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Paul S. Hlina1,†, Mary Ann E. Feist2, Derek S. Anderson1, Paul B. Marcum3, Reed J. Schwarting1, Nicholas P. Danz1 1Lake Superior Research Institute, University of Wisconsin–Superior

Barstow Hall 4 Belknap and Catlin Ave

P.O. Box 2000, Superior, WI 54880 2Wisconsin State Herbarium, University of Wisconsin-Madison Department of Botany, Birge Hall 430 Lincoln Drive, Madison, WI 53706

3Illinois Natural History Survey, Prairie Research Institute University of Illinois at Urbana-Champaign 1816 South Oak Street, Champaign, IL 61820


The vascular plants of the Bois Brule River watershed are listed, and over 160 years of change in plant communities observed is described. The watershed covers approximately 51,300 ha in northwestern Wisconsin, primarily in Douglas County with a short arm extending eastward into Bayfield County. The Bois Brule River travels southwest to the northeast 71 km from its headwaters and drains into Lake Superior. The diverse landscape supports boreal forest, northern mesic forest, northern wet-mesic forest, pine barrens, and other forested and non-forested communities. Five generalized changes in the watershed were noted: (i) the pine barrens community declined by more than 95%, (ii) the northern wet-mesic forest (dominated by Thuja occidentalis) immediately surrounding the river has been reduced to a narrow band, (iii) the large complex of conifer wetlands is greatly reduced, (iv) northern hardwood swamp (dominated by Fraxinus nigra), Alnus incana thickets, and the boreal forest in the lower reaches of the forest have been reduced and converted largely to timber production, and (v) old growth forest has been reduced to less than 1% of its pre-EuroAmerican settlement extent. A total of 839 vascular plant species have been documented in the watershed, 747 of them during our survey. Additionally, we documented 233 species new to the watershed, of which 53 are new county records and 13 are listed as endangered, threatened, or special concern in Wisconsin. This study has shown that the Bois Brule River watershed harbors a diverse assemblage of plants and is worth further conservation action. It is recommended that additional survey work continue in the future to inform and guide land managers.

KEYWORDS: Flora of Wisconsin, land cover change, pre-settlement conditions, Brule River Survey.

* Data sets used in this article are available upon request from the Lake Superior Research Institute at the University of Wisconsin-Superior, Superior, Wisconsin. † Author for correspondence ( Page  25 2020 THE GREAT LAKES BOTANIST


This is the second of two articles (the first of which is Hlina et al. 2020) reporting on a three-year project to re-survey and analyze vegetative and land use changes in the exceptional Bois Brule River watershed in northwestern Wisconsin over the last 160 years. The watershed of the Bois Brule River (hereafter referred to as the Brule River) exhibits an exceptionally diverse array of habitat types and outstanding water resources that support fish and wildlife species and numerous rare flora and fauna. The watershed is on the southwest side of Lake Superior, in northwestern Wisconsin. The watershed crosses through three ecoregions: Lake Superior clay plains, covering the northern third, the Bayfield sand plains from the southwest to the northeast and the Milles Lacs uplands to the west. The consistent flow of cold waters of the Brule River moving through hundreds of feet of outwash sandy plains to arise on the valley floor is a product of the cedar swamps in the headwaters and groundwater-connected springs. The vegetative cover of the Brule River watershed is exceptional, with large tracts of lowland forest at the region of the headwaters and old growth pine forest extending to Winneboujou, an unincorporated community located where County Highway B crosses the Brule River. The narrowing lower reaches of the watershed consist of boreal forest heavily influenced by Lake Superior.

For more than a century, the Brule River has been recognized as one of the premier trout streams of the Upper Midwest, with presidents of the United States and influential families fishing its waters and occupying its shores. By the late 1930s, managers, residents, and visitors noticed a significant decline in the fisheries, even after in-stream improvements and heavy stocking programs. These early observations led to the first comprehensive study of the Brule River watershed in northwest Wisconsin. As Schneberger and Hasler (1944) noted:

The need for an intensive study on this stream became evident when it was realized that during a five-year period extending from 1937 to 1941, a total of $34,247.67 was expended for the planting of fish and that stocking was not bringing about the desired results of maintaining or improving fishing.

The Brule River was studied from 1942–1945 and 1954 using a watershed approach. The Brule River Survey started in 1942 as a collaborative study conducted by the Wisconsin Conservation Commission, the University of Wisconsin- Madison, University of Wisconsin-Superior (then the Superior State Teacher’s College), and private citizens (Schneberger and Hasler 1942). The survey examined the physical, biological, and chemical characteristics of the Brule River watershed and resulted in a series of eleven papers: Bean and Thomson (1944), Churchill (1945), Evans (1945), Fassett (1944), Fischthal (1945), O’Donnell (1944), O’Donnell (1945), O’Donnell and Churchill (1954), Schneberger and Hasler (1942), Thomson (1944), Thomson (1945).

The survey’s objective was to pinpoint environmental disturbances that may be impacting fish populations. Botanists Norman Fassett and John Thomson conducted vegetative studies and floristic inventories of the Brule River valley and its forests, pine barrens, and wetland communities to document change and


provide baseline vegetative data. This work provided a critical view of the vegetation in the watershed in the early 1940s.

These earlier studies lend themselves to follow-up research to document vegetative changes, which we have now undertaken some seventy years later, the results of which are reported in this article. The objectives of this study were 1) to undertake a comprehensive floristic inventory of the forests of the Brule River watershed, focusing on complete species identification and voucher specimens,

2) to make qualitative summaries of floristic changes between our time and that of Fassett and Thomson, and 3) to characterize large-scale land cover changes in the watershed over three time periods: 1852–1856, 1932–1943, and 2014–2017. METHODS AND MATERIALS

Study Site

The Brule River watershed is in northwestern Wisconsin, primarily in Douglas County with a short arm extending eastward into Bayfield County. The watershed consists of more than 51,300 ha of forests, barrens, lakes, spring ponds, and the Brule River itself, which traverses 71 km, including 32 km as a steep river gorge draining into Lake Superior. The river channel is remarkable in that it flows northeastward in the ancient channel of the much larger Glacial Lake Duluth outlet that in glacial times (approximately 9,500–10,000 years ago) flowed southwesterly in what is now the St. Croix River (Clayton 1984; LaBerge 1994). The headwaters of both rivers are in the Divide Swamp, which is part of the Brule Glacial Spillway State Natural Area. The Brule River State Forest follows the river gorge and consists of 19,020 ha of forest lands managed by the State of Wisconsin for harvest, management and protection. There are four state natural areas in the state forest: Brule Glacial Spillway 1,070 ha, Mott’s Ravine 265 ha, Brule River Boreal Forest 263 ha and Brule Rush Lake 9 ha and 21 additional primary sites that offer further conservation potential within its boundaries (O’Connor 2016).

Between 2003 and 2016, more than 1,214 ha within the Brule River State Forest were removed from their earlier status as areas of priority conservation and restoration activities (O’Connor 2016). Simultaneously, the state of Wisconsin has recently increased timber production lands from 27% to 58% within the Brule River State Forest with most of the increase taken from the natural land’s designation (Wisconsin Department of Natural Resources 2017). These policy shifts have a high potential to reverse natural succession away from multi-age and old growth forest communities with high diversity towards an earlier successional sere, thereby creating a greater likelihood of a simplified forest, which already occurs on 35% of the watershed.


The general climate of the watershed is continental. Based on weather data from Gordon, Wisconsin (located at the southern end of the watershed), the mean annual precipitation is 81 cm. On average, July is the wettest month with a mean of 11.4 cm of precipitation. The mean annual temperature is 5.0°C; July is the warmest month with a mean of 19.9°C, and January is the coldest with a mean of –12.3°C. The length of the frost-free growing season has ranged from 45 days to 145 days with an average of 118 days per year (Midwestern Regional Climate Center 2017a).

In contrast, the area of the watershed to the north is influenced by Lake Superior. The large body of water provides an oceanic-like microclimate that moderates the climate, making winters warmer and summers cooler in the areas near the lake. According to weather data from Superior, Wisconsin (located to the west of the watershed and on Lake Superior), the mean annual precipitation is 78 cm. On average, September is the wettest month with a mean of 10.4 cm of precipitation. The mean annual temperature is 5.2°; July is the warmest month with a mean of 19.1°C and January is the coldest with a mean of –10.1°C. The length of the frost-free growing season has ranged from 64 days to 189 days with an average of 148 days per year based on data collected from 1981 to 2010 (Midwestern Regional Climate Center 2017b).

The forests on the Brule River watershed exhibit responses to three microclimates that dictate forest composition. In the lower reaches of the river where forest is restricted to a narrow steep val


ley and is influenced by the cooler temperatures of Lake Superior, boreal forest is found. Old growth coniferous bogs and swamps persist in the headwaters, where seepage springs deliver cool enriched mineral waters to a dense quagmire of Thuja occidentalis, Picea mariana, Larix laricina, and Alnus incana. Due to the foresight of early scientists and land managers, large tracts of this forest have been protected (Thomson 1945). The landscape rising above the middle reaches of the river exhibits yet a third microclimate, consisting of nutrient-poor, outwash glacial sand plains from the last glacial period (Sweet 1880; Clayton 1984; LaBerge 1994). In these pine barrens, drought resistant, fire-dependent species continue to shift through a mosaic of dwarf pine trees, scrub oaks, and open prairie. Today, most of this land consists of pine plantations (Pinus banksiana and P. resinosa) and is managed by private companies or state and county forest agencies.


The landscape of the watershed has been shaped by Precambrian lava flows, faulting, sedimentation, long epochs of erosion, and, finally, Pleistocene glaciation. Late in the Precambrian period, about 1.1 billion years ago, tectonic forces began rifting the North American craton (Laurentia), near present-day Lake Superior. The rifting extended 2,200 km southwest to present-day Kansas and 800 km southeast through present-day Michigan (Dott and Attig 2004). From these fissures, lava was released and spread across the region for millions of years. At the close of the Precambrian a shallow sea flooded the area laying down layers of sedimentary rocks. Over time these sediment deposits buried the underlying Keweenawan basalt. The sheer weight of the lava flows, and sediments formed the Lake Superior syncline, a down-warping of the earth’s surface (Laberge 1994). Faulting would occur numerous times along the syncline. The Douglas Fault would later up-thrust the underlying basalt and distort the Keweenawan sandstones, creating the steep river valley in the lower reaches of the watershed (Bean and Thomson 1944, LaBerge 1994).

While these Precambrian events provide the underlying structure for the Brule River valley, a recent period of glaciation called the Wisconsinan, shaped the landscape with deposition of outwash sands and glacial till. Clayton (1984) describes numerous epochs of the ice sheets advancing and retreating. The last phase of the Wisconsinan glacial period occurred 25,000 years ago with the advancement of the Superior lobe. It scraped its way down the length of the valley that was to become the Lake Superior basin. Around twelve thousand years ago, Glacial Lake Duluth began to form, roughly 180 m higher than present-day Lake Superior. Over the next one thousand years, the Superior Lobe started melting for the final time. The initial meltwater formed a channel flowing from the southwest to the northeast. A spillway would eventually form to the south called the Brule/St. Croix spillway (Dott and Attig 2004). This new spillway would drain Glacial Lake Duluth to the south. Eventually (approx. 9,500 years ago), the continual erosive force of meltwater carved a deep channel that resulted in the stream flow reversing and now flowing northeast back to Glacial Lake Duluth. When the glaciers fully receded, a divide formed out of which the Brule and St. Croix rivers flow in opposite directions today (Bean and Thomson 1944).

Based on this geology and subsequent deposit of sediments, the Brule River watershed can be split into three main sections starting at the southwest and moving northeast.

1) At the headwaters of the watershed, the gradient is very gentle and flat. The river meanders slowly and arises out of a complex of conifer swamps dominated by Thuja occidentalis, Larix laricina and Picea mariana. Surrounding this boggy lowland, at least 30.5 m of outwash sands were deposited by streams draining the melting waters (Clayton 1984). Springs in the headwaters region are direct indications of the groundwater flow resulting from these outwash plains. 2) In the middle section of the river, the sandy outwash is reduced and replaced with glacial till and occasional sandstone outcrops (Bean and Thomson 1944). The sandstone exposures provide many of the rapids and falls seen in this section of the river. Many of the lakes present today (e.g., Lake Nebagamon and Lake Minnesuing) were likely depressions on the landscape prior to glaciation and that filled once the melting ice retreated (Clayton 1984). Additionally, this area of the valley readily stores water in hundreds of small wetlands. 3) Finally, in its lower reaches, the river flows through lacustrine deposits of red clay accumulated in Glacial Lake Duluth (9,500 years ago) (Clayton 1984). As the ice melted and retreated, clay deposits mixed with iron oxide were exposed, giving the soils their characteristic red brick color. Clay soils are characterized by the small grain size, a high water-holding capacity, and an elevated cation- exchange with nutrients in the soil (U.S. Environmental Protection Agency 1980). These heavy soils are impermeable, poorly drained, high in nutrients, and remain cool and moist throughout the grow Page  28 28 THE GREAT LAKES BOTANIST Vol. 59

TABLE 1. Number of floristic inventory survey sites in each forest community type. Thomson sites are the sites selected in the forest community types surveyed by Thomson (1945). New sites are the additional sites first surveyed in the present study.

Forest Community Type Thomson Sites New Sites Total Boreal Forest 6 4 10 Northern Dry Forest – 5 5 Northern Dry-Mesic Forest – 6 6 Northern Hardwood Swamp 6 1 7 Northern Mesic Forest 2 7 9 Northern Wet Forest – 6 6 Northern Wet-Mesic Forest 3 9 12 Pine Barrens 4 2 6

ing season, which influences the species composition of the boreal forests they support (Epstein 2017).

Land Cover Analysis

To characterize the early vegetation of the Brule region, we used ArcGIS to digitize maps published by Fassett (1944) and Thomson (1945) that correspond to the time periods 1852–1856 and 1932–1943. We established a boundary of the current Brule River watershed based on recent elevation data using an automated watershed creation tool in ArcGIS. We cross-walked current land cover type information from Wiscland 2.0 data at a 600 dpi resolution (Wisconsin Department of Natural Resources 2016a) into the cover type categories used by Fassett (1944) and summarized the areas in each land cover type for all time periods as a way of inferring qualitative changes in the region. Strict area-based comparisons with early maps are subject to a moderate level of inaccuracy due to comparatively coarse tools used prior to the development of GIS; however, Thomson (1945) stated that the rates of error were below 7% for the early maps.

Floristic Inventories

Floristic inventories were conducted between May and September in 2015, 2016, and 2017 to document the flora of 61 sites, across eight terrestrial forest community types in the Brule River watershed: boreal forest, northern dry forest, northern dry-mesic forest, northern mesic forest, northern hardwood swamp, northern wet-mesic forest, northern wet forest and pine barrens (Table 1, Figure 1). These forest community types are described and classified in a Wisconsin Department of Natural Resources publication on Wisconsin’s natural communities (Epstein 2017). In 2015–2017, 21 sites were chosen and surveyed in forested community types originally surveyed by Thomson (1945). Although Thomson (1945) did not disclose his site locations, we used geographical and locational information indicated on his more than 500 herbarium voucher specimens, which are in the Donald W. Davidson Herbarium at the University of Wisconsin-Superior (SUWS) and the Wisconsin State Herbarium at the University of Wisconsin-Madison (WIS). Sites were further refined by inspecting forest stand compartment maps from the Wisconsin Department of Natural Resources, followed by an on-site visit to confirm representative types.

Thomson’s community species lists were created by assigning his herbarium specimens to specific forest community types. Additional species noted by Thomson (1944, 1945) but not represented by specimens were added to these community lists. Thomson combined northern wet-mesic and northern wet forest as conifer bog forest. We used his label information and our professional judgment to separate these communities into the northern wet and northern wet-mesic forest classification of today.

Forty additional sites were added to the study to provide a greater geographic representation of the eight forest types in the watershed. These sites were selected by viewing digital land cover maps to identify potential sites, followed by on-site visits to confirm representative habitats.

All sites were a minimum of five acres in size. Sites were selected if the following characteristics were evident: (i) boreal forest, a strong conifer component was present in the understory and the forest was approaching a mid-late successional sere; (ii) northern dry forest dominated by Pinus banksiana and P. resinosa, while devoid of P. strobus; (iii) northern dry-mesic forest, a component of


FIGURE 1. Brule river watershed. Survey sites are indicated by black dots, and the forest community type of each is indicated by the following codes: BF=boreal forest, NDF=northern dry forest, NDMF=northern dry-mesic forest, NHF=northern hardwood forest, NMF=northern mesic forest, NWF=northern wet forest, NWMF=northern wet-mesic forest, PB=pine barrens.


FIGURE 2. Northern wet-forest communities dominated by Picea mariana was one of the eight forest community types surveyed. Photo by Reed J. Schwarting.

Pinus strobus was found in the canopy, along with Pinus resinosa and/or Pinus banksiana as associates; (iv) northern hardwood swamps dominated by Fraxinus nigra; (v) northern mesic forest dominated by Acer saccharum or Tsuga canadensis with or without Betula alleghaniensis, Fraxinus americana or Tilia americana; (vi) northern wet forest dominated by Picea mariana or Larix laricina, with a canopy at least 6 m in height (Figure 2); (vii) northern wet-mesic forest dominated by Thuja occidentalis; and (viii) pine barrens consisted of mosaics of prairie-like openings, scrub oaks, and small stands of pines.

At each site, a floristic inventory was completed by a team of professional botanists from Illinois and Wisconsin. The team meandered through each site compiling a species list and collecting voucher specimens until no new species were observed (Figure 3). At the end of each survey, all observed species were assigned an abundance designation in one of four categories: abundant (A) = locally dominant species and those species that were widely distributed and often found growing in large quantities, common (C) = widely distributed and often found but not in abundant quantities, occasional (O) = not widespread but found in small numbers, rare (R) = rare to the site, only a small population or very few individuals found. These codes were applied to each site within a forest community type and then combined to create a complete species list for each community type. The abundance codes were applied subjectively by the team, but a conscientious effort was made for consistency. These descriptors are subjective estimates and should not be confused with quantitative cover values.

Voucher specimens were added to the archival Thomson collection housed at SUWS and the general herbarium collection at WIS (Figure 4). Duplicates were prepared and given to the Illinois Natural History Survey Herbarium (ILLS). Nomenclature primarily follows Voss and Reznicek (2012), which includes most Wisconsin species. Judziewicz et al. (2014) was used for all the grass species and for those species not included, the Flora of North America (Flora of North America Editorial Committee 1993+) was consulted.

Additional surveys were conducted at known and potential localities for rare and notable plants


FIGURE 3. Field team of Paul Marcum, Loy R. Phillippe, and Mary Ann Feist collecting voucher specimens to add to the Thomson archival collection. Photo by Derek S. Anderson.


FIGURE 4. Heuchera richardsonii. A July 1943 Brule River Survey collection by Dr. John Thomson found in the pine barrens community at the abandoned Volker farm field. It was preserved specimens like these that narrowed our designated sites for repeat surveys. Digital image from Univesity of Wisconsin-Superior, Superior.

of the watershed. Rare plants included any plant species listed on the Wisconsin Natural Heritage Working List (Wisconsin Department of Natural Resources 2016d). Notable species are uncommon species found for the first time in the watershed and may also be new to Douglas County. Rare and notable plant populations were identified and documented by recording latitude and longitude, population size, phenology, and site characteristics and by taking photographs.


