower Michigan
Pennfield Bog          PEN    SW Lower Michigan
Buck Hollow            BUC    SE Lower Michigan
Inverness Mud Lake     IML    N Lower Michigan
Bryant's Bog           BRY    N Lower Michigan
Dingman Bog            DIN    N Lower Michigan
Gate's Bog             GAT    N Lower Michigan
Mud Lake (open mat)    MLO    N Lower Michigan
Mud Lake (treed)       MLT    N Lower Michigan
Penny Lake Bog         PLB    N Lower Michigan
Green Pond Bog         GPB    S New York
Spruce Pond Bog        SPB    S New York
Tiny Cedar Pond        TCP    S New York
Uttertown Bog          UB     N New Jersey
Mishaps Bog            MB     N New Jersey
Lost Lake Bog          LLB    N New Jersey
Pequawket Bog          PQB    Central New Hampshire

Peatland name        source

Burket Bog           Present study
Dutch Street Bog     Present study
Leatherleaf Bog      Present study
Yost Bog             Present study
Shoemaker Bog        Present study
Fern Lake Bog        Andreas & Bryan (1990)
Flatiron Lake Bog    Andreas & Bryan (1990)
Triangle Lake Bog    Andreas & Bryan (1990)
Portage Bog          Brewer (1966)
North Bog            Keough & Pippen (1981)
South Bog            Keough & Pippen (1981)
Pennfield Bog        Crow (1969a)
Buck Hollow          Bailey (1967)
Inverness Mud Lake   Bevis (1960)
Bryant's Bog         Schwintzer (1981)
Dingman Bog          Schwintzer (1981)
Gate's Bog           Schwintzer (1981)
Mud Lake (open mat)  Schwintzer (1981)
Mud Lake (treed)     Schwintzer (1981)
Penny Lake Bog       Schwintzer (1981)
Green Pond Bog       Lynn & Karlin (1985)
Spruce Pond Bog      Lynn & Karlin (1985)
Tiny Cedar Pond      Lynn & Karlin (1985)
Uttertown Bog        Lynn & Karlin (1985)
Mishaps Bog          Lynn & Karlin (1985)
Lost Lake Bog        Lynn & Karlin (1985)
Pequawket Bog        Fahey & Crow (1995)
Table 9

Prevalent vascular plants from 27 leatherleaf bogs in six general
localities in the eastern United States: 1) northern Indiana (present
study), 2) southern Michigan (Bevis 1960, Brewer 1966, Bailey 1967, Crow
1969a, Keough & Pippen 1981), 3) northern Michigan (Coburn et al. 1932,
Schwintzer 1981), 4) northeast Ohio (Andreas & Bryan 1990), 5) southern
New York & northern New Jersey (Lynn & Karlin 1985), and 6) New
Hampshire (Fahey & Crow 1995). Number in parentheses is the number of
peatlands included for that region. Numbers in the table show the
percent of the peatlands that harbored the respective taxon. In regions
where only one bog was represented, a "P" indicates that a given species
was present.

                                1 (5)  2 (5)  3 (7)  4 (3)  5 (6)  6 (1)

Widespread species
 Acer rubrum                      100     80     17    100    100      P
 Andromedia glaucophylla           60     40     50     --     67      P
 Calla palustris                   --     20     --     67     67      P
 Chamaedaphne calyculata          100    100    100    100    100      P
 Drosera intermedia                40     20     17     33    100      P
 Drosera rotundifolia              40     40     33     67    100      P
 Gaylussacia baccata               20     --     17     33    100     --
 Larix laricina                    --     40     33     67     83      P
 Rhynchospora alba                 --     20     17     67     83      P
 Sarracenia purpurea               80     40     33     67    100      P
 Vaccinium macrocarpon             60     40     17    100    100      P
 Vaccinium oxycoccos               60     40     33     33    100      P
 Aronia melanocarpa                60     40     17     67     50      P

Northern species
 Eriophorum spissum                --     --     67     --     --     --
 Ledum groenlandicum               --     --     50     --     --     --
 Smilacina trifolia                --     --     33     --     --      P
 Vaccinium angustifolium           --     --     83     --     --     --
 Vaccinium myrtiloides             --     --     83     --     --     --

"Southern" species
 Cephalanthus occidentalis         80     --     --     --     33      P
 Decodon verticillata              40     20     --     67     83      P
 Dulichium arundinaceum            80     60     --     33     17      P
 Hypericum virginicum              80     20     --    100    100     --
 Toxicodendron vernix              80     20     --     67     50      P
 Vaccinium corymbosum              40     40     --    100    100      P
 Woodwardia virginica             100     60     --    100     17      P

Northern & Great Lakes species
 Betula pumila                     --     20     33     --     --     --
 Carex oligosperma                 60     40     67     --     --      P

Northern & northeastern
 species
 Carex trisperma                   --     --     50    100     67      P
 Kalmia polifolia                  --     --     67     --     50      P
 Picea mariana                     --     --     50     --     83      P

Northeastern species
 Carex canescens                   --     --     --    100    100      P
 Gaylussacia frondosa              --     --     --     --     67     --
 Kalmia angustifolia               --     --     --     --    100      P
 Peltandra virginica               --     --     --     33     83      P
 Picea rubens                      --     --     --     --     50     --
 Rhododendron viscosum             --     --     --     --    100     --

Southern Great Lakes species
 Carex rostrata                    80     --     --     --     --     --
 Iris versicolor                   60     --     --     --     --     --
 Nuphar advena                     60     --     --     --     --     --
 Scirpus cyperinus                100     40     --     --     --     --
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Article Details
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Author:Wujek, Daniel E.
Publication:Proceedings of the Indiana Academy of Science
Article Type:Statistical Data Included
Geographic Code:1U3IN
Date:Jan 1, 2001
Words:15890
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