We calculated various statistics to characterize the flora within each forest community type and made comparisons with similar calculations that we applied to Thomson’s data and voucher specimens for each forest community type (Thomson 1944, Thomson 1945). Such comparisons were made for all community types except northern dry forest and northern dry-mesic forest, which had few representative sites remaining in the watershed. We calculated average species richness per site and total species richness across all sites within each community as a basic measure of floristic diversity.

A floristic quality assessment (FQA) was used to provide an ecological condition assessment based on all species in the forest. This assessment was originally developed in the late 1970’s in the Chicago region to identify protection-worthy lands with a simple, repeatable, quantitative method (Swink and Wilhelm 1979). They assigned a Coefficient of Conservatism (C), a number from 0 to 10 for each species representing the Illinois flora. In the early 2000’s, Wisconsin’s expert botanists convened and did the same for Wisconsin’s vascular plants (Bernthal 2003). This list is now maintained by the Wisconsin Department of Natural Resources (Wisconsin Department of Natural Resources 2016c).

The FQA relies on two measurements, the Mean C and the Floristic Quality Index (FQI). The FQI is sensitive to the total hectares surveyed as species richness increases as the sample area increases (Matthews 2005). We were unable to determine Thomson’s sample areas in 1944–45 and chose the Mean C as a reliable metric in comparing the two time periods at the forest community scale. Species that are relatively tolerant of anthropogenic disturbance have low C-values, whereas species that are less tolerant of anthropogenic disturbance have high C-values (Spyreas 2019). We calculated a Mean Coefficient of Conservatism (C) for each forest type community. Mean C is the


arithmetic average of the C values across the total number of plant species observed (n) by the cumulative surveys by forest type. We calculated versions of C for all species (Ct) and for native species only (Cn). Since non-native species have a Coefficient of Conservatism of zero, Ct will always be less than Cn.


C = (C1+ C2+ C3 + ... + Cn) ÷ n

Further, we determined the percentage of non-native species, as well as the presence and abundance of prevalent ground layer species. In each forest community type, we classified prevalent ground layer species as those found in 80% or more of the surveyed sites with an abundance code of

(A) or (C) in at least 50% of them. Most prevalent ground layer species are those with an abundance code of (A) or (C) and occur in 80% or more of the surveyed sites. We used Sørensen Similarity Coefficients (Sørensen 1948) to identify sites that were compositionally dissimilar from others in their corresponding forest community type. The calculated coefficient ranges from 0 to 1, where a higher value indicates a higher percentage of shared species between the two sites. We calculated the coefficient for all pairs of sites within each forest community type. We removed sites in each community type where the coefficient was less than 0.4 from further comparisons. This process resulted in the removal of three sites that were likely misclassified, one each from the northern wet-mesic forest, northern hardwood swamp, and pine barrens communities.


Vegetation Cover Map Analysis (1852–2017)

As with all regions throughout the Midwest, land cover changes over the past 160 years have been substantial in the Brule River watershed. Today public lands encompass 60% (30,729 ha) of the watershed and include: Brule River State Forest, Douglas County Forestry and Parks and Bayfield County Forestry. The remaining 40% (20,570 ha) of the land in the watershed is controlled by private land holdings including timber management, logging companies and shoreline and streambank landowners. Based on comparisons of land cover maps across time periods (Table 2, Figure 5), we noted five general trends:

1. There was a shift from open pine barrens and dry forest to managed pine plantation in the southeastern extent of the watershed. These pine barrens declined from over 16,187 ha in 1852–1856 and 12,140 ha in 1932–43 to approximately 908 ha today. 2. Old-growth (northern dry-mesic forest) had been reduced by 75% of its pre-EuroAmerican settlement coverage by the 1940s. As a result of private and public protection of these old growth forests in land trusts and managed natural lands there has been a slight rebound with 2.6% (1,325 ha) of land represented in the watershed. 3. The early surveys depict a narrow strip of northern wet-mesic forest surrounding the Brule River from the headwaters area extending northeast up to Big Lake (Thomson 1945). Thomson (1945) noticed on-going harvesting in this forest in the 1940s that resulted in a substantial narrowing of this band. In 1945, the Brule River State Forest expanded its boundaries to include these headwater Page  34 34 THE GREAT LAKES BOTANIST Vol. 59

TABLE 2. Land cover area of historical community types and their current equivalents in three different time periods, illustrating vegetation changes in the Brule River watershed. The cover area for 1852–1856 and 1932–1943 were mapped originally in Fassett (1944) and are digitized for our comparisons here. The areas in the column headed 2014–2016 (Fassett) were determined using Wiscland

2.0 and cross-walked to Fassett’s cover types. The areas in the column headed 2014–2016 (Current) refer to Epstein’s (2017) natural communities’classification in discussing the eight forested communities in the watershed.

Historical Mapped Community Classification 1852–1856 Land Cover in hectares 2014–2016 1932–1943 (Fassett) 2014–2016 (Current) Current Community Classification Aspen 7609 15168 13529 13302 Northern hardwood aspen 226 Northern hardwood oak-maple Bog conifer 6702 3571 3401 971 Northern wet-mesic forest 1936 Northern wet forest 495 Muskeg Lowland hardwood 230 4988 5420 5420 Northern hardwood swamp Maple-yellow birch 924 192 750 750 Northern mesic forest Pine forest 3649 881 17293 2649 Northern dry-mesic forest 14644 Northern dry forest – pine plantation Pine-hardwood 6717 2096 4480 4480 Northern mixed/conifer hardwood Pine barren 16882 12285 908 908 Pine barren Spruce-fir forest 3413 432 1877 1877 Boreal forest Cleared 0 4010 2165 427 Developed 1725 Agriculture Willow, alders, etc. 0 59 574 273 Alder thicket 44.5 Northern wet meadow Marsh 85 13 Shrub carr 244 Open bog Open water 1490 1482 915 915 Open water Maple coppice 45 642 0 Small fir & aspen 0 1248 0 Totals 47054 47768 51312 51300


FIGURE 5. Land cover changes in the Brule River watershed from 1852 to 2016. Page  36 36 THE GREAT LAKES BOTANIST Vol. 59

forests, offering protection and management (Wisconsin Department of Natural Resources 2017).

4.A large conifer bog complex north of Lake Nebagamon was depicted on the 1852–1856 map. By the 1940s, there was a substantial decline from Thuja occidentalis to a lower quality forest of northern hardwood swamps dominated by Fraxinus nigra and Alnus incana. Remnants of the original forest are present, but in even smaller parcels. 5. Boreal forest extending from the mouth of the river towards the southwest has been greatly diminished, though recovery is occurring, and some stands are approaching old growth. Substantial forest lands were initially cleared for farming, while thousands of hectares have been placed into managed timber production consisting primarily of Populus spp., Quercus spp. and Acer spp. Forest Inventory Summaries

During his surveys in 1944 and 1945, Thomson collected 523 specimens representing 376 species. In this study, surveying in similar forest community types but with greater sampling effort, we collected over 2,200 specimens representing 747 species (16.6% of them non-native). A review of Thomson’s collections and other herbarium records revealed documented records of 92 species not found during our surveys. These are listed in Table 3. This brings the total of species documented for the watershed to 839. From our surveys, we recorded 53 new records for Douglas County (17% of them non-native species) (Table 4, Figure 6). Further, we collected 233 species (5% of them non-native) from the Brule River watershed for the first time. Three concurrent studies on the watershed were further referenced (Hlina et al. 2018a; Hlina et al. 2018b; Schwarting et al. 2018) in creating the comprehensive species list of the Brule River watershed in Appendix 1. An additional 18 species (Table 5) were seen and recorded in these studies but we failed to collect, and no historical herbarium records were found; therefore, these species are not included in the comprehensive species list (Appendix 1). Thomson vouchered enough species from the boreal forest, northern wet-mesic forest, pine barrens, northern hardwood swamps, northern mesic forest, and northern wet forest communities for us to compare his collection data with our observed and vouchered specimens.

Boreal forest averaged 132 species per site, with a range of 93–182 species. Dominant tree species include Populus tremuloides, Abies balsamea, Picea glauca, and Pinus strobus. Community richness was 66 families, 192 genera, 362 species of which 14.9% are non-native. The five most dominant families were Cyperaceae (11%), Asteraceae (10%), Rosaceae (8%), Poaceae (8%) and Ranunculaceae (5%). Twelve prevalent ground layer species were recorded (Appendix 2), with four species being most prevalent: Eurybia macrophylla, Maianthemum canadense, Pteridium aquilinum, and Aralia nudicaulis. Thomson’s community richness consisted of 38 families, 79 genera, and 105 species, 5% of which are non-native (Thomson 1945). Seventy-nine percent of prevalent species were also surveyed by Thomson. The value for Cn was 5.3, as compared to 4.9 for Thomson’s survey, while Ct was 4.5, as compared to 4.7 (Figure 7).


TABLE 3. Plant taxa previously documented from the Brule River Watershed but not encountered during the 2015–2016 survey. Non-native species are indicated with an asterisk (*). Tephroseris palustris is listed in Wisconsin as Special Concern and was last seen in the watershed by L.S. Cheney in 1897.

Family Species

PTERIDOPHYTES Cystopteridaceae Cystopteris tenuis Equisetaceae Equisetum ×ferrissii Lycopodiaceae Dendrolycopodium obscurum Lycopodiaceae Huperzia selago Lycopodiaceae Lycopodiella inundata Ophioglossaceae Botrychium lanceolatum Ophioglossaceae Sceptridium multifidum

DICOTS Amaranthaceae *Froelichia gracilis Anacardiaceae Rhus × pulvinata Apiaceae *Pastinaca sativa Apiaceae Sanicula odorata Araliaceae Aralia hispida Asteraceae *Achillea ptarmica Asteraceae Ambrosia psilostachya Asteraceae *Artemisia pontica Asteraceae *Artemisia vulgaris Asteraceae *Centaurea jacea Asteraceae Cirsium discolor Asteraceae *Crepis tectorum Asteraceae *Grindelia squarrosa Asteraceae Liatris ligulistylis Asteraceae Rudbeckia laciniata Asteraceae Symphyotrichum boreale Asteraceae Symphyotrichum pilosum Asteraceae Tephroseris palustris Betulaceae Alnus viridis Boraginaceae Lithospermum caroliniense Boraginaceae *Lithospermum officinale Brassicaceae *Erysimum cheiranthoides Brassicaceae Rorippa palustris Brassicaceae *Sisymbrium altissimum Brassicaceae *Thlaspi arvense Caprifoliaceae Symphoricarpos occidentalis Caryophyllaceae *Silene dichotoma Caryophyllaceae *Stellaria borealis Caryophyllaceae *Stellaria graminea Fabaceae Lespedeza capitata Fabaceae *Robina pseudoaccacia Fabaceae *Trifolium campestre Fabaceae *Trifolium pratense Fabaceae *Vicia villosa Gentianaceae Gentiana rubricaulis Lamiaceae *Ajuga genevensis Linderiaceae Lindernia dubia Molluginaceae *Mollugo verticillata Onagraceae Oenothera perennis Penthoraceae Penthorum sedoides Plantaginaceae *Linaria vulgaris

(Continued on next page)


TABLE 3. (Continued)

Family Species

Plantaginaceae Nuttallanthus canadensis Plantaginaceae Veronica peregrina Plantaginaceae *Veronica serphyllifolia Polygalaceae Polygala sanguinea Polygonaceae Polygonum achoreum Polygonaceae *Polygonum aviculare Portulaceae *Portulaca oleracea Ranunculaceae Clematis occidentalis Rhamnaceae Ceanothus americanus Rosaceae Agrimonia striata Rosaceae Aronia × prunifolia Rosaceae Crataegus succulenta var. macracantha Rosaceae *Filipendula rubra Salicaceae *Salix alba Urticaceae Urtica dioica


Alismataceae Alisma triviale Alismataceae Sagittaria cuneata Convallariaceae Maianthemum stellatum Cyperaceae Carex arcta Cyperaceae Carex cryptolepis Cyperaceae Carex houghtoniana Cyperaceae Carex lurida Cyperaceae Carex pellita Cyperaceae Cyperus lupulina Cyperaceae Eleocharis obtusa Cyperaceae Schoenoplectus pungens Cyperaceae Scirpus microcarpus Eriocaulaceae Eriocaulon aquaticum Hydrocharitaceae Elodea nuttallii Juncaceae Juncus balticus Juncaceae Juncus interior Juncaceae Juncus vaseyi Orchidaceae Calopogon tuberosus Orchidaceae Goodyera pubescens Orchidaceae Malaxis unifolia Orchidaceae Plantathera clavellata Poaceae Agrostis hyemalis Poaceae Alopecurus aequalis Poaceae Ammophila breviligulata Poaceae Dichanthelium columbianum Poaceae *Echinochloa crusgalli Poaceae Eragrostis hypnoides Poaceae *Poa annua Potamogetonaceae Stuckenia filiformis


TABLE 4. New vascular plant records for Douglas County. Non-native species are indicated by an asterisk (*). The Douglas County collection of Lactuca hirsuta is the second known collection from the state of Wisconsin.

Family Taxon

Asteraceae Bidens discoidea Asteraceae Helianthus hirsutus Asteraceae *Hieracium lachenalia Asteraceae Lactuca hirsuta Asteraceae Krigia biflora Asteraceae Pseudognaphalium obtusifolium Asteraceae Symphyotrichum urophyllum Boraginaceae *Myosotis arvensis Brassicaceae Cardamine pratensis var. palustris Caprifoliaceae Triosteum aurantiacum Caryophyllaceae *Arenaria serpyllifolia Caryophyllaceae *Dianthus barbatus Caryophyllaceae *Gypsophila muralis Caryophyllaceae Moehringia lateriflora Caryophyllaceae *Spergularia rubra Cyperaceae Carex alopecoidea Cyperaceae Carex backii Cyperaceae Carex bromoides var. bromoides Cyperaceae Carex echinodes Cyperaceae Carex hirtifolia Cyperaceae Carex muehlenbergii Cyperaceae Carex normalis Cyperaceae Carex ormostachya Cyperaceae Carex radiata Cyperaceae Carex rosea Cyperaceae Carex sprengelii Cyperaceae Carex tribuloides Cyperaceae Carex × knieskernii Droseraceae Drosera intermedia Ericaceae Pyrola americana Fabaceae Hylodesmum glutinosum Fabaceae *Vicia cracca Gentianaceae Gentiana alba Grossulariaceae Ribes lacustre Hemerocallidaceae *Hemerocallis fulva Juncaceae Juncus brachycephalus Lamiaceae Clinopodium vulgare Lythraceae Decodon verticillatus Onagraceae Oenothera clelandii Orobanchaceae Conopholis americana Plantaginaceae Plantago rugelii Plantaginaceae *Veronica longifolia Poaceae Bromus latiglumis Poaceae Bromus pubescens Poaceae Cinna arundinacea Poaceae Dichanthelium linearifolium Poaceae Dichanthelium oligosanthes Poaceae Elymus wiegandii Poaceae Torreyochloa pallida Potamogetonaceae Potamogeton nodosus Potamogetonaceae Potamogeton oakesianus Rosaceae Crataegus submollis Smilacaceae Smilax illinoensis


FIGURE 6. Conopholis americana, one of 53 new county records for Douglas County in northwestern Wisconsin. Photo by Derek S. Anderson.

TABLE 5. Species seen during this study but not vouchered and for which no herbarium records from the Brule River watershed are known. An asterisk (*) indicates a non-native species.

Family Species

Apiaceae Asteraceae Asteraceae Asteraceae Brassicaceae Brassicaceae Cyperaceae Cyperaceae Cyperaceae Fabaceae Lamiaceae Orchidaceae Poaceae Ranunculaceae Rosaceae Salicaceae Smilacaceae Vitaceae

Osmorhiza longistylis Erigeron philadelphicus Helianthus pauciflorus Heliopsis helianthoides Boechera grahamii Cardamine diphylla Carex blanda Carex laxiflora Cyperus schweinitzii Dalea purpurea

*Nepeta cataria Spiranthes cernua Elymus villosus Anemone acutiloba

*Pyrus communis Salix nigra Smilax ecirrhata Vitis riparia


FIGURE 7. Ct and Cn values compared with the Thomson data of 1944–45. There was no comparison data for the northern dry and northern dry-mesic forests communities, as these were not forest types that Thomson recognized.

Northern wet-mesic forest averaged 98 species per site, with a range of 67– 125 species. Dominant tree species include Thuja occidentalis and Abies balsamea. Community richness consisted of 64 families, 163 genera and 299 species, of which 4.3% are non-native. The five dominant families were Cyperaceae (12.3%), Asteraceae (8%), Rosaceae (7%), Ericaceae (6%) and Poaceae (5%). Eleven prevalent ground layer species were recorded (Appendix 2), with five species being most prevalent: Rubus pubescens, Coptis trifolia, Maianthemum canadense, Trientalis borealis, and Cornus canadensis. Thomson’s community richness consisted of 40 families, 71 genera, and 83 species, of which 3% were non-native (Thomson 1945). Eighty percent of prevalent species we recorded were also surveyed by Thomson. The value for Cn was 6.1, as compared to 6.9 for Thomson’s survey, while Ct was 5.7, as compared with 6.7 (Figure 7).

Pine barrens averaged 87 species per site, with a range of 62–111 species. Dominant tree species included Pinus resinosa and Pinus banksiana. Community richness consisted of 54 families, 137 genera, and 207 species, of which 16.4% are non-native. The four dominant families were Asteraceae (20%), Poaceae (13.8%), Rosaceae (10.5%), and Cyperaceae (5.7%). Sixteen prevalent ground layer species were recorded (Appendix 2), with eleven species being most prevalent: Comptonia peregrina, Corylus americana, Prunus pumila, Rubus flagellaris, Carex pensylvanica, Vaccinium angustifolium, Quercus macrocarpa, Quercus ellipsoidalis, Andropogon gerardii, Danthonia spicata, and Monarda fistulosa. Thomson’s community richness consisted of 39 families, 94 genera, and 127 species, of which 17% were non-native (Thomson 1945). Sixty-three percent of prevalent species recorded, were also surveyed by Thom


son. Cn was 4.8, compared to 4.9 from Thomson’s survey. Ct was 4.1, compared to 4.0 (Figure 7).

Northern hardwood swamp averaged 93 species per site, with a range of 71– 130 species. Dominant tree species included Fraxinus nigra and Thuja occidentalis. Community richness consisted of 66 families, 170 genera, and 307 species, of which 10.7% are non-native. The four dominant families were Cyperaceae (13%), Asteraceae (9%), Poaceae (8%), and Rosaceae (7%). Ten prevalent ground layer species were recorded (Appendix 2), with four species being most prevalent: Carex stipata, Rubus pubescens, Carex intumescens, and, Onoclea sensibilis. Thomson’s community richness consisted of 33 families, 47 genera, and 52 species with no non-native species (Thomson 1945). Twenty-five percent of our prevalent species were also surveyed by Thomson. Cn was 5.6 compared to 4.9 in Thomson’s. Ct was 5.0 compared to 4.9 (Figure 7).

Northern wet forest averaged 36 species per site, with a range of 23–71 species. Dominant tree species included Picea mariana and Larix laricina. Community richness consisted of 44 families, 93 genera, and 156 species, of which 6.4% were non-native. The five dominant families were Cyperaceae (18%), Rosaceae (11%), Ericaceae (7%), Asteraceae (6%), and Poaceae (6%). Four prevalent ground layer species were recorded (Appendix 2), with two species being the most prevalent: Rhododendron groenlandicum and Maianthemum trifolium. Thomson’s community richness consisted of 16 families, 25 genera, and 34 species with no non-native species (Thomson 1945). One hundred percent of our prevalent species were also surveyed by Thomson. Cn was 6.1, compared to

7.0 in Thomson’s survey. Ct was 5.7, compared to 7.0 (Figure 7). Northern mesic forest averaged 85 species per site, with a range of 42–144 species. Dominant tree species included Acer saccharum, Tilia americana, and Acer rubrum. Community richness consisted of 59 families, 147 genera, and 242 species, of which 10% were non-native. The five dominant families were Cyperaceae (11%), Poaceae (9%), Asteraceae (8%), Rosaceae (7%), and Ranunculaceae (6%). Six prevalent ground layer species were recorded (Appendix 2), with three species being most prevalent: Acer saccharum, Maianthemum canadense, and Clintonia borealis. Thomson’s community richness consisted of 31 families, 45 genera, and 59 species, of which 3% were non-native (Thomson 1945). Eighty-three percent of our prevalent species were also surveyed by Thomson. Cn was 5.6 for both time periods. Ct was 4.9, compared to 5.5 in Thomson’s survey (Figure 7).

Although we could not do comparisons for the remaining forest types, we provide summary data from our surveys for two additional forest types. Northern dry forest averaged 75 species per site, with a range of 45–111 species. Dominant tree species included Pinus resinosa and P. banksiana. Community richness consisted of 51 families, 134 genera, and 209 species, of which 15.8% were non-native. The five dominant families were Asteraceae (11%), Poaceae (11%), Rosaceae (11%), Cyperaceae (6%), and Ericaceae (6%). Seven prevalent ground layer species were recorded (Appendix 2), with three species being most prevalent: Maianthemum canadense, Pteridium aquilinum, and Oryzopsis asperifolia. Cn for all northern dry forest species was 5.2, while Ct was 4.4 (Figure 7).


Northern dry-mesic forest averaged 94 species per site, with a range of 39– 128 species. Dominant tree species included Pinus resinosa, P. banksiana, P. strobus, and Betula papyrifera. Community richness consisted of 65 families, 163 genera, and 263 species, of which 12.9% were non-native. The five dominant families were Asteraceae (10%), Rosaceae (9%), Poaceae (8%), Cyperaceae (6%), and Ericaceae (5%). Seven prevalent ground layer species were recorded (Appendix 2), with four species being most prevalent: Maianthemum canadense, Pteridium aquilinum, Aralia nudicaulis, and Eurybia macrophylla. Cn was 5.5, while Ct was 4.8 (Figure 7).

The following species were documented by Thomson (1945), but not found by our 2015–2017 surveys: Agrimonia striata, Alnus viridis, Botrychium lanceolatum, B. matricariifolium, Botrypus virginianus, Calopogon tuberosus, Carex retrorsa, Celastrus scandens, Clematis occidentalis, Geum fragarioides, Grindelia squarrosa, Liatris ligulistylis, Nuttallanthus canadensis, Osmorhiza longistylis, Rudbeckia laciniata, Symphoricarpos occidentalis, Symphyotrichum boreale.

Among various plant growth forms, graminoid species displayed substantial proportional compositional increases between the two time periods (1945, 2017) in the boreal forest, northern hardwood swamp and northern wet-mesic forest, while there was only a slight increase of graminoids in the northern mesic forest. There was no change in plant growth forms for the pine barrens and northern wet forest (Table 6).

Appendix 3 summarizes these details for all forest types surveyed.

Rare and Notable Species

Between 2015 and 2017, we found thirteen species listed as rare in Wisconsin (Wisconsin Department of Natural Resources 2016c): Asclepias ovalifolia (Figure 8), Callitriche hermaphroditica, Calypso bulbosa, Carex backii, Coptidium lapponicum, Cypripedium parviflorum var. makasin, Eriophorum chamissonis (Figure 9), Geum macrophyllum var. macrophyllum, Petasites frigidus var. sagittatus, Pyrola minor, Ribes oxyacanthoides, Rhynchospora fusca, and Vaccinium vitis-idaea. Six rare species were found in the northern wet-mesic forest, four in the boreal forest, one in the pine barrens, one in the northern hardwood swamp and one in the northern wet forest. These areas may be the last refuge in the Brule River watershed, if not in the State of Wisconsin, for the species Coptidium lapponicus and Calypso bulbosa and many other rare plant assemblages. An additional ten notable species with low occurrences were added to the list (Table 7).


We could not determine the exact methods that Thomson (1944, 1945) used to survey the flora of each of these communities. Upon close examination of his herbarium specimens and his papers (Thomson, 1944; Thomson 1945), it


TABLE 6. Number of species of five growth forms represented in species lists for 1944–1945 and 2015–2016 in each of the eight forest community types. Data for 1944–1945 are omitted for northern dry forests and northern dry-mesic forests due to lack of historical data.

1944–1945 2015–2016 Growth Form Number of Species Percentage (%) Number of Species Percentage (%) Boreal forest Tree 16 15 25 7 Shrub 21 21 62 17 Forb 57 54 198 55 Graminoid 8 8 70 19 Vine 3 3 7 2 Totals 105 100 362 100 Northern wet-mesic forest Tree 6 7 19 6 Shrub 14 17 57 19 Forb 55 67 162 54 Graminoid 8 9 57 19 Vine 0 0 4 2 Totals 83 100 299 100 Pine barrens Tree 7 6 13 6 Shrub 19 15 28 14 Forb 81 64 121 58 Graminoid 18 14 43 21 Vine 1 1 2 1 Totals 126 100 207 100 Northern hardwood swamp Tree 8 15 20 7 Shrub 7 13 51 16 Forb 33 64 167 54 Graminoid 2 4 64 21 Vine 2 4 5 2 Totals 52 100 100 Northern mesic forests Tree 8 13 18 7 Shrub 8 14 44 18 Forb 34 58 126 53 Graminoid 8 14 51 21 Vine 1 1 1 1 Totals 59 100 242 100

(Continued on next page)


TABLE 6. (Continued)

1944–1945 2015–2016 Growth Form Number of Species Percentage (%) Number of Species Percentage (%) Northern wet forest Tree 7 21 13 8 Shrub 9 26 38 24 Forb 10 29 65 42 Graminoid 8 24 40 26 Vine 0 0 0 0 Totals 34 100 156 100 Northern dry forest Tree 15 7 Shrub 43 21 Forb 105 50 Graminoid 46 22 Vine 0 0 Totals 209 100 Northern dry-mesic forest Tree 24 9 Shrub 51 19 Forb 143 54 Graminoid 41 16 Vine 4 2 Totals 263 100

appears that Thomson may have had some bias towards collecting along roadways and in abandoned fields. Furthermore, Thomson collected on his own on 53 field days spanning a three-year period, whereas our research teams consisted of groups of surveyors and professional botanists surveying the equivalent of 180 field days spanning a three-year period. The level of effort far exceeded Thomson’s and has likely contributed to the discrepancy between the measurements of species richness, species composition, and species abundance for the two time periods.

When examining changes in the function, structure, and growth of the groundcover stratum, a clear pattern emerges. First, the graminoids have become more dominant throughout several forest types (Table 6). Recent authors (Rooney and Waller 2003; Rooney 2009; Burton et al. 2014) suggest that graminoids are better able to recover from browsing pressure than most forbs, which leads to their increased dominance in today’s forests because of an increase in herbivory by white-tailed deer. A second trend observed was an increase in the total number of tree species in most forest types (Table 6), while species percentages as a portion of forest composition remained the same across all community types. A likely explanation is the exhaustive nature of our studies


FIGURE 8. Asclepias ovalifolia in a population of more than 200 individuals located in a recently harvested pine plantation in the pine barrens community type. Photo by Derek S. Anderson.

FIGURE 9. Eriophorum chamissonis in a population of hundreds found in one small bog near the shores of Lake Nebagamon in the western portion of the watershed. Photo by Reed J. Schwarting.


TABLE 7. Rare, threatened, endangered species in the watershed. Also included in the list are notable species which are uncommon species found for the first time in the watershed and may also be a new Douglas County record. In the State Status column, SC = Special Concern, THR =Threatened, END = Endangered, N = Notable.

Number of Year Last State Occurrences Observed Status Comments Asclepias ovalifolia Callitriche hermaphroditica Calypso bulbosa var. bulbosa 1 4 2 2015 2016 2016 THR SC THR 100 plants cold spring waters 125 plants in 1996; Carex assiniboienensis Carex backii Carex vaginata Carex × knieskernii Coptidium lapponicus Cypripedium parviflorum var. makasin Cypripedium parviflorum var. pubescens Cypripedium reginae Dryopteris fragrans Eriophorum chamissonis 1 2 Several 1 3 3 1 2 1 1 2016 2015 2016 2016 2015 2016 2016 2016 2015 2017 N SC N N END SC N N N SC two plants in 2016 boreal tributary ravine bedrock glade scattered several plants >250 plants numerous numerous two locations rock outcropping hundreds of plants in Epilobium palustre Gentiana alba 1 1 2017 2017 SC N one bog one plant several plants on riverbank Geum macrophyllum var. macrophyllum Huperzia selago Lactuca hirsuta 1 1 1 2016 1996 2015 SC SC N one plant a few dozen Platanthera huronensis 1 2015 N Pyrola minor Petasites frigidus var. sagittatus Rhynchospora fusca Ribes oxyacanthoides Taxus canadensis Tephroseris palustris Vaccinium vitis-idaea 1 3 1 1 3 1 1 2015 2015 2016 2017 2015 1897 2015 END THR SC THR N SC END unknown several populations hundreds of plants several plants several plants 70–100 plants

and the large percentage of trees that were found at the seedling stage; these seedlings may have been discounted or overlooked by Thomson.

We observed 124 non-native species throughout all the forest communities we surveyed. Of these, Rhamnus cathartica poses the greatest threat to the lower reaches of the boreal forest, growing in thickets that replace native understory shrubs and forbs. Iris pseudacorus is found from the southern edge of Big Lake to the mouth of the river. There is an active invasive plant management program on Lake Minnesuing, and to a lesser extent on Lake Nebagamon, for controlling these populations, while the Brule River community is just becoming aware of the threat. Myosotis scorpioides and Nasturtium officinale are naturalized from near the headwaters to the mouth of the river and have likely replaced some native aquatic plant populations (e.g., Ranunculus aquatilis, Callitriche palustris, and Callitriche hermaphroditica). Bromus inermis, Phalaris arundinacea, and Centaurea stoebe are found growing along disturbed road, trails, ditches, parking


areas, and other disturbed areas, but the populations are well managed. Two emerging invasive plant threats to the watershed include the recent appearance of small populations of Sorbus sorbifolia and Valeriana officinalis that were found embedded in both the boreal forest and the northern wet-mesic forest. Lastly, a private landowner informed us of a large population of Berberis thunbergii planted decades earlier within the watershed, but our survey teams did not observe this species, and as a result it is not included on our list in Appendix 1.

Boreal Forest

By the time of the Thomson survey, the boreal forest was greatly diminished due to cutover with early successional species such as Abies balsamea and large stands of Populus spp. present. Boreal forest occupied approximately 6.9% of the watershed (Fassett 1944). Dominant species were Betula papyrifera, Populus grandidentata, Populus tremuloides, and Prunus pensylvanica. Ground cover species of importance were Eurybia macrophylla, Diervilla lonicera and Pteridium aquilinum (Thomson 1945).

The present-day boreal forest covers 3.6% (1,877 ha) of the watershed and is found north of the Copper Range. It is characterized as gradually sloping to the northeast within a gentle terrain, bisected by numerous steep ravines. A contiguous second- and third-growth aspen forest continues to dominate the forest today with some Abies balsamea and Picea glauca in the understory (Wisconsin Department of Natural Resources 2016b). Other areas have lost timber due to an alteration of the hydrology caused by heavy equipment resulting in swamping that favors thickets of Alnus incana and Salix spp.

Boreal forest Ct values indicate a slight decrease in floristic quality. Thomson did not discover as many non-native plants in these early successional forests (only 5%) as we did in today’s boreal forest (14.9%). Thomson classified the boreal forest as “The Aspen Association,†since the boreal forest composition was greatly compromised from early logging activities and was dominated by hardwoods in the 1940s (Fassett 1944, Thomson 1945). Family dominance has changed from Rosaceae, Asteraceae, Ranunculaceae, and Salicaceae in the 1940s to Cyperaceae, Asteraceae, Rosaceae, Poaceae, and Ranunculaceae today. The graminoids represent almost 20% of the boreal forest flora today, in contrast to only 7% in the 1940s. Many earlier inventories often overlooked graminoid species and that may explain the changes we observed. Another observation is that insect pollination dependent families (e.g., Rosaceae and Ranunculaceae) have declined, while wind-pollinated families (e.g., Cyperaceae and Poaceae) have substantially increased. These findings are consistent with the trends found during a project that re-surveyed the 1959 baseline data collected by John T. Curtis for selected northern forests in Wisconsin (Rooney et al. 2004).

The best examples of remnant boreal forest stands are along steep ravines near the mouth of the river and extending inland for several miles. The cooler climate and red clay soils dictate which tree species can be sustained in this part of the watershed. In these areas we see the returning prominence of Pinus strobus, Abies balsamea, and Picea glauca, while Betula papyrifera has decreased in abundance (Hlina et. al. 2018a). In the deep creek ravines along tributaries, dom


inant trees include Thuja occidentalis, Abies balsamea, Picea glauca, and Populus tremuloides. The boreal forest on the Brule River is slowly recovering, and opportunities for restoration are high. In the Biotic Inventory of the Brule River State Forest O’Conner (2016) states:

The Brule River State Forest offers the single best opportunity for clay plain boreal forest restoration on state-owned land on the entire Superior Coastal Plan Ecological Landscape and possibly North America.

Such restoration will ensure the continued biodiversity, complexity, and health of this rare plant community in Wisconsin for future generations.

Northern Wet-Mesic Forest

As noted earlier, Thomson (1945) lumped northern wet forest and northern wet-mesic forest into one plant community type, which he called conifer bog. The dominant species he noted were Abies balsamea, Larix laricina, Picea mariana, Thuja occidentalis, and Sphagnum spp. He noted two shrubs, Chamaedaphne calyculata and Rhododendron groenlandicum, as significant components of the understory. The coverage of this forest was 6.8% during his survey (Fassett 1944).

In the present day, northern wet-mesic forest encompasses 967 ha, approximately 2% of the watershed in the headwater region of the Brule River and is dominated by old-growth even-aged stands of Thuja occidentalis. From 1942 to 1945, Thomson observed that area residents and farmers were harvesting the cedar in this area for fence posts and other uses, exposing the sphagnum hummocks and pools to wind and sun resulting in severe desiccation and recommended that all cutting cease (Thomson 1945). Thomson recognized the importance of this area and the direct influence it had on sustaining water quality, water flow, and stable temperatures needed to support brown, brook, and rainbow trout populations in the river. This forest is now approaching 200 years of age and exhibits little apparent regeneration in the last 70–80 years. Beals et al. (1960) and others (Alverson et al. 1988; Van Deelen 1999; Rooney et al. 2002; Forester et al. 2008) have documented the impact of deer populations on the ability of Thuja occidentalis to regenerate. Due to the lack of regeneration, the Thuja occidentalis swamps, which are the largest in the state, are vulnerable to disappearance in the next 50–75 years (Bushman 2006; Scheller and Mlandenoff 2005, 2008). Based on our survey data and observations, this forest is most likely to be replaced by Abies balsamea and Acer rubrum trees with large patches of Alnus incana thickets.

Northern wet-mesic forest Ct values depict a decrease in floristic quality from

6.9 to 5.8 and in Cn values from 7 to 6.1. Non-native species slightly increased temporally, from 3% to 4.3%, but this alone does not explain the discrepancy in floristic quality between the two time periods. As was standard in the 1940s, and as noted earlier, Thomson classified both the northern wet forest (Picea mariana and Larix laricina dominant) and northern wet-mesic forest (Thuja occidentalis dominant) as one community type called conifer bog, though the former has a greater percentage of conservation species. In Vegetation of Wisconsin, Curtis Page  50 50 THE GREAT LAKES BOTANIST Vol. 59

(1959) compares these two-forest community types and finds only a 50% similarity between them, and therefore separated them into northern wet forest and northern wet-mesic forest. This difference perhaps explains the wide differences in C values. Family dominance has also changed from Cyperaceae, Orchidaceae, Caprifoliaceae, Rosaceae, and Ericaceae to Cyperaceae, Asteraceae, Rosaceae, Poaceae, and Ericaceae. Graminoid dominance has increased from 11% to 17%. Surprisingly Maianthemum canadense, Aralia nudicaulis, and Acer spicatum, which are highly prevalent today, were not recorded in the 1940s. It is possible that these species were simply overlooked. Another substantial decline was found in Orchidaceae, dropping from 6.0% to 2.9%. Three orchid species that were present in the 1940s but are absent today are Calopogon tuberosus, Goody- era pubescens, and Spiranthes cernua. Calypso bulbosa, a state threatened orchid, is probably near extinction in the Brule River watershed (E. J. Judziewicz, personal communication 2016). Judziewicz studied the same cedar swamps in the mid-to late 1990s and found hundreds of individuals of Calypso bulbosa (Epstein et. al. 1999). During the field seasons of 2015 and 2016 only four plants were found, two of which were sterile and two in flower. Moreover, Rooney and Waller (2003) and Rawinski (2008) described the effects of high deer densities on the understory flora of forested ecosystems, which may explain some of this decline, as well as a drop in the Ct values of northern wet-mesic forest.

Pine Barrens

Pine barrens were frequently burned and were mostly treeless at the time of the Brule River surveys of the 1940s; the last big fire occurred in 1936 (Fassett 1944). Thomson (1945) and Fassett (1944) documented a much more intact pine barrens than we find today, noting many characteristics such as shrubby jack pine, scattered red pine savannas, and vast open barrens. They reported the pine barrens covered 25.7% of the total watershed in 1938; today pine barrens cover approximately 1.8% (908 ha). Lost to history is the full extent of the flora of the pine barrens as it existed in 1854. The land that was historically pine barrens is now managed as pine plantation monocultures predominantly by private timber companies, the Brule River State Forest, and Douglas and Bayfield counties. The C-values show little difference between the 1940s and now and may indicate that the pine barrens remain a community in recovery.

Thomson’s (1945) description of the Pinus banksiana community distinguished between two paths of succession, both essentially leading to a Pinus banksiana, P. resinosa, Quercus spp. community based upon the intensity and intervening time between fires. He noted the dominant woody plants in scattered stands as Pinus banksiana and Quercus ellipsoidalis. Ground cover species of importance were Corylus americana, Danthonia spicata, Comptonia peregrina, Pteridium aquilinum, Quercus macrocarpa, Salix humilis, and Vaccinium angustifolium. (Thomson 1945). Many associate species are prairie species adapted to drier conditions, low soil fertility, and fire.

Pine barrens Ct and Cn values show little change between the two time periods. These values are low relative to other forest types and may indicate an overabundance of species tolerant of anthropogenic disturbances with low C values


in Asteraceae and Poaceae. These species accounted for more than 30% of the flora. Pine barrens of today are embedded in large acreages of pine plantation, resulting in the highest percentage of non-native species (16.4%) of all the communities surveyed. Family dominance has remained the same (Asteraceae, Poaceae, and Rosaceae). Hieracium aurantiacum (orange hawkweed) is the only non-native species that ranked high enough to make the prevalent ground layer species list for any community in the Brule River watershed. Another interesting observation is the successful spread of Carex pensylvanica, a native sedge that was not recorded by Thomson, though its presence in other adjacent forested communities may explain its presence in the pine barrens.

Due to the smaller open areas and savanna-like conditions, the globally rare pine barrens are home to a wide variety of wind-dispersed non-native plants (e.g., Agrostis gigantea, Centaurea stoebe, and Cirsium arvense). Fire suppression activities have altered the pine barrens landscape by eliminating large patches (950–1500 ac) of open habitat (Radeloff 2000; Grossman and Mladenoff 2007; Scheller and Mladenoff 2008). It is doubtful that the historical shifting mosaic of oak and pine savanna surrounded by large open patches will be achieved without further use of fire and other management tools by land managers. Open barrens in the watershed occur only in small parcels today, and not as the vast terrain of yesteryear, that numerous wildlife species such as sharp- tailed grouse, bobolink, and savannah sparrow require (Radeloff et al.1999). The best remaining examples of pine barrens in the Brule River watershed are found at Mott’s Ravine State Natural Area and in patches between forests of recent harvests. It was at Mott’s Ravine that Lactuca hirsuta was found, only the second collection of this species for Wisconsin. The 2003 Brule River State Master Plan includes plans to modestly increase the acreage of pine barrens found at Mott’s Ravine (Van Horn et al. 2003).

Northern Dry Forest

Northern dry forest prior to extensive landscape disturbance was comprised of large stands of Pinus resinosa interspersed within pine barrens. The forest covered approximately 12% of the watershed. It was quickly harvested during the lumbering activities in the late 19th century and converted to jack and red pine plantations by the late 1930s. The five stands we surveyed were either Pinus resinosa plantations greater than 100 years old or small pockets of older P. resinosa and P. banksiana intermixed with scrub oak (Quercus macrocarpa and

Q. ellipsoidalis). In the barrens area of the watershed, these forest plantations are heavily represented and occupy 28.5% (14,614 ha) of the watershed. Thomson treated this forest as a successional stage within the pine barrens complex and hence one-to-one comparison was not possible with our data. This forest, as expected, had one of the lowest diversities of all the communities and a higher level of non-native species compared to the other communities. The understory of the northern dry forest is dominated by Acer rubrum and Abies balsamea saplings and hazelnut (Corylus spp.), with pine regeneration not apparent. The Brule River State Forest has identified three primary sites for northern dry forest needing protection (Wisconsin Department of Natural Resources 2017). These sites Page  52 52 THE GREAT LAKES BOTANIST Vol. 59

are exceptionally small and subject to catastrophic damage from extreme weather events. Opportunities for management, including the use of fire to increase open conditions for pine seedlings, are suggested in the 2016 Biotic Inventory of the Brule River State Forest (O’Connor 2016).

Northern Dry-Mesic Forest

At time of settlement these valuable northern dry-mesic forests encompassed 7.8% of the watershed and by the 1940’s had declined to 1.8% of the area. Today, this forest type has slightly rebounded and occupies approximately 2.6% (1,325 ha) of the watershed, with most old growth northern dry-mesic forest in land trust stewardship. Due to generations of human activity with the creation of trails, roads, homes, outbuildings, and earlier landscape plantings, a suite of nonnative species was found. Except for Berberis thunbergii, none of these species have compromised the ecological integrity of this old growth forest. Berberis thunbergii, which was planted in the area as an ornamental, has increased its spread in recent years into the surrounding forest. Species richness remains high (263 species), but much of the gain compared with Thomson’s survey has come from non-native species (12.9% of the current total). Acer rubrum and Abies balsamea seedlings are dominant in the understory and are poised to become the replacement forest in the absence of Pinus strobus and P. resinosa regeneration. Though floristic richness remains high, 40% of species in this forest were found only once during the surveys of this forest type and at a relatively low abundance.

Northern Hardwood Swamp

Fassett (1944) and Thomson (1945) defined northern hardwood swamp as a lowland hardwood association, a minor component of the watershed, occupying only 6.7% of the total watershed area. Thomson (1945) noted that the dominant tree species are Acer rubrum, Fraxinus nigra, and Ulmus americana. Associated woody species included: Abies balsamea, Alnus incana, Betula papyrifera, and Populus balsamifera. Ground cover species were numerous and many of the species commonly associated with wet habitats, such as Carex tuckermanii, Carex crinita, Scutellaria lateriflora, Galium trifidum, and Micranthes pensylvanica were observed. Surprisingly, only 25% of prevalent ground layer species were common to both surveys. The discrepancy might be explained through the diversity of plant associations we observed in the thirteen northern hardwood swamp sites surveyed by our team.

Today, a large area of high-quality northern hardwood swamp exists in the watershed covering 10% (5,420 ha) of the watershed. The present-day northern hardwood swamp community has 307 species, of which 10.7% are non-native, which is in the lower range among the communities surveyed. This community harbored the greatest number of Carex species (32), as well as large numbers of shade tolerant specialists. By far the greatest threat to these lowland swamps is the invasion of the emerald ash borer that causes high mortality in all species of Fraxinus. Our survey did not find any evidence of the beetles. Likewise, no signs


of the beetle have ever been found in the Brule River State Forest (D. Schulz, personal communication 2017).

Based on our survey results, it is likely that these lowlands, if compromised by the beetle, may be replaced by Acer rubrum, Abies balsamea, and the shrub Alnus incana, as these species are the most well-represented woody species in understory layers. This could have a profound effect on the structure, function, and ecological integrity of this forest.

If Fraxinus nigra suffers increased mortality from an emerald ash borer invasion, it is likely that severe hydrological changes will occur in this wetland forest. Slesak et al. (2014) showed that in an infected ash forest, flood conditions lasted six to eight weeks longer, which would have serious consequences to the establishment of other tree species and the groundcover vegetation that grows based on hydrological regimes (saturation, very wet, wet, and moderately wet). In the Brule River region, these new conditions may favor weedy aquatic and wet meadow graminoid species, such as Phalaris arundinacea, Typha spp., and Phragmites australis that are able to easily colonize from adjacent landscapes. Some long-term projects evaluating the success of anticipated replacement species (e.g., Quercus bicolor, Celtis occidentalis) have been conducted or are underway in similar regional forests (Looney et al. 2015, Rooney et al. 2015). These two species remain south of the tension zone in Wisconsin and still have no presence in the Brule watershed.

Northern Mesic Forest

Fassett’s (1944) land cover analysis depicted this forest type as having little importance in the watershed, with less than 1.7% of the land cover. Northern mesic forest stands were predominately located on the north side of Lake Minnesuing and to the east of the Brule River near and on the Copper Range and are known locally as Sugar Camp Hill. Dominant tree species included Acer rubrum, Acer saccharum, Betula alleghaniensis, Betula papyrifera, Ostrya virginiana, Pinus strobus, and Tilia americana (Thomson 1945). The associated ground cover species consisted of species associated with maple–basswood forest, such as Dryopteris carthusiana, Actaea pachypoda, Botrypus virginianus, Sanicula marilandica, and Lysimachia ciliata.

Thomson (1945) noted that the large sugar bush trees (Acer spp.) were gone, as was the sugar camp of earlier days. The topsoil was lost, first by logging and then with subsequent fires resulting in erosion. Most of the landscape of the 1940s was covered with a coppice scrubby growth of Acer spp., and Thomson predicted it would be many years before this forest recovered (Thomson 1945). Surprisingly, Thomson did not include Tsuga canadensis in his description of this forest. However, on May 9, 1944, Thomson did collect a voucher specimen on the bank of a stream crossing the Copper Range (Appendix 1). Later, Davidson et al. (1973) documented a range extension for Tsuga canadensis at one of the stands at the westernmost boundary of the watershed near Lake Minnesuing. Today, Tsuga canadensis is known to extend further west into St. Louis County in Minnesota and several others.

Today, northern mesic forest, typically dominated by Acer saccharum, Betula


alleghaniensis, and Tilia americana, comprises 1.5% of the watershed, covering 750 ha. As in yesteryear, this mesic forest is located along the Copper Range and on the southwestern side of Lake Minnesuing. Sugar Camp Hill on top of the Copper Range was last logged in 1933 and has slowly been regenerating a maple–basswood forest. Today, these stands have moved to a later successional stage and are slated to be harvested in the next decade. In the Lake Minnesuing area, some mesic forest stands have a component of Tsuga canadensis. These are privately and publicly owned, and most are of poor quality. A few are represented by multi-age stands with relatively high species diversity. A patch of Adiantum pedatum was found in a deep gully in such a stand and represents only the third known occurrence of this species in the watershed.

Northern Wet Forest

Thomson (1945) lumped this forest community type together with northern wet-mesic forest community type into what he called conifer bog. This is described in more detail above under northern wet-mesic forest.

Today, northern wet forest encompasses 3.8% (1,896 ha) of the watershed and consists of weakly minerotrophic, conifer-dominated, acid peatlands located north and south of the Lake Nebagamon region where the water table is near the surface or where drainage is somewhat impeded. A large forested complex consisting of a mix of Fraxinus americana, Thuja occidentalis, Larix laricina, and Picea mariana intergrade into a mosaic in this region, with species individually responding to gradients of pH, water depth, the presence of Sphagnum spp., and available nutrients. This forest is uncommon in the watershed, except in the headwaters region and is distinct with a specialized associations of plant species. However, even this forest has increased from zero non-native species to 6.4% between the two studies. An area deserving further protection occurs along the east side of Degerman Road, north of Lake Nebagamon. In this region we observed a northern wet forest dominated by a canopy of Picea mariana with ample regeneration of 3.6–4.6 m tall Thuja occidentalis and uncommon forbs underneath; this is the only area in the watershed known to have significant Thuja occidentalis regeneration.


Overall, our study has shown that the Brule River watershed is of exceptional quality, but also an area at high risk. Many challenges exist for sustaining, maintaining, and restoring its natural forest communities. The boreal forest remains susceptible to severe erosion of clay banks as rain events and snow melts intensify under changing climate scenarios. The northern wet-mesic swamps are even- aged and apparently not regenerating naturally. Northern hardwood swamps are likely to decline substantially in the watershed over the next 50 years from emerald ash borer infestations. Old growth northern dry-mesic forest will not likely regenerate in the absence of fires and are a declining presence in the watershed


forests. The pine barrens mosaic remains diminished and the area continues to be dominated by Pinus resinosa and P. banksiana plantations with only small opportunities to expand.

The purpose of this study was to document the existing flora, make comparisons with earlier surveys, and make qualitative and quantitative data available for future researchers. We highly recommend that a similar survey be made one or two decades after this study to document changes and inform land managers and decision-makers with new information for making the difficult decisions they will face in the future.


The following organizations, agencies and individuals assisted in financially supporting the project, and we are forever grateful.

Grants: University of Wisconsin Foundation–Dr. Donald Davidson Memorial Fund; Enbridge EcoFootprint Grant; Wisconsin Coastal Management Grant Program (grant no. NA15NOS4190094).

Organizations: Brule River Coalition, Brule River Preservation, Brule River Sportsman’s Club, Friends of the Brule River, Winneboujou Club, The Nature Conservancy, Cedar Island Conservancy, Douglas County Land and Water Conservation Department, West Wisconsin Land Trust, Wisconsin Department of Natural Resources (Brule River State Forest, State Natural Areas Program, Bureau of Natural Heritage Conservation).

We extend our deepest appreciation to Dr. Michael Huft, editor of the Great Lakes Botanist and the anonymous reviewer for adding insight, extensive notes and suggestion for both conciseness and clarification for this article and its companion article, “Analyzing Vegetative Cover of the Bois Brule River Watershed Re-Visited in Northwestern Wisconsin, Part I: Forest Stand Changes (1968 to 2016).â€

We honor the memories of Dr. John W. Thomson, Jr., and Dr. Donald W. Davidson and hope we’ve done justice in interpreting their data and work, as without this data, none of this work would have been possible. We were honored and grateful to have the expertise of Dr. Loy Richard Phillippe of the Illinois Natural History Survey, whose more than 44,000 collections in his extraordinary career provided specimen details we often didn’t know or see. We also are deeply grateful to Dr. Emmet Judziewicz, professor emeritus, University of Wisconsin-Stevens Point. Emmet was always our lead botanist finding the unusual or a niche upon a niche finding that obscure plant. I still don’t know how he sighted Carex ×knieskernii among the Carex arctata and Carex castanea in an upland white cedar swamp. Thank you, Emmet!

Thank you to Dr. Matt Tenyck for his comments on earlier drafts of this article and his unending support for numerous requests. We are thankful to Dr. Brenda Molano-Flores and Ms. Jean Mengelkoch for their assistance with preparing plant specimens for the project. We also want to thank UW-Superior alumni and students who worked on this project—Stephanie Glass, Research Tech 1, and UW-Superior natural science students Paige Kent, Mike Krick, Adam Krick, Daniel Gil De La Madrid, and Molly Bergman for working a lot of extra hours to ensure the quality of the reported data and meeting every unreasonable demand we asked of them and then some. You have all been so awesome, generous, and kind, and it has been a privilege and an honor to learn from you, guide you, and share with you the diversity of one of the most pristine landscapes in Wisconsin. Let our paths cross again in the wild places on the Brule.


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Page  59 APPENDIX 1. Vascular plant taxa documented for the Brule River Watershed. Nomenclature follows Voss and Reznicek (2012) for species known to occur in Michigan. Nomenclature for other species follows the published volumes of the Flora of North America (Flora of North America Editorial Committee 1993+) and Judziewicz et al. (2014) was used for grasses. The list is organized by major plant groups, then alphabetically by family and species within each major group. Non-native species are indicated by an asterisk (*) in front of the name.

For taxa collected during the current project, the collection number is prefaced by the initial of the last name of the primary collector: A = Anderson; F = Feist; H = Hlina; M = Marcum; P = Phillipe; S = Schwarting. Earlier collections are indicated by an italicized collection number and by the full name of the collector with the exception of Thomson, whose collections are denoted with a T. Links to the voucher specimens can be found at Consorium of Midwest Herbaria (2020).

The Habitat column indicates the most common communities in which a particular species was observed, using the following abbreviations: BF = Boreal Forest; WC = White Cedar; PB = Pine Barrens; NDF = Northern Dry Forest; NDMF = Northern Dry-Mesic Forest; NHS = Northern Hardwood Swamp; NMF = Northern Mesic Forest; BSS = Black Spruce/Tamarack Forest; AP = Aquatic Plants (Brule River, Lake Minnesuing, Lake Nebagamon); MB = Mud Bank Plants; W-NF = Wetlands -Non-forested; LSS = Lake Superior Shoreline; D = Disturbed Site.

The Status column indicates whether a taxon is on Wisconsin’s rare species list (END = Endangered; THR = Threatened; SC = Special Concern), is a new record for the Brule River watershed (indicated by WS), or is a new county record (indicated by CR).

Taxon Habitat Status


ATHYRIACEAE (LADYFERN FAMILY) Athyrium filix-femina (L.) Roth (common lady fern); F6037, F6074, F6411, F6489, M6844, M6882, F6489, BF, NDMF T5159, T5525

CYSTOPTERIDACEAE (Brittle Fern Family)

Cystopteris fragilis (L.) Bernh. (brittle bladder fern); M7134, M7135, F5654 BG WS Cystopteris tenuis (Michx.) Desv. (MacKay's brittle fern); Sommerville 95 NMF Gymnocarpium dryopteris (L.) Newm. (common oak fern); F6008, F6093, M6863, T5503 BF, NMF

DENNSTAEDTIACEAE (Bracken Fern Family) Pteridium aquilinum (L.) Kuhn (bracken fern); F6409, S109, T5088, T5100 PB, NDMF

DRYOPTERIDACEAE (Wood Fern Family) Dryopteris carthusiana (Vill.) H. P. Fuchs (spinulose wood fern); F6002, F6005, F6092, F6360, M6974, BF, NMF

M7056, M7138, M6845, M6862, Ts.n. Dryopteris cristata (L.) A. Gray (crested shield fern); F6360, M6838, M6860, T5507 WC, NHS Dryopteris fragrans (L.) Schott (fragrant fern); F5658 BG Dryopteris intermedia (Willd.) A. Gray (evergreen wood fern); F6336, F5624, P43857 BF, NDM

(Continued on next page)


Page  60 APPENDIX 1. (Continued)

Taxon Habitat Status

EQUISETACEAE (Horsetail Family) Equisetum arvense L. (common horsetail); H3568 BF, NMF Equisetum fluviatile L. (river horsetail); F6579, H3635, H3789, M7295 BSS. AP Equisetum hyemale L. (scouring rush); H3985, H3565 NHS, NMF Equisetum laevigatum A. Braun (smooth horsetail); H3706 NMF WS Equisetum scirpoides Michx. (dwarf scouring rush); F5618, F5644, F6139, M6721 WC Equisetum sylvaticum L. (woodland horsetail); F5634, M6720, P43787 BF, NMF Equisetum ×ferrissii Clute (woodland horsetail); T5549 NDF

ISOETACEAE (Quillwort Family) Isoetes echinospora Durieu (spiny spored quillwort); S233 AP

LYCOPODIACEAE (Club-Moss Family)

Dendrolycopodium dendroideum (Michx.) A. Haines (tree club moss); A2630, F6101, F6331, M6649, P43848,

P44191 BF, NDMF

Dendrolycopodium hickeyi (W.H. Wagner, Beitel & R.C. Moran) A. Haines (Hickey's tree club moss); H3874, WC, NDMF WS

H3979, M6849

Dendrolycopodium obscurum (L.) A. Haines (ground pine); Conklin 404, Salomaki 97/36 WC

Diphasiastrum complanatum (L.) Holub (northern ground-cedar); M6766 NDF, NDMF WS

Diphasiastrum digitatum (Dill. ex A.Braun) Holub (trailing ground-pine); A2628, H3513, M7248, S113, S287 NDMF

Diphasiastrum tristachyum (Dill.) Holub (northern ground-pine); M7211 PB

Huperzia lucidula (Michx.) R.Trevis. (shining club moss); H3656, F5610, Ts.n. WC, NHS

Huperzia selago (L.) Bernhardi (fir club moss); Clark 1065 WC

Lycopodiella inundata (L.) Holub (bog club moss); T5360 W-NF

Lycopodium clavatum L. (running ground pine); F6332, H3518, M6650, M7212, P43866 NDF, NDMF

Spinulum annotinum (L.)A. Haines (stiff clubmoss); F5615, F6100, F6328, M6651, M7114, M6866, P44105,

T5341, T5495 WC, NDMF

ONOCLEACEAE (Sensitive Fern Family)

Matteuccia struthiopteris (L.) Todaro (ostrich fern); P43774, F6513, F6570, Ts.n. NHS, NMF

Onoclea sensibilis L. (sensitive fern); F6118, F6375, F6496, M6969, P43775, P44156, Ts.n. BF, NHS

OPHIOGLOSSACEAE (Adder's-Tongue Family)

Botrychium lanceolatum (S.G.Gmel.) Angstr. (lace-leaved moonwort); T5577 NMF

Botrypus virginianus (L.) Michx. (rattlesnake fern); S127, T5391, T5559 WC

Sceptridium multifidum (S. G. Gmel.) M. Nishida (leathery grape fern); T5400 PB



OSMUNDACEAE (Royal Fern Family) Osmunda claytoniana L. (interrupted fern); S120 BF, NMF Osmunda regalis L. (royal fern); S191 WC, NHS Osmundastrum cinnamomeum (L.) C. Presl (cinnamon fern); F6319, S110 WC, BSS

POLYPODIACEAE (Polypody Fern Family) Polypodium virginianum L. (common polypody fern); F5645.1, M7119, T5311 BG

PTERIDACEAE (Maidenhair Fern Family) Adiantum pedatum L. (maidenhair fern); S142 NMF

SELAGINELLACEAE (Spikemoss Family) Selaginella rupestris (L.) Spring (rock spikemoss); F5651, T5142, T5200 PB

THELYPTERIDACEAE (Marsh Fern Family) Phegopteris connectilis (L.) Slosson (northern beech fern); F6011, F6073 H3607, M6874, T5504 BF, NMF Thelypteris palustris Schott (marsh fern); S186, T5528 WC, NHS

WOODSIACEAE (Woodsia Family) Woodsia ilvensis (L.) R. Br. (rusty cliff fern); F5646, F5650.1, T5335 BG


CUPRESSACEAE (Cypress Family) Thuja occidentalis L. (white cedar); M7288, P44145, Ts.n. WC, BF

PINACEAE (Pine Family) Abies balsamea (L.) Mill. (balsam fir); S34, S45, H4288, T499, T5500 BF,WC,NDFM Larix laricina (Du Roi) K. Koch (tamarack); M7030, M7163, M7306, F6428, F6376, P44097, T s.n. BSS, WC

*Picea abies (L.) H. Karst (Norway spruce); S86 BF Picea glauca (Moench) Voss (white spruce); S38, S46 BF, NDMF WS Picea mariana (Mill.) Britton, Sterns & Poggenb. (black spruce); F6404, M7262, M7309, T5497 BSS, WC Pinus banksiana Lamb. (jack pine); M6691, M6810, P44006, T5223, T5698 PB, NDMF Pinus resinosa Aiton (red pine); F5652 PB, NDMF Pinus strobus L. (white pine); P43779 NDMF, WC

*Pinus sylvestris L. (Scotch pine); S289 BF Tsuga canadensis (L.) Carrière (eastern hemlock); F6098, S119, T5501 NMF WS

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Page  62 APPENDIX 1. (Continued)

Taxon Habitat Status

TAXACEAE (Yew Family)

Taxus canadensis Marshall (American yew); H3418, P43784, Ts.n. BF, WC


ADOXACEAE (Moschatel Family)

Sambucus canadensis L. (American elderberry); H2397 WC, BSS WS Sambucus racemosa L. (red-berried elder); F5625, F6060, S69 BF, NMF Viburnum lentago L. (nanny berry); F6571, H3951, P43884, S133, T5518 BF, WC Viburnum rafinesquianum Schult. (arrow-wood); P43783, P44212, F6348, F6600 BF, NHS WS Viburnum trilobum Marshall (American high-bush cranberry); H3954, P44153, F6595, T5477 BF, WC

AMARANTHACEAE (Amaranth Family)

Chenopodium album L. (lamb's quarters); S278 PB Chenopodium simplex (Torr.) Raf. (maple leaf goosefoot); F6125 BF WS *Froelichia gracilis (Hook.) Moq. (cottonweed); Judziewicz 12125, Sumlan 737 PB


Rhus typhina L. (staghorn sumac); H4108, T5257 BF, NDF Rhus x pulvinata Greene (hybrid sumac); T5500 NMF Toxicodendron rydbergii (Rydb.) Greene (western poison-ivy); S129 NDF, NDMF WS

APIACEAE (Parsley Family)

Angelica atropurpurea L. (purple-stemmed angelica); S265 W-NF Cicuta bulbifera L. (bulblet water hemlock); H3910, M7242, M7281, F6523, T5091 W-NF, NHS Cicuta maculata L. (water hemlock); F6089, M6864, P44146 BF, NHS Heracleum maximum Bartram (cow parsnip); H4072, M6730, T5540 W-NF Osmorhiza claytonii (Michx.) C. B. Clarke (hairy sweet cicely); F6335, S128, T5538 NMF *Pastinaca sativa L. (wild parsnip); T5255 D Sanicula marilandica L. (black snakeroot); F6079, M6765, M6808, M7003, T5482 BF, NMF Sanicula odorata (Raf.) Pryer & Phillippe (clustered black snakeroot); Christensen s.n. BF Sium suave Walter (water parsnip); H3624, P44070, T5320 W-NF, NHS Zizia aurea (L.) W. D. J. Koch (common golden alexanders); P43867 BF WS



APOCYNACEAE (Dogbane & Milkweed Family)

Apocynum androsaemifolium L. (spreading dogbane); H4046, S205 PB, NDF Apocynum cannabinum L. var. hypericifolium A. Gray (clasping dogbane); S181 BF WS Asclepias exaltata L. (poke milkweed); H3511, H3616, M7216 NDMF, PB WS Asclepias incarnata L. (swamp milkweed); M7247, T5290, T5547 W-NF, NHS Asclepias ovalifolia Decne (dwarf milkweed); H3386, H3387 PB THR, WS Asclepias syriaca L. (common milkweed); H4044, S254 BF


Ilex mucronata (L.) M. Powell, V. Savolainen & S. Andrews (mountain holly); A2441, F6432, F6458, M6662, WC, BSS M6827, M7162, P44133, T5482 Ilex verticillata (L.) A. Gray (winterberry); F6061, F6146, F6432, M6823, M7181, M7289, P43868, P44160, BF, NMF, NHS P44185, P44208, T5096

ARALIACEAE (Ginseng Family)

Aralia hispida Vent. (bristly sarsaparilla); T5256 NMF Aralia nudicaulis L. (wild sarsaparilla); F6308, F6407, P43746, T5479 BF, NDMF, NMF Aralia racemosa L. (American spikenard); F6407, M7126, P43860, T5099, T5545 BF, NDMF, NMF Hydrocotyle americana L. (marsh pennywort); H3556, M6842 WC Panax trifolius L. (dwarf ginseng); H3566, H3816, S194, T5418 NMF WS


Asarum canadense L. (Canadian wild-ginger); F5631, H3562, H3857, S103 WC, NDMF

ASTERACEAE (Sunflower Family)

Achillea millefolium L. (common yarrow); H3487, S170 PB, NDF

*Achillea ptarmica L. (sneezewort); Gerst s.n. D Ambrosia artemisiifolia L. (ragweed); H4061 PB, NCF WS Ambrosia psilostachya DC. (western ragweed); T s.n. NDMF Anaphalis margaritacea (L.) Benth. (pearly everlasting); H3962, M7231 PB, NDF Antennaria howellii Greene (small pussy's toes); F5657, M6696, P44035, P44043, T5328, T5329, T5372, T5375 PB, NDF Antennaria neglecta Greene (cat's foot); H3582, S95 BF Antennaria parlinii Fernald (smooth pussy's toes); M6693, P43760, P44022, T5205, T5374 PB, NDMF

*Arctium minus Bernh. (common burdock); S270 BF WS *Artemisia pontica L. (Roman wormwood); T5270 PB *Artemisia vulgaris L. (mugwort); T5116 NDMF

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Page  64 APPENDIX 1. (Continued)

Taxon Habitat Status

Bidens beckii Spreng. (water beggar-ticks); F6515, T5367 AP Bidens cernua L. (nodding beggar's tick); F6553, M7278, T5359 W-NF Bidens connata Willd. (purple-stemmed tickseed); H3791 W-NF-NHS Bidens discoidea (Torr. & A. Gray) Britton (swamp beggar's tick); H3938 WC, NHS CR Bidens frondosa L. (common beggar's tick); F6138, F6555, M6848, M7240, P44139, Ts.n. W-NF, NHS Bidens tripartita L. (straw-stem beggar-ticks); H3534 W-NF Bidens vulgata Greene (tall beggar-ticks); H3484 W-NF, NHS WS

*Centaurea jacea L. (brown knapweed); T5076 PB, NDF *Centaurea stoebe L. (spotted knapweed); H4088, T5190 PB, NDF *Cirsium arvense (L.) Scop. (Canada thistle); H3834, M7228 NDF, BF

Cirsium discolor (Willd) Spreng. (field thistle); Mitchell 87 PB Cirsium muticum Michx. (swamp thistle); F6494, H3585, H3690, Ts.n. NHS, BF *Cirsium vulgare (Savi) Ten. (bull thistle); S264 PB, NDMF Conyza canadensis (L.) Cronq. (horseweed); M6778, P43898, T5544 PB

*Crepis tectorum L. (hawk's beard); T5268 PB Doellingeria umbellata (Mill.) Nees (flat-topped aster); F6080, M6832, M6885, P43869, P44136, P44178, BF, NMF

T5139, T5566 Erechtites hieraciifolius (L.) DC. (burnweed); H3987 PB Erigeron annuus (L.) Pers. (annual fleabane); S193, S207 BF WS Erigeron glabellus Nutt. (streamside fleabane); S283, T5548 PB Erigeron strigosus Willd. (daisy fleabane); M6746, M6790, T5182, T5387 PB, NDMF Eupatorium perfoliatum L. (boneset); P44213, S274 W-NDF Eurybia macrophylla (L.) Cass. (big-leaved aster); F6072, P44167, T5517 BF, NDF Euthamia graminifolia (L.) Nutt. (grass-leaved goldenrod); H3725, P43906, T s.n. PB, NDMF Eutrochium maculatum (L.) E. E. Lamont (spotted Joe-pye-weed); M6858, P44158, T5298 W-NF *Gnaphalium uliginosum L. (cud weed); H3963, T5195 NDMF *Grindelia squarrosa (Pursh) Dunal (gumweed); T5434 NMF Helianthus giganteus L. (giant sunflower); P43881, M7263, F6598, T5449 BF, NDMF Helianthus hirsutus Raf. (hairy sunflower); H3486 BF CR Helianthus occidentalis Riddell (western sunflower); M6745, M6785, T5130 PB Helianthus strumosus L. (pale-leaved sunflower); H3738, M6792 PB

*Hieracium aurantiacum L. (devil's paintbrush); F6361, M6788, M6999, P43897 PB, NDF, NDMF WS *Hieracium caespitosum Dumort. (yellow hawkweed); M6743, M7000, P44004 PB, NDF



*Hieracium lachenalii Suter (common hawkweed); F6029, F6097, P43865 BF, NDMR CR

*Hieracium piloselloides Vill. (glaucous king-devil); H4095, S135 NHS, NMF WS Hieracium scabrum Michx. (rough hawkweed); P43891, P43907, M7111, S29, T5483 BF, PB Hieracium umbellatum L. (Canada hawkweed); F6076, M6793 PB, NMF WS Krigia biflora (Walter) S. F. Blake (false-dandelion); H3842, P44013 PB CR Lactuca biennis (Moench) Fernald (tall blue lettuce); S177 PB, NHS WS Lactuca canadensis L. (Canada lettuce); P43854, T5186, T5187 PB, NMF Lactuca hirsuta Nutt. (hairy tall lettuce); M6749 PB WS

*Leucanthemum vulgare Lam. (ox-eye daisy); S134, T5543 PB, NDF Liatris aspera Michx. (rough blazing star); M6781, M7226, P43904 PB Liatris ligulistylis (A.Nelson) K.Schum. (meadow blazing star); T5071 PB

*Matricaria discoidea DC (pineapple weed); S2111 D Packera aurea (L.) Ã. Löve & D. Löve (golden ragwort); M6970, M6977, M7043, T5559 NDF, NHS Packera paupercula (Michx.) Ã. Löve & D. Löve (northern ragwort); F6353, P44018, P44050, S173, PB, NHS

T5332, T5515 Petasites frigidus (L.) Fries (sweet colt's-foot); H3555 BF, NDMF Petasites sagittatus (Pursh) A. Gray (arrowhead sweet colt's foot); Photo BF THR Prenanthes alba L. (white lettuce); F6086, P44169, S21, T5441 BF, NMF Pseudognaphalium obtusifolium (L.) Hilliard & B. L. Burtt (cat's-foot; fragrant cudweed); F6087, PB, NDMF CR

M6794, P43919 Rudbeckia hirta L. (black-eyed Susan); H4063, T5388 NDF Rudbeckia laciniata L. (cut-leaved coneflower); T5523, T5524 BF Solidago canadensis L. (Canada goldenrod); S280, T5516 BF, NMF Solidago flexicaulis L. (zig-zag goldenrod); F6123 BF, NMF Solidago gigantea Aiton (giant goldenrod); F6077, F6112, F6568, M7227, P43871, P44144, P44175, T5379 BF, NMF Solidago hispida Willd. (hairy goldenrod); P43910, T5125 PB, NDF Solidago juncea Aiton (early goldenrod); H3388, P43895, T5122 PB Solidago nemoralis Aiton (gray goldenrod); H3527, M6776, M7223, P43894, T5121, T5123 PB Solidago ptarmicoides (Torr. & A. Gray) B. Boivin (upland white goldenrod); H4094, T5133, T5380 PB Solidago speciosa Nutt. (showy goldenrod); M6796, M7224 PB Solidago uliginosa Nutt. (bog goldenrod); M6877, P44149, T5120, T5340 WC, NHS, BSS

*Sonchus arvensis L. (field sow-thistle); F6126, F6584 BF *Sonchus oleraceus L. (common sow-thistle); H4117 D WS Symphyotrichum boreale (Torr. & A. Gray) Ã. Löve & D. Löve (northern bog aster); T5124, T5140 WC

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Page  66 APPENDIX 1. (Continued)

Taxon Habitat Status

Symphyotrichum ciliolatum (Lindl.) Ã. Löve & D. Löve (northern heart-leaved aster); M7221, M7225, BF, PB, NDF

P43863, T5136Symphyotrichum laeve (L.) Ã. Löve & D. Löve (smooth aster); M6740, P43912, T5185, T5132 PB Symphyotrichum lanceolatum (Willd.) G. L. Nesom var. lanceolatum (panicled aster); F6111, F6585, W-NF, NHS

M6789, S25, T5138, T5368 Symphyotrichum lateriflorum (L.) Ã. Löve & D. Löve (calico aster); F6075, P43870, P44150, P44182, S22 BF, NMF Symphyotrichum ontarionis G.L. Nesom (Ontario aster); H4020 BF WS Symphyotrichum oolentangiense (Riddell) G. L. Nesom (sky blue aster); M6777, P43889, T5567 PB Symphyotrichum pilosum (Willd.) G. L. Nesom (frost aster); T5135 PB Symphyotrichum puniceum (L.) Ã. Löve & D. Löve (swamp aster); M6822, M6868, M7239, P44138, NHS, W-NF

T5485, T5565

Symphyotrichum urophyllum (DC.) G. L. Nesom (arrow-leaved aster); H4170, H4615 PB CR *Tanacetum vulgare L. (common tansy); F6583, H4106 BF, NDF WS *Taraxacum officinale F. H. Wigg (common dandelion); F5636 BF, NMF WS

Tephroseris palustris (L.) Rchb. (marsh groundsel); Cheney 7419 W-NF *Tragopogon dubius Scop. (lesser goat's beard); P44018, P44050, P44016, S277, T5252 PB WS

BALSAMINACEAE (Touch-Me-Not Family) Impatiens capensis Meerb. (orange jewel-weed); F6506, F6574, M7315, Ts.n. NHS, WC

BERBERIDACEAE (Barberry Family) Caulophyllum thalictroides (L.) Michx. (blue cohosh); S72, T5568 BF

BETULACEAE (Birch Family) Alnus incana (L.) Moench (speckled alder); F5642, F6071, F6511, F6533, M6836, M7253, P44098, BF, NHS, W-NF

P44211, T5052Alnus viridis (Chaix) DC. (green alder); Ts.n. NDMF Betula alleghaniensis Britton (yellow birch); H3973, S122, T5492, T5540 NMF Betula papyrifera Marshall (paper birch); M6677 BF, NDF Betula pumila L. (bog birch); F6378, S87 BSS Carpinus caroliniana Walter (American hornbeam); H3439 NFM, NDMF Corylus americana Walter (American hazelnut); P43918, M7144, M7213, P44009, P44038, F6354, PB, NDMF

P44186, T5382, T5545 Corylus cornuta Marshall (beaked hazelnut); F6410, F6342, M7172, P44187, Ts.n. BF, NMF Ostrya virginiana (Mill.) K. Koch (ironwood); M7011, M7140, P44189, T5537 BF, NMF



BORAGINACEAE (Borage Family)

Cynoglossum boreale Fernald (northern wild comfrey); F6049, H4050 BF, NDMF Lithospermum canescens (Michx.) Lehm. (hoary puccoon); M6687, M6798, P44039, S96, T5227, T5371, T5383 PB, NDF Lithospermum caroliniense (J. F. Gmel.) MacMill. (hairy puccoon); T5377 PB

*Lithospermum officinale L. (gromwell); Epstein s.n. PB *Myosotis arvensis (L.) Hill (field forget-me-not); P43762 BF CR *Myosotis scorpioides L. (forget-me-not); F6142, M6965, M7264, Ts.n. BF, AP, WC *Myosotis sylvatica Hoffm. (garden forget-me-not); H3817 BF, NMF WS

BRASSICACEAE (Mustard Family)

Arabidopsis lyrata (L.) O'Kane & Al-Shehbaz (sand cress); H3493, T5560 PB WS *Barbarea vulgaris W. T. Aiton (yellow rocket); S58, S101 BF, NHS *Berteroa incana (L.) DC. (hoary alyssum); S286, T5563 NDMF

Cardamine concatenata (Michx.) O.Schwarz (toothwort); H3895, S73 BF WS Cardamine pensylvanica Willd (Pensylvania bitter-cress); H3869, M6964 BF, NHS WS Cardamine pratensis L. var. palustris Wimm. & Grab (cuckoo flower); H3896 BF CR

*Erysimum cheiranthoides L. (wormseed mustard); Gerst s.n. D *Lepidium densiflorum Schrad. (small pepper grass); H3380, S281, T5201 D *Nasturtium officinale W. T. Aiton (water cress); H3658, M7271, T5272 AP, NHS, WC

Rorippa palustris (L.) Besser (yellow cress); T5179 D *Sisymbrium altissimum L. (tumble mustard); Gerst s.n. D *Thlaspi arvense L. (penny-cress); Gerst s.n. D

Turritis glabra L. (tower mustard); H3376, H3672, M6820, S210 PB

CABOMBACEAE (Water-shield Family)

Brasenia schreberi J.F. Gmel. (water-shield); H3642 AP WS

CAMPANULACEAE (Bell Flower Family)

Campanula aparinoides Pursh (marsh bellflower); F6554, T5085 NHS, W-NF

*Campanula rapunculoides L. (creeping bellflower); H3704, S275 NDF, NDMF WS Campanula rotundifolia L. (harebell); M6738, M7155, P44055, T5384 PB, NDF, NDMF Lobelia inflata L. (Indian tobacco); H3856, M7161 PB, NDF, NDMF Lobelia spicata Lam. (spiked lobelia); S290 D WS


Humulus lupulus L. (common hops); H3861, F6578, T5355 BF

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Page  68 APPENDIX 1. (Continued)

Taxon Habitat Status

CAPRIFOLIACEAE (Honeysuckle Family)

Lonicera canadensis Marshall (American fly honeysuckle); F5616, P43759, P43773, M6663, M6963, BF, WC

F6309, M7053, T5390 Lonicera dioica L. (mountain honeysuckle); H3825 NDF, NDMF Lonicera hirsuta Eaton (hairy honeysuckle); H3368, F6144, F6359, M6888 PB, NDF, NDMF WS

*Lonicera morrowii A. Gray (Asian fly honeysuckle); H3968 BF, NHS WS Lonicera oblongifolia (Goldie) Hook. (swamp fly honeysuckle); S214, T5239 BF, WC, NDMF *Lonicera tatarica L. (Tartarian honeysuckle); F6059 BF, NHS WS Lonicera villosa (Michx.) Schultes (mountain fly honeysuckle); F6025, M6668, S31, S60, T5237 BF, NHS

*Lonicera ×bella Zabel (hybrid honeysuckle); S132 BF, NMF WS Symphoricarpos albus (L.) S. F. Blake (snowberry); H3865, M6819, M7217, T5181 PB, NDF, NDMF Symphoricarpos occidentalis Hook. (wolfberry); T5480 NDMF Triosteum aurantiacum E. P. Bicknell (early horse-gentian); H3521, P43873 BF CR


*Arenaria serpyllifolia L. (thyme-leaved sandwort); S97 BF CR *Cerastium fontanum Baumg. (mouse-ear chickweed); H3897, S112, T5198, T5324 NDF, NDMF *Dianthus barbatus L. (sweet-William); H3666, H3667 NMF CR *Gypsophila muralis L. (cushion baby's breath); M6855 D CR Moehringia lateriflora (L.) Fenzl (wood sandwort); P43772 BF CR *Scleranthus annuus L. (knawel); H3908, M6756, S76, T5094 PB, NDF Silene antirrhina L. (sleepy catchfly); M6758 PB WS *Silene dichotoma Ehrh. (forked catchfly); Fassett 9226 PB *Silene latifolia Poir. (white campion); S160, T5258 PB, NDF, NMDF *Silene vulgaris (Moench) Garcke (bladder campion); H3728, S201, T5392 PB, NDMF *Spergularia rubra (L.) J.Presl & C.Presl (red sand spurry); P44057, S152 NDF CR Stellaria borealis Bigelow (northern stitchwort); Davis s.n., Koch 5800 WC *Stellaria graminea L. (starwort); Gerst s.n. D Stellaria longifolia Willd. (long-leaved chickweed); M6709, M7034, M6980, M7062, S176, T5260 WC, NHS

CELASTRACEAE (Bittersweet Family)

Celastrus scandens L. (American bittersweet); F6596, T5354 BF




Ceratophyllum demersum L. (coon's tail); F6559, H3761, H3577, T5192, T5272 AP

CISTACEAE (Rock-Rose Family)

Crocanthemum bicknellii (Fernald) Janch. (Bicknell's rock-rose); H3934, M6732 PB WS Crocanthemum canadense (L.) Britton (common frostweed); P43902, P43902, T5393 PB Lechea intermedia Britton (intermediate pinweed); M6803, M6816, M7156, P43921 PB, NDF WS

CONVOLVULACEAE (Morning Glory Family)

Calystegia spithamaea (L.) Pursh (low bindweed); M6751, P44011, S144, T5333, T5514 PB, NDF, NDNF

CORNACEAE (Dogwood Family)

Cornus alternifolia L. f. (pagoda dogwood); P44188, T5522 BF, WC, NMF Cornus canadensis L. (bunchberry); F6137, F6314, F6434, M6869, M7005, M7027, M7174, P43761, P44194, BF, WC, NMF

T5138Cornus foemina Mill. subsp. racemosa (Lam.) J. S. Wilson (gray dogwood); H4045, S208, T5253 BF, NDF Cornus rugosa Lam. (round-leaved dogwood); F6078, F6510, F6593, H3975, H3709, T5404 BF, NDMF Cornus sericea L. (red-osier dogwood); F6048, F6116, F6484, F6566, M6989, M7287, T5081, T5563 BF, NHS, BSS


Echinocystis lobata (Michx.) Torr. & A. Gray (wild cucumber); H4053 BF, NHS

DIERVILLACEAE (Bush-honeysuckle Family)

Diervilla lonicera Mill. (bush honeysuckle); F6134, F6143, F6345, M7121, P43908, T5403 PB, NDF

DROSERACEAE (Sundew Family)

Drosera intermedia Hayne (spoon-leaved sundew); M7193, P44069, P44126 W-NF CR Drosera rotundifolia L. (round-leaved sundew); F6403, M6901, F6460, T5314, T5551 WC, BSS, W-NF

ELAEAGNACEAE (Oleaster Family)

Shepherdia canadensis (L.) Nutt. (soapberry); F6044, H3373, H3554, Ts.n. BF

ERICACEAE (Heath Family)

Andromeda glaucophylla Link (bog rosemary); A2609, F6437, F6442, P44094, P44122, T5550 BSS, W-NF Arctostaphylos uva-ursi (L.) Spreng. (bearberry); P44026, T5097 PB, NDF Chamaedaphne calyculata (L.) Moench (leatherleaf); F6374, F6433, F6449, M7022, M7182, M7251, P44080, WC


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Page  70 APPENDIX 1. (Continued)

Taxon Habitat Status

Chimaphila umbellata (L.) W. P. C. Barton (pipsissewa); M6764, M7150, M7189, P44045, P44190 BF,NDF,NDMF Epigaea repens L. (trailing arbutus); H3510, P44041, S59, T5106, T5306 PB, NDF Gaultheria hispidula (L.) Bigelow (creeping snowberry); F5633, F6013, F6422, M7025 WC, BSS Gaultheria procumbens L. (wintergreen); F6102, M6679, M7024, M7151, M7169, P44028, P44192 BF, WC, PB Hypopitys monotropa Crantz. (pinesap); A2612, M7152 WC, NMF Kalmia polifolia Wangenh. (bog-laurel); M7021, M7185, M7255, F6377, P44090, P44121, T5313, T5549 WC, BSS, W-NF Moneses uniflora (L.) A. Gray (one-flowered pyrola); F5609, F6006, F6018, F6321, H3829, P43864, BF, WC, NDF

T5104, T5241 Monotropa uniflora L. (Indian-pipe); F6007, F6416, M7130, P44172, P44193 WC, NDF, BSS Orthilia secunda (L.) House (one-sided pyrola); F5617, F6004, M6667, M6676, M6763, M6829.1, M6870, WC, NDMF

P44168, H3830, T5389 Pyrola americana Sweet (American wintergreen); H3525, M6762, M7145, M7148, P43874, P44171 BF, NDMF CR Pyrola asarifolia Michx. (pink shinleaf); F6131, H3801, H3845, T5511, T5561 BF, NMF Pyrola chlorantha Sw. (green shinleaf); S32, S78, T5245 BF, WC Pyrola elliptica Nutt. (large-leaved shinleaf); F6032, F6090, F6127, M6834, M7128, P44042, P44164 BF, NDMF Pyrola minor L. (snowline wintergreen); M6829.2 WC END, WS Rhododendron groenlandicum (Oeder) Kron & Judd (labrador tea); F6390, M6894, M7017, M7165, F6367, BSS, NHS

M7256, F6429, P44071 Vaccinium angustifolium Aiton (early low blueberry); F6402, F6421, M6660, M6680, M7020, P43767, BF, WC, NDMF

P44002, P44127, T5183, T5184, T5336 Vaccinium macrocarpon Aiton (large cranberry); F6438, F6472, P44118, P44130 WC WS Vaccinium myrtilloides Michx. (velvet-leaf blueberry); F6431, M6661, M7019, P43788, P44008 BSS, WC WS Vaccinium oxycoccos L. (small cranberry); F6396, F6438, F6472, M7026, M7178, M7254, P44065, T5552 WC, BSS Vaccinium vitis-idaea L. (lingonberry); Photo WC END


Euphorbia glyptosperma Engelm. (rib-seed sand mat); S260 PB Euphorbia maculata L. (spotted sand-mat); S259 PB

FABACEAE (Pea and Bean Family)

Amphicarpaea bracteata (L.) Fernald (American hog peanut); F6085, F6586, T5343 NDMF, NMF Astragalus canadensis L. (Canadian milkvetch); F6051 BF WS Hylodesmum glutinosum (Willd.) H. Ohasi & R.R. Mill (pointed tick-trefoil); H3883 BF CR Lathyrus japonicus Willd. (beach pea); F6597 LSS



Lathyrus ochroleucus Hook. (cream pea); F5666, F6364, P44029, T5325 BF, NDF, NMF Lathyrus venosus Willd. (veiny pea); S255, T5509, T5518 BF Lespedeza capitata Michx. (round headed bush clover); T5348 PB *Lotus corniculatus L. (bird's foot trefoil); S150 BF, NDMF WS *Lupinus polyphyllus Lindl. (garden lupine); S130 NMF *Medicago lupulina L. (black medic); H4065 BF WS *Melilotus albus Medik. (white sweet-clover); S203 BF, NHS WS *Melilotus officinalis (L.) Pall (yellow sweet-clover); S151 D WS *Robinia pseudoacacia L. (black locust); T5169 NDMF *Securigera varia (L.) Lassen (crown-vetch); H4107 BF WS *Trifolium arvense L. (rabbit-foot clover); S158, S268, S284 BF WS *Trifolium aureum Pollich (hop clover); M7157, S197, T5386 BF, NDF

*Trifolium campestre Schreb. (low hop clover); Gerst s.n. NDF *Trifolium hybridum L. (alsike clover); S269, T5520 NDF *Trifolium pratense L. (red clover); Christensen s.n. BF, NDF *Trifolium repens L. (white clover); S192 BF, NDF

Vicia americana Willd. (American vetch); M7002, M7157, S148, T5576 BF, NMF *Vicia cracca L. (cow vetch); M7153 NDF CR *Vicia sativa L. (common vetch); H4064 D *Vicia villosa Roth (hairy vetch); T5524 D

FAGACEAE (Beech Family)

Quercus ellipsoidalis E. J. Hill (northern pin oak); P43855, T5102, T5189 PB, NDMF Quercus macrocarpa Michx. (bur oak); T5218, T5264, T5349, T5542, T5578 PB, NDMF Quercus rubra L. (red oak); P44214, T5515 PB, NDMF

GENTIANACEAE (Gentian Family)

Gentiana alba Nutt. (pale gentian); H4055 W-NF CR Gentiana andrewsii Griseb. (bottle gentian); H1924 W-NF Gentiana rubricaulis Schwein. (red-stemmed gentian); T5509 W-NF Halenia deflexa (Sm.) Griseb. (spurred gentian); M6711, M6853, M6887, P44176 BF, NDMF

GERANIACEAE (Geranium Family)

Geranium bicknellii Britton (Bicknell's geranium); H3796, M7133, T5161 PB, BG

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Page  72 APPENDIX 1. (Continued)

Taxon Habitat Status

GROSSULARIACEAE (Gooseberry Family)

Ribes americanum Mill. (wild black current); F5635, S54, T5334, T5521 BF, NMF Ribes cynosbati L. (prickly wild gooseberry); F5648, H3710, M7129 BF, NMF Ribes glandulosum Grauer (skunk current); A2443, A2445, M6646, M6654.2, M6973, T5537, T5555 WC, NHS Ribes hirtellum Michx. (swamp gooseberry); F5665, F6483, M6655, M7060, S49, Ts.n. WC, NMF, BSS Ribes hudsonianum Richardson (northern black current); F5606, F6024, F6039, M6645 WC, NHS Ribes lacustre (Pers.) Poir. (bristly black current); M6729, P43782 BF, WC CR Ribes oxyacanthoides L. (northern gooseberry); M6988, S51, S91 NHS THR, WS Ribes triste Pall. (swamp red current); F5619, F6054, F6502, M6647, M6654.1, M7061, T5537 BF, WC, NHS

HALORAGACEAE (Water-milfoil Family)

Myriophyllum heterophyllum Michx. (various leaved water-milfoil); H3578 AP WS Myriophyllum sibiricum Komarov (spiked water-milfoil); F6562, F6530, H3580, M7312, T5193, T5273 AP Myriophyllum tenellum Bigelow (slender water-milfoil); S234, T5541 AP Myriophyllum verticillatum L. (water-milfoil); F6562 AP

HYPERICACEAE (St. John's Wort Family)

Hypericum ascyron L. (giant St. John's-wort); F6514, S271, T5288 W-NF Hypericum canadense L. (Canadian St. John's-wort); H3699 W-NF WS *Hypericum perforatum L. (common St. John's-wort); H4105, M7160 PB, NDF WS Triadenum fraseri (Spach) Gleason (marsh St. John's-wort); F6461, F6541, M7244, M7308, P44084, W-NF P44200, T52890

LAMIACEAE (Mint Family)

Agastache foeniculum (Pursh) Kuntze (blue giant hyssop); M6757, T5508 PB

*Ajuga genevensis L. (bugle); Clark 1264 BF Clinopodium vulgare L. (wild basil); H3374, H3749 NMF CR Dracocephalum parviflorum Nutt. (American dragonhead); M6818 PB WS *Galeopsis tetrahit L. (hemp-nettle); M7230, Ts.n. PB, NDMF

*Glechoma hederacea L. (creeping-Charlie); S106 BF, NDMF Lycopus americanus W. P. C. Barton (common water horehound); H3846, S285, T5482 NHS, W-NF Lycopus uniflorus Michx. (northern bugleweed); F6542, M6847, M6893, M7259, P44075, P44140, NHS, W-NF

T5273, T5481 Mentha canadensis L. (wild mint); S272, T5480 BF, NHS, WC



Monarda fistulosa L. (wild bergamot); H3721, M7222, T5077 PB, NDF Prunella vulgaris L. (self-heal); S180, T5263 NDMF, NDF, PB Scutellaria galericulata L. (marsh skullcap); F6550, H3772, P44091, Ts.n. WC, NHS, W-NF Scutellaria lateriflora L. (mad-dog skullcap); F6507, F6550, M6861, M7276, P44101, P44162 NHS, NMF Stachys arenicola Britton (marsh hedge nettle); H3947 PB, NHS Stachys tenuifolia Willd. (smooth hedge nettle); H4103, M6783 WC, PB

LENTIBULARIACEAE (Bladderwort Family) Utricularia intermedia Hayne. (northern bladderwort); F6538, M7233, P44135 AP WS Utricularia minor L. (lesser bladderwort); F6563, H3646, H3695, H3972 AP WS Utricularia vulgaris L. subsp. macrorhiza (J. Le Conte) R. T. Clausen (common bladderwort); F6531, AP H3620, P44216, S240

LINDERNIACEAE (False Pimpernel Family)

Lindernia dubia (L.) Pennell (false pimpernel); T5484 W-NF

LINNAEACEAE (Twinflower Family)

Linnaea borealis L. (twinflower); F6304, M6760, M7008, P44032, T5248 NDMF, WC, BG

LYTHRACEAE (Loosestrife Family) Decodon verticillatus (L.) Elliott (swamp loosestrife); S243 W-NF CR *Lythrum salicaria L. (purple loosestrife); F6477, H4115, S239 W-NF

MALVACEAE (Mallow Family) Tilia americana L. (basswood); Koch 12273 NMF, BF

MENYANTHACEAE (Buckbean Family) Menyanthes trifoliata L. (buckbean); P44089, S168, T5560 W-NF, BSS

MOLLUGINACEAE (Carpetweed Family) Mollugo verticillata L. (carpetweed); T5174, T5082 PB, NDF

MYRICACEAE (Bayberry Family) Comptonia peregrina (L.) J. M. Coult. (sweet fern); F6405, M6686, M6734, P43905, P44001, T5519 PB, BF Myrica gale L. (sweet gale); F6535, M7265 WC, NHS

MYRSINACEAE (Myrsine Family) Lysimachia ciliata L. (fringed loosestrife); F6151, S28 BF, NDMF

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Page  74 APPENDIX 1. (Continued)

Taxon Habitat Status

Lysimachia quadrifolia L. (whorled loosestrife); A2483, M7149, P43851, P43900, T5513, T5541 PB, NDF Lysimachia terrestris (L.) Britton, Sterns & Poggenb. (swamp candles); A2608, F6456, M7180, P44072, NHS, W-NF

P44123, P44197, T5561Lysimachia thyrsiflora L. (tufted loosestrife); F6391, H3623, M7059, P44141, S171, T5302 WC, W-NF Trientalis borealis Raf. (star flower); P43752, P44034, T5316 WC, PB, NDMF

NYMPHAEACEAE (Water Lily Family) Nuphar microphylla (Pers.) Fernald (small yellow pond lily); S250 AP Nuphar variegata Durand (yellow pond lily); F6527,S249, T5486 AP Nymphaea odorata Aiton. (fragrant water-lily); H3655, P44199, S236 AP WS

OLEACEAE (Olive Family) Fraxinus americana L. (white ash); S121 NMF Fraxinus nigra Marshall (black ash); M6967, M7054, M7236, T5546, T5557 NHS, WC Fraxinus pennsylvanica Marshall (green ash); F6582, S44, T5364 BF, NMF

ONAGRACEAE (Evening-Primrose Family) Chamerion angustifolium (L.) Holub (fireweed); A2606, M6791, M7192 PB, NDF WS Circaea alpina L. (small enchanter's nightshade); M6843, M6889, M7052, P44106, S14, T5090, T5246 WC, NHS Circaea canadensis (L.) Hill (broad-leaf enchanter’s-nightshade); H3708, H3877, M7139 NHS, NMF WS Epilobium ciliatum Raf. (willow herb); F6135, F6569, M7279, M7314, T5274 WC, NHS Epilobium coloratum Biehler (cinnamon willow herb); M6878, P44159 WC, NHS Epilobium leptophyllum Raf. (American marsh willow-herb); F6476, M6833, M6857, M7299 WC, W-NF Epilobium palustre L. (marsh willow-herb); M6896, S190 WC Ludwigia palustris (L.) Elliott (marsh purslane); H3588 W-NF WS Oenothera biennis L. (common evening primrose); H3726, H4100, P43899 PB, NDMF Oenothera clelandii W. Dietr., Raven & W.L. Wagner (evening primrose); M6744 PB CR Oenothera perennis L. (small evening primrose); Goessl 7644 Oenothera villosa Thunb. (evening primrose); M6780 PB WS

OROBANCHACEAE (Broom-rape Family) Agalinis paupercula (A. Gray) Britton (smooth false foxglove); H1918 W-NF Conopholis americana (L.) Wallr. (American cancer-root); A2604, M7100, M7110 NMF CR

*Euphrasia stricta J. F. Lehm. (drug eye-bright); M6856, P44215 PB WS



Melampyrum lineare Desr. (cow-wheat); A2488, F6412, M6733, M6761, M7142, M7190, P44023, T5562 PB, NDF Pedicularis canadensis L. (wood-betony); F6355, H3398, M6695, P44046, T5317 PB, NDF

OXALIDACEAE (Wood-sorrel Family) Oxalis acetosella L. subsp. montana (Raf.) D. Löve (mountain wood-sorrel); M6890, P44111, S179 WC, NHS WS Oxalis dillenii Jacq. (southern yellow wood-sorrel); M6805 BF, WC WS Oxalis stricta L. (wood-sorrel); H4073, S198 PB, NDF WS

PAPAVERACEAE (Poppy Family) Capnoides sempervirens (L.) Borkh. (pink corydalis); F5650, M7132, T5338, T5423 BG Sanguinaria canadensis L. (bloodroot); H3564 BF

PENTHORACEAE (Stonecrop Family) Penthorum sedoides L. (ditch stonecrop); T5283 W-NF

PHRYMACEAE (Lopseed Family) Mimulus glabratus Kunth (James' monkey-flower); H4054, M7305, T5191 W-NF Mimulus ringens L. (monkey-flower); F6560, T5284 WC

PLANTAGINACEAE (Plantain Family) Callitriche hermaphroditica L. (autumnal water starwort); H4016, M7291, T5278, T5357, T5510 AP SC Callitriche palustris L. (water starwort); F6015, M6903, M7270, T5280, T5281, T5378 AP Chelone glabra L. (turtle head); F6113, F6582, F6594, M6831, M6872, M7241, M7280, P44147 BF, W-NF Hippuris vulgaris L. (common mare's will); M7297 AP WS

*Linaria vulgaris Mill (butter and eggs); T5565 D

Nuttallanthus canadensis (L.) D. A. Sutton (blue toad-flax); Judziewicz 11400 PB, NDF *Plantago major L. (broad-leaved plantain); P43858 PB, NDF WS *Plantago patagonica Jacq. (woolly plantain); M6804 PB WS

Plantago rugelii Decne. (American plantain); H3903, P44165 BF, NDF CR

Veronica beccabunga L. var. americana Raf. (American brooklime); F6003, M6900, M7268 WC, NHS *Veronica longifolia L. (garden veronica); F6034 BF CR *Veronica officinalis L. (common speedwell); F6150, H3479, M7001, M7095 BF, NDMF

Veronica peregrina L. (purslane speedwell); Judziewicz 10701, Koch 12277 D Veronica scutellata L. (marsh speedwell); P44096 W-NF WS Veronica serpyllifolia L. (thyme-leaved speedwell); Gilbert,s.n. BF, NDMF

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Page  76 APPENDIX 1. (Continued)

Taxon Habitat Status

POLYGALACEAE (Milkwort Family) Polygala paucifolia Willd. (fringed polygala); F5620, F6315, P43850 WC, NDMF Polygala polygama Walter (racemed milkwort); M6737, M6797, M6813, P44052, S216, T5394 PB, NDF Polygala sanguinea L. (field milkwort); Garske 770 D

POLYGONACEAE (Buckwheat Family) Fallopia cilinodis (Michx.) Holub (fringed black bindweed); F6334, F6499, M7124, P43861 WC, NHS WS

*Fallopia convolvulus (L.) Ã. Löve (black bindweed); M6814, P43911, S204 BF, PB Fallopia scandens (L.) Holub (false buckwheat); F6334, H4101 BF, NDMF Persicaria amphibia (L.) Delabare (water smartweed); P44210, S247, T5550 BSS

*Persicaria hydropiper (L.) Delabare (marsh-pepper knotweed); M7286 W-NF WS Persicaria lapathifolia (L.) Delabare (nodding smartweed); F6537, T5178 W-NF

*Persicaria maculosa Gray (curly-top knotweed); H3969, H4104 W-NF WS Persicaria pensylvanica (L.) M. Gómez (Pensylvania knotweed); H3529 WC WS Persicaria punctata (Elliott) Small (dotted smartweed); H3765 W-NF WS Persicaria sagittata (L.) H. Gross (arrow-leaved tear-thumb); M6865, P44157, T5286 WC, W-NF Polygonella articulata (L.) Meisn. (coastal joint weed); M6801, P43887 PB WS Polygonum achoreum S. F. Blake (leathery knotweed); T5171 PB

*Polygonum aviculare L. (prostrate knotweed); T5101, T5198 NDF *Rumex acetosella L. (sheep sorrel); S89, S282, T5323 PB, NDF *Rumex crispus L. (curly dock); H4049, S154, T5233 BF, NHS *Rumex obtusifolius L. (bitter dock); H3745 BF WS

Rumex orbiculatus A. Gray (great water dock); F6522, M7243, M7269, P44161, T5083 W-NF

PORTULACACEAE (Purslane Family) *Portulaca oleracea L. (purslane); T5381 PB

RANUNCULACEAE (Buttercup Family) Actaea pachypoda Elliott (doll's-eyes); H3608, H3715 NMF WS Actaea rubra (Aiton) Willd. (red baneberry); F6132, F6313, F6480, M6876, T5519, T5530, T5546 BF, WC, NDMF Anemone americana (DC.) H. Hara (round-leaved hepatica); F5612, S41, T5307 BF, NMF Anemone canadensis L. (Canada anemone); S94, S138, T5326 BF, NDMF Anemone cylindrica A. Gray (thimbleweed); F6047 PB WS Anemone quinquefolia L. (wood anemone); F5622, F6351, M6653, M6682, M7007, P43753, T5309, BF, NDF, NMF



T5505, T5222 Anemone virginiana L. (thimbleweed); F6128, S23 BF Aquilegia canadensis L. (wild columbine); F6333, F5653, T5327 BF, NDF, NMF Caltha palustris L. (marsh marigold); F5638, F6322, M6652, M6962, M7042 WC, W-NF Clematis occidentalis (Hornem.) DC. (purple clematis); Sasse, s.n. WC Clematis virginiana L. (virgin's bower); F6572, H3939, H3980, H4056, T5326 BF, NHS Coptidium lapponicum (L.) Rydb. (Lapland buttercup); F6366 WC END Coptis trifolia (L.) Salisb. (goldthread); F5607, M6644, P44112, T5092, T5093, T5232 BF, WC, NMF Ranunculus abortivus L. (kidney-leaved buttercup); M6986, T5422 BF, NDMF

*Ranunculus acris L. (tall buttercup); F6082, M6961, S115 BF, NMF Ranunculus hispidus Michx. (bristly buttercup); H3594, H3716, H3820, M6971 BF,NHS,NMF WS Ranunculus longirostris Godr. (aquatic buttercup); H3759, H3862, M7277, S221, T5166, T5279, T5295 AP Ranunculus pensylvanicus L. f. (bristly buttercup); H3984, M7311 BF, NHS, NMF Ranunculus recurvatus Poir. (hooked buttercup); F6500, H3596, M6719, M7057 BF, NHS, NMF WS Ranunculus sceleratus L. (celery-leaf buttercup); H3396 WC WC Thalictrum dasycarpum Fisch. & Avé-Lall. (purple meadow-rue); F6120, F6512, M6715, M6867, BF, WC, NHS

T5490, T5507 Thalictrum dioicum L. (early meadow-rue); F5629, F6350, M6653, M6682, P43753 BF, WC, NHS WS

RHAMNACEAE (Buckthorn Family)

Ceanothus americanus L. (New Jersey tea); Clark 98 NDF Ceanothus herbaceus Raf. (Jersey tea); P43911, P44054, T5331 PB Rhamnus alnifolia L'Her. (alder-leaved buckthorn); A2444, H3587, M6657, S188, T5240, T5529 BF, NHS

*Rhamnus cathartica L. (common buckthorn); M7232, S167, S288 BF, NDMF, NHS, NMF

ROSACEAE (Rose Family)

Agrimonia gryposepala Wallr. (tall agrimony); F6045, F6081, F6497, P43872, S24 BF, NHS, NMF Agrimonia striata Michx. (roadside agrimony); T5269 WC Amelanchier arborea (F. Michx.) Fernald (common serviceberry); F6344, F6435, H3365, S62 BF,WC,NDMF WS Amelanchier bartramiana (Tausch) M. Roem. (mountain Juneberry); H3599 BSS Amelanchier interior Nielsen (inland serviceberry); F6481, H3904, M6669 WC Amelanchier laevis Wiegand (smooth serviceberry); H3597 BF, WC, NDMF Amelanchier sanguinea (Pursh) DC. (round-leaved serviceberry); F5655, F6043, F6344, M6671, M7120, BF, WC, NDMF

P43999, P44044, T5514Amelanchier spicata (Lam.) K. Koch (shadbush serviceberry); M6697 PB

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Page  78 APPENDIX 1. (Continued)

Taxon Habitat Status

Aronia melanocarpa (Michx.) Elliott (chokeberry); P44204 NDF, BSS Aronia x prunifolia (Marshall) Rehder (chokeberry); T5554 W-NF Comarum palustre L. (marsh cinquefoil); A2607, F6399, F6453, F6547, M7166, P44077, P44124 WC, NHS, BSS Crataegus chrysocarpa Ashe (hawthorn); A2545, P43748 BF Crataegus punctata Jacq. (dotted hawthorn); F6601 BF WS Crataegus submollis Sarg. (northern red haw); A2546, H4021 BF CR Crataegus succulenta subsp. macracantha (hawthorn); T5488 BF Drymocallis arguta (Pursh) Rydb. (prairie cinquefoil); M6752, P43920 PB WS

*Filipendula rubra (Hill) B. L. Rob. (queen-of-the-prairie); Clark 1344 D Fragaria vesca L. (woodland strawberry); F6311, M7131, S156, T5277 BF, WC, NMF Fragaria virginiana Mill. (wild strawberry); F5643, M6685, M7013, P43749, T5221, T5401 BF, WC, NMF Geum aleppicum Jacq. (yellow avens); F6055, F6083, F6492, M7108, T5275, T5512 BF, NDF, NDMF Geum canadense Jacq. (white avens); F6487 BF, NHS WS Geum fragarioides (Michx.) Smedmark (barren strawberry); F6305, P43790, P44015, P44025, S43, T5531 PB, NDF Geum laciniatum Murray (rough avens); F6492, M6993 NHS WS Geum macrophyllum Willd. (large-leaved avens); (H3964, F6010) WC, NDMF SC, WS Geum rivale L. (purple avens); M6726, M6859, M6898, M7044 BF, WC, NHS

*Potentilla argentea L. (silvery cinquefoil); H3893, T5516 PB, NDF Potentilla norvegica L. (rough cinquefoil); S155, S266, T5188 PB, NDF

*Potentilla recta L. (rough-fruited cinquefoil); S219 PB, NDF WS Potentilla simplex Michx. (common cinquefoil); F6356, P44014, P44051 PB, NDF WS Prunus americana Marshall (American plum); S52 BF, WC WS Prunus nigra Aiton (Canada plum); H3482, H3806, H3833, S26, T5356, T5376 BF, NDMF Prunus pensylvanica L. f. (pin cherry); H3603, M6664, M7122, M7191, T5087, T5370 BF, WC, NMF Prunus pumila L. (sand cherry); M6678, P44000, T52226, T5405 PB, NDF Prunus serotina Ehrh. (black cherry); S117, T5300 BF, NHS, NMF Prunus virginiana L. (chokecherry); F5637, F6340, F6482, F6599, M6659, M6698, P43785 BF, PB, NHS Rosa acicularis Lindl. (bristly rose); F5656, F6019 BF, WC, PB WS Rosa blanda Aiton (smooth rose); A2484, A2487, F6122, F6580, M6996, P44049, T5513 BF, PB, NDMF Rosa carolina L. (pasture rose); H3664, M6747, M6795 BF, PB, NDMF WS Rubus allegheniensis Porter (common blackberry); F6352, F6358, M7143, P44030, T5517 BF, PB, NDF Rubus canadensis L. (Canadian highbush blackberry); H3880, S93 NDF, NDMF Rubus flagellaris Willd. (short-stalk dewberry); H3384, H3385, M6767, P43903, P44003, P44036 PB, NDF WS



Rubus hispidus L. (swamp dewberry); H3686 WC WS Rubus parviflorus Nutt. (thimbleberry); M7125 BF, WC, NDMF WS Rubus pubescens Raf. (dwarf raspberry); F5640, M6656, M6723, M6968, M7045, S61, T5320 BF, WC, NDMF Rubus setosus Bigelow (bristly blackberry); F6379, F6387, H4018, S195 PB, BSS Rubus strigosus Michx. (red raspberry); M6990 BF, PB, NMF Sibbaldiopsis tridentata (Aiton) Rydb. (three-toothed cinquefoil); A2489, M6736, M6806, P44053, Ts.n. PB

*Sorbaria sorbifolia (L.) A.Braun (false spiraea); S258 BF Sorbus americana Marshall (American mountain ash); M6884 BF,NDMF,NHS,NMF WS

*Sorbus aucuparia L. (Eurasian mountain ash); S145 NDMF WS Spiraea alba Du Roi (white meadowsweet); F6465, F6546, M7170, M7267, P44079, P44113 WC, NHS, BSS WS Spiraea tomentosa L. (steeplebush); H2179, H4113 BSS WS

RUBIACEAE (Madder Family)

Galium aparine L. (cleavers); H3898, T5538 BF Galium asprellum Michx. (rough bedstraw); F6107, F6485, F6565, T5271, T5448 BF, WC, NMDF Galium boreale L. (northern bedstraw); S218 BF, PB Galium labradoricum (Wiegand) Wiegand (northern bog bedstraw); F6536, F6543 WC WS Galium tinctorium L. (stiff bedstraw); F6469, S141, S163, T5084 BF, NHS Galium trifidum L. (small bedstraw); F6536, F6543, M6982, M7304 BF, WC, NHS Galium triflorum Michx. (fragrant bedstraw); F6022, F6038, F6362, M7012, P44184, T5247 BF, NMF, NHS Houstonia longifolia Gaertn. (long-leaved bluets); P43915, T5170 PB Mitchella repens L. (partridgeberry); F6136, P43757, T5421 BF, WC, NDMF

SALICACEAE (Willow Family) Populus balsamifera L. (balsam poplar); S50, T5556 BF, NDF, NHS Populus grandidentata Michx. (big-tooth aspen); H3372, H3375, S104, T5523 BF, NDMF Populus tremuloides Michx. (quaking aspen); H4743 BF, NDMF, NHS

*Salix alba L. (white willow); T5363 BF Salix bebbiana Sarg. (Bebb's willow); F6509, H3657, H3988 BF, NHS, NMF Salix candida Willd. (sage-leaved willow); H2703 W-NF WS Salix discolor Muhl. (pussy willow); F6020, M6665, S33, T5484 BF, WC, NHS Salix eriocephala Michx. (heart-leaved willow); S159, T5484 NDF, NDMF Salix exigua Nutt (sandbar willow); F6587, S146 W-NF Salix humilis Marshall (prairie willow); M6699, P43747, S37, T5095, T5494, T5502, T5510, T5547 PB, NDMF Salix lucida Muhl. (shining willow); F6470, T5487 W-NF

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Taxon Habitat Status

Salix pedicellaris Pursh (bog willow); H3610, P44082, F6397, F6454, F6475 BSS Salix petiolaris Sm. (slender willow); F6473, S40 BSS Salix pyrifolia Andersson (balsam willow); F6370, F6474, M6670, P44095 P44206 WC, BSS WS

*Salix × fragilis L. (hybrid crack willow); S164 BF

SANTALACEAE (Sandalwood Family) Comandra umbellata (L.) Nutt. (bastard toadflax); F6363, H3477, H4047, T5330 PB, NDF, NDMF

SAPINDACEAE (Soapberry Family) Acer negundo L. (box elder); S126 BF, NHS WS Acer rubrum L. (red maple); P43750 BF, NDMF, NMF WS Acer saccharum Marshall (sugar maple); S92, S108 BF, NMF, NHS WS Acer spicatum Lam. (mountain maple); F5628, F5660, F6001, F6119, F6323, M6892, M7123, P43862 BF, NDF, NDMF

SARRACENIACEAE (Pitcher-Plant Family)

Sarracenia purpurea L. (pitcher plant); F6401, F6444, M7188, P44067, P44131, T5567 BSS

SAXIFRAGACEAE (Saxifrage Family) Chrysosplenium americanum Hook. (golden saxifrage); H3936, M6846, T5108, T5124 BF, WC, NHS Heuchera richardsonii R. Br. (alum root); H3393, H3552, P43917, T5204, T5301 PB Micranthes pensylvanica (L.) Haw. (swamp saxifrage); F5654, M6972, M7047, T5026 BF, WC, NHS Mitella diphylla L. (bishop's cap); F5621, F5641 BF, WC Mitella nuda L. (naked miterwort); F5613, H3379, H3481, M6673, T5249, T5417 BF, WC, NHS

SCROPHULARIACEAE (Figwort Family) Scrophularia lanceolata Pursh (early figwort); F6573, H4109, S136, S206, T5564 D *Verbascum thapsus L. (mullein); H4062, S276, Ts.n. PB

SOLANACEAE (Nightshade Family) Physalis virginiana Mill. (Virginia ground cherry); H3888, M6750, S217, T5180, T5407 PB *Solanum dulcamara L. (bittersweet night-shade); F6130, F6567 BF, WC, NHS

THYMELACEAE (Mezereum Family) Dirca palustris L. (leatherwood); A2601, H3589, M7098 BF, NDF, NMF

ULMACEAE (Elm Family) Ulmus americana L. (American elm); H3978, T5533 BF, NDF, NHS



URTICACEAE (Nettle Family) Laportea canadensis (L.) Wedd. (wood nettle); H3983, S165, T5554 BF, WC, NHS Urtica dioica L. (stinging nettle); T5574 BF, NHS

VALERIANACEAE (Valerian Family) *Valeriana officinalis L. (garden valerian); F6109, F6575, P43877 BF, NDF, WC

VERBENACEAE (Vervain Family) Verbena hastata L. (blue vervain); H3744, S267, T5345 NHS, W-NF

VIOLACEAE (Violet Family) Viola adunca Sm. (hook-spur violet); M6681, M7141, M7229 PB, NDF, NDMF WS Viola blanda Willd. (sweet white violet); H3583, H3899, Koch 7731 BF, WC Viola cucullata Aiton (blue marsh violet); H3399, S100 BF, WC, NHS WS Viola labradorica Schrank (dog violet); F5626, H3600, H3613, M7014, P43754, P44174, T5231 BF, PB, NDMF Viola macloskeyi F. E. Lloyd (smooth white violet); F5627, H3583, H3899, M6666, S77, S99, T5398 BF, WC, NHS Viola pedata L. (bird's-foot violet); M6689, S74, T5220 PB, NDF Viola pubescens Aiton (yellow wood violet); F5630, P43776, T5536, T5308 BF, NMF, NDF Viola renifolia A. Gray (kidney-leaved violet); H3498, H3900, S63 BF, WC, NDMF WS Viola sororia Willd. (common blue violet); H4034, S71 BF, ND, NHS WS

VITACEAE (Grape Family) Parthenocissus inserta (A. Kern.) Fritsch (grape woodbine); P43885 BF WS Parthenocissus quinquefolia (L.) Planch. (Virginia creeper); S182 BF, WC, NDMF WS


ACORACEAE (Sweet Flag Family) Acorus americanus (Raf.) Raf. (sweet-flag); H2834, T5491 AP *Acorus calamus L. (sweet-flag); H4150, S166 AP

ALISMATACEAE (Water-Plantain Family) Alisma triviale Pursh (northern water plantain); T5351, T5483 AP Sagittaria cuneata E. Sheld. (arum-leaved arrow-head); T5259, T5477 AP Sagittaria latifolia Willd. (arrow-head); F6516, H3774, H3850, T5194, T5292 AP Sagittaria rigida Pursh (stiff arrow-head); F6517, H3629, H3771, H3863, T5282, T5291, T5293, T5339 AP

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Taxon Habitat Status

ALLIACEAE (Onion Family) Allium tricoccum Aiton (wild leeks); H3711, M7107, S39, T5539 BF, NMF

ARACEAE (Arum Family) Arisaema triphyllum (L.) Schott (jack-in-the-pulpit); F6338, M6852, T5425 BF, WC, NHS Calla palustris L. (wild calla); F6384, F5663, M6839, M7173, M7307, P44074, P44125, P44155, WC, NHS, BSS

T5251, T5551 Lemna minor L. (common duckweed); F6540, M7237, M7266 AP WS Lemna trisulca L. (star duckweed); H3576, M7292 AP WS Lemna turionifera Landolt (red duckweed); H3955, T5163 AP Spirodela polyrrhiza (L.) Schleid. (greater duckweed); F6532, H3640, T5163 AP Symplocarpus foetidus (L.) W.P.C. Barton (skunk cabbage); S172, T5552 NHS

CONVALLARIACEAE (Lily-of-the-valley Family) Clintonia borealis (Aiton) Raf. (blue-bead lily); F6346, F6414, M6824, M6854, M6895, M7117, BF, WC M7175, P43765, P44179, T5426 *Convallaria majalis L. (European lily-of-the-valley); S131 BF, NHS Maianthemum canadense Desf. (wild lily-of-the-valley); F6094, F6365, F6413, M6347, M7116, BF, NDF, NMF

P43768, P44024, T5319, T5337 Maianthemum racemosum (L.) Link (false Solomon's-seal); F6365, H3545, P44183 BF, NDF, NMF WS Maianthemum stellatum (L.) Link (starry false Solomon's-seal); Schlapper, 35/03 PB Maianthemum trifolium (L.) Sloboda (false may flower); F6393, F6420, M7023, P44088, T5321, T5553 BSS, WC Polygonatum pubescens (Willd.) Pursh (downy Solomon's-seal); F5659, F6349, M7118, P43858 BF, NDF, NDMF WS Streptopus lanceolatus (Aiton) Reveal (rose twisted stalk); F6339, H3475, M6850, P43778, S18, T5416 BF, NDF, NDMF Uvularia grandiflora Sm. (bellwort); F5645, F6056, F6113, S66 NDMF, NMF Uvularia sessilifolia L. (sessile bellwort); F6117, F6357, F6408, M6799, M7066, M7115, P43786, T5312 BF, NDF, NDMF

CYPERACEAE (Sedge Family) Bulbostylis capillaris (L.) C. B. Clarke (dense tuft hair sedge); A2486 PB Carex adusta Boott (lesser brown sedge); M6759, P44027, P44037, T5405 NDF Carex alopecoidea Tuck. (foxtail sedge); H3811 NMF CR Carex aquatilis Wahlenb. (water sedge); H3999 BF, WC WS Carex arcta Boott (northern cluster sedge); Christensen s.n. NMF Carex arctata Boott (droopingwoodlandsedge);F5661,F6027,F6095,F6153, F6310,M7096, M7010, T5242 BF,NDF,NDMF



Carex assiniboinensis W.Boott (Assiniboine sedge); A3747, H3628, H3750 BF WS Carex aurea Nutt. (golden sedge); F6046 BF Carex backii W. Boott (Rocky Mountain sedge); A2603, F5649, H3419, M7127 BG SC, CR Carex bebbii (L. H. Bailey) Fernald (Bebb's sedge); H3907, M6704, T5173 WC Carex bromoides Willd. (brome-like sedge); H4007, H4010, M7039 BF, NHS, NMF CR Carex brunnescens (Pers.) Poir. (brownish sedge); F6041, M6705, M6957, M7046 BF, NHS, NMF Carex canescens L. (silvery sedge); M6987, M7031, M7038 WC, NHS, BSS Carex castanea Wahlenb. (chestnut sedge); F6152, M6707, M7004, M7094, M7109, P43751, P43764 BF, WC, NHS Carex chordorrhiza L. f. (cord-root sedge); F6425, H3783, M7257 BC, BSS WS Carex communis L. H. Bailey (fibrous root sedge); F5662, F6325, H3605, H3606, P43766, S55, T5478 BF, NDMF Carex comosa Boott (bristly sedge); H3909, M7235, P44154 NHS WS Carex crawfordii Fernald (Crawford's sedge); H3776, H4040, M6800 PB Carex crinita Lam. (fringed sedge); F6490, M6953, M7040, M7102, P44109, P44148, T5521 BF, WC, NHS Carex cryptolepis Mack. (northeastern sedge); Goessl 7675, Sulman 747 W-NF Carex debilis Michx. (northern weak sedge); M7168 NDMF, NMF Carex deflexa Hornem. (northern sedge); F6327, F6330, F6343, M6658, M7063 WC, NMF Carex deweyana Schwein. (Dewey's sedge); F6033, M6716, M7064, P43758, S57 BF, NDF, NMF Carex disperma Dewey (soft leaf sedge); F5632, F6012, M6643, M6712 WC, BSS Carex eburnea Boott (bristle-leaf sedge); H3966 WC WS Carex echinata Murray (star sedge); H3400, F6464 BF, WC, NHS WS Carex echinodes (Fernald) P.Rothr., Reznicek, & Hipp. (marsh straw sedge); F6062 BF CR Carex foenea Willd. (bronze-headed oval sedge); F6312, F6337, H3491 PB Carex gracillima Schwein. (graceful sedge); F6040, F6063, F6310, M6713, M6958, M7006, M7050, BF, NDMF, NMF

M7113, P43763 Carex granularis Willd. (limestone meadow sedge); F6050 BF, W-NF WS Carex gynandra Schwein. (nodding sedge); F6053, F6070, M6717, M6727, M6880, M6899 BF, NHS, BSS Carex hirtifolia Mack. (hairy sedge); H3812 BF CR Carex houghtoniana Dewey (Houghton's sedge); Davis s.n., Koch 5730 W-NF Carex hystericina Willd. (bottlebrush sedge); F6023, M6722, M6897, M7300, T5177 WC, NHS, AP Carex interior L. H. Bailey (inland sedge); F6000, M6728, S118 WC, NHS Carex intumescens Rudge (greater bladder sedge); A2602, F6148, F6307, F6508, M6706, M6960, M6995, BF, NDMF, NMF

M7006, M7041, M7092, M7176, P44103, T5250, T5539 Carex lacustris Willd. (lake sedge); F6371, F6549, S125 NHS, W-NF Carex lasiocarpa Ehrh. (woolly fruit sedge); F6381, F6451, F6471, P44129 BSS, W-NF WS

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Taxon Habitat Status

Carex leptalea Wahlenb. (bristly stalked sedge); F6009, M6873, M6956, M7048, P44102 WC, BSS Carex leptonervia (Fernald)Fernald(nervelesswoodlandsedge);F6026,F6324,H3592,H3601,M7036,P44102 BF, WC, NHS Carex limosa L. (muck sedge); F6439, H3617, P44087 W-NF WS Carex lurida Wahlenb. (shallow sedge); A459 W-NF Carex magellanica Lam. (boreal bog sedge); F6028, F6398, F6427, F6468, M6826, M6904, M7028, BSS, WC, W-NF

M7179, M7261 Carex muehlenbergii Schkuhr ex Willd. (Muhlenberg's sedge); M6753 PB CR Carex normalis Mack. (greater straw sedge); F6147 BF, NHS CR Carex oligosperma Michx. (few seed sedge); F6380, F6452, M7033, M7186, M7260, P44073, P44117 BSS, WC Carex ormostachya Wiegand (necklace spike sedge); F6324, P43777 BF, NDMF CR Carex pauciflora Lightf. (few flowered sedge); M7194, P44063 W-NF Carex peckii Howe (Peck's sedge); F5614, F6306, H3604, H3797 NMF, BF, NDMF WS Carex pedunculata Willd. (long-stalk sedge); F5608, M6648, P43756 BF, WC, NMF WS Carex pellita Willd. (broad-leaved wooly sedge); Koch 5718, Freckmann 29090 WC Carex pensylvanica Lam. (Pennsylvania sedge); M6684, P43780, P44031, S75 PB, NDMF, NMF Carex projecta Mack. (necklace sedge); F6486, F6503, M6710, M6714, M6978, M6985, M6994, M6998, BF, NHS, NMF

M7099, M7105, M7158, P44142 Carex pseudo-cyperus L. (false bristly sedge); M7302 AP Carex radiata (Wahlenb.) Small (eastern star sedge); H3998, M6981, M6983 BF, NHS CR Carex retrorsa Schwein. (deflexed bottlebrush sedge); F6140, H3748, T5527 BF, WC, W-NF Carex rosea Willd. (rosy sedge); H3595 BF, NHS CR Carex rostrata Stokes (beaked sedge); H2775, S273 BSS, WC WS Carex scabrata Schwein. (eastern rough sedge); F6084, H4011 BF, NMF Carex scoparia Willd. (broom sedge); H2302 BF, NMF, NHS Carex siccata Dewey (dry-spiked sedge); H3809, P44017 PB WS Carex sprengelii Spreng. (long-beaked sedge; Sprengel's sedge); H3805, P43791 BF CR Carex stipata Willd. (common fox sedge); F6495, M6955, M6725, P44107 BF, NDF, NMF Carex stricta Lam. (tussock sedge); M7032, T5261 BSS, WC, W-NF Carex tenera Dewey (quill sedge); F6145, P43998 BF, PB Carex tenuiflora Wahlenb. (sparse flower sedge); F6400 WC Carex tonsa (Fernald) E. P. Bicknell (shaved sedge); M6683, P44047, S56 PB, NDF Carex tribuloides Wahlenb. (blunt brome sedge); H4031 W-NF CR Carex trisperma Dewey (three seeded sedge); F6392, F6419, M7018, M7101, M7167, T5244 WC, BSS, NHS



Carex tuckermanii Dewey (Tuckerman's sedge); M6954, M7103, M7238, T5520 NHS, NMF Carex umbellata Willd. (early oak sedge); F5647 BF WS Carex utriculata Boott (yellow lake sedge); F6372, F6424, F6452, F6525, M7183, P44085, P44115 NHS, BSS, W-NF Carex vaginata Tausch (sheathed sedge); H3403, F6017 WC Carex vesicaria L. (blister sedge); M6979, P44205 NHS Carex viridula Michx. (little green sedge); S80 WC Carex vulpinoidea Michx. (fox sedge); H3662 W-NF Carex ×knieskernii Dewey (hybrid sedge); H3714, M7009 NMF CR Cyperus houghtonii Torr. (Houghton's nut sedge); M6817, P43893 PB WS Cyperus lupulinus (Spreng.) Marcks (slender sand sedge); Epstein s.n. PB Dulichium arundinaceum (L.) Britton (three-way sedge); H3632, P44195, S202, T5366, T5566 NHS, W-NF Eleocharis acicularis (L.) Roem. & Schult. (needle spike rush); S232 NHS, AP Eleocharis erythropoda Steud. (bald spike rush); H4024, T5176 AP, W-NF Eleocharis intermedia Schult. (intermediate spike rush); H3958, H4151, T5285 AP Eleocharis obtusa (Willd.) Schult. (blunt spike rush); Svenson (1971-01-01) AP, W-NF Eleocharis ovata (Roth) Roem. & Schult. (oval spike rush); H3758 W-NF Eleocharis palustris (L.) Roem. & Schult. (spike rush); F6556, M7296, P44196 AP, W-NF Eriophorum angustifolium Honck. (narrow leaf cotton grass); F6406, H4002, S82, T5230 BSS, W-NF Eriophorum chamissonis C.A. Meyer (Chamisso's cotton grass); H2152, S81, S107 W-NF Eriophorum gracile W. D. J. Koch (slender leaf cotton grass); F6443, H3622, H3723 W-NF Eriophorum tenellum Nutt. (conifer cotton grass); A2610, F6446, M7258, P44068, P44092, P44119, P44201 W-NF WS Eriophorum vaginatum L. (tussock cotton grass); F6389, F6445, M7029, T5229, T5572 WC, BSS Eriophorum virginicum L. (tawny cotton grass); A2611, F6418, F6447, M7187, M7249, P44066, P44116 BSS, W-NF Eriophorum viridicarinatum (Engelm.) Fernald (green-keeled cotton grass); S187, T5238 W-NF Rhynchospora alba (L.) Vahl (white beak sedge); F6448, P44086 W-NF WS Rhynchospora fusca (L.) W. T. Aiton (brown beak sedge); A2614, F6463, H3697 W-NF SC Schoenoplectus acutus (Bigelow) Ã. Löve & D. Löve (hard-stem bulrush); H3891 AP WS Schoenoplectus pungens (Vahl) Palla (chair-maker's rush); Salomaki 97/26 AP Schoenoplectus smithii (A. Gray) Sojak var. setosus (Fernald) S.G. Smith (Smith's bulrush); A2633, P44202 AP WS Schoenoplectus tabernaemontani (C. C. Gmel.) Palla (soft stem bulrush); F6520, H3769, M7301, S242, AP

T5358, T5479 Scirpus atrocinctus Fernald (black-girdled wool-grass); H3859, M7177 NHS, BSS WS Scirpus atrovirens Willd. (black bulrush); F6121, P44181 BF, NMF, NHS WS Scirpus cyperinus (L.) Kunth (wool-grass); F6440, M7284, P44099, P44132, P44163 BF, NHS, BSS

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Taxon Habitat Status

Scirpus microcarpus J. Presl & C. Presl (panicled bulrush); Allen s.n., Gerst s.n., Koch 12276 WC, W-NF Trichophorum alpinum (L.) Pers. (alpine bulrush); A2613, F6462 W-NF

ERIOCAULACEAE (Pipewort Family) Eriocaulon aquaticum (Hill) Druce (pipewort); T5542 AP

HEMEROCALLIDACEAE (Day-lily Family) Hemerocallis fulva (L.) L. (orange daylily); S263 BF CR

HYDROCHARITACEAE (Frog's-bit Family) Elodea canadensis Michx. (common waterweed); F6526, F6552, H3928, M7272, T5165, T5275, T5294 AP Elodea nuttallii (Planch.) H. St. John (slender waterweed); Ts.n. AP Najas flexilis (Willd.) Rostk. & Schmidt (slender naiad); H3633, S226 AP Vallisneria americana Michx. (eel-grass); F6551, M7275, Ts.n. AP

IRIDACEAE (Iris Family) *Iris pseudacorus L. (yellow-flag); S143 AP, W-NF Iris versicolor L. (wild blue-flag); F6386, F6441, M6840, M6976, M7245, M7252, P44081, P44128, AP, W-NF T5266, T5556 Sisyrinchium montanum Greene (mountain blue-eyed grass); P44019, P44020, T5481 PB

JUNCACEAE (Rush Family) Juncus balticus Willd. (arctic rush); Lahti 80 AP Juncus brachycephalus (Englem.) Buchenau (small headed rush); F6521 WC, BSS, W-NF Juncus brevicaudatus (Englem.) Fernald (narrow-panicle rush); F6467, H3700, P44203 WC, AP, W-NF WS Juncus effusus L. (soft-stem rush); F6478, H3974, M6991, M7294 NHS, BSS, WC WS Juncus greenei Oakes & Tuck (Greene's rush); H2724, H3996 WC WS Juncus interior Wiegand (inland rush); Fields 108 D Juncus nodosus L. (joint rush); H3995, M7298 AP Juncus tenuis Willd. (path rush); S199, T5197 BF, NDF, NMF Juncus vaseyi Engelm. (Vasey's rush); Judziewicz 11717 W-NF SC Luzula acuminata Raf. (hairy wood rush); F5611, M6642, P43755 BF, NDMF, NMF Luzula multiflora (Ehrhart) Lej. (common wood rush); P43789, S42 BF, NDF WS

LILIACEAE (Lily Family) Erythronium americanum Ker Gawl. (yellow trout lily); P43781, S47 BF WS



Lilium michiganense Farw. (Michigan Lily); F6035 BF, NHS Lilium philadelphicum L. (wood lily); M6735, T5369, T5379 PB

ORCHIDACEAE (Orchid Family) Arethusa bulbosa L. (dragon's mouth); S184 WC Calopogon tuberosus (L.) Britton, Sterns & Poggenb. (grass pink); Ts.n. W-NF Calypso bulbosa (L.) Oakes (fairy slipper); Photo WC END Corallorhiza maculata (Raf.) Raf. (spotted coral-root); F6415, Ts.n. BF, NDMF Corallorhiza trifida Châtel (early coral-root); M6675, M7058, P44108, S178 WC, NHS Cypripedium acaule Aiton (moccasin flower); F6430, M6825, M7016, P44137, T5402 WC Cypripedium parviflorum Salisb. var. makasin (Farw.) Sheviak (small yellow lady's slipper); F6369 WC SC Cypripedium parviflorum Salisb. var. pubescens (Willd.) O. W. Knight (yellow lady's slipper); Photo WC Cypripedium reginae Walter (showy lady's slipper); P43876 BF SC Goodyera pubescens (Willd.) R. Br. (downy rattlesnake plantain); Gockman s.n. NDF Goodyera repens (L.) R. Br. (creeping rattlesnake plantain); H3553, M6851, T5078 NDMF Goodyera tesselata Lodd. (tesselated rattlesnake plantain); H3480, M7214 PB, NDMF Malaxis unifolia Michx. (green adder's mouth); T5396, T5408 WC Neottia cordata (L.) Rich. (heart-leaved twayblade); A2442, F5623, M6674 WC Platanthera aquilonis Sheviak (northern green orchid); M6828, M6902 WC WS Platanthera clavellata (Michx.) Luer (club-spur orchid); Judziewicz 11977, Nekola s.n. WC Platanthera huronensis (Nutt.) Lindl. (green bog orchid); P44110, P44143, S175, T5353 NHS Platanthera obtusata (Pursh) Lindl. (blunt-leaved orchid); F6368, M6830, S212, T5105, T5385, T5558 WC Platanthera psycodes (L.) Lindl. (purple fringed orchid); M7246, S262 W-NF Pogonia ophioglossoides (L.) Ker Gawl. (rose pogonia); F6466 W-NF Spiranthes lacera (Raf.) Raf. (northern slender lady's tresses); H4093 PB

POACEAE (Grass Family)

*Agrostis gigantea Roth (red top); H3671, M7112, T5143, T5530 PB, NDMF, NMW Agrostis hyemalis (Walter) Britton, Sterns & Poggenb. (tickle grass); T5410, T5411, T5679 PB, NDF Agrostis perennans (Walter) Tuck. (autumn bent grass); F6141, M6837, M6881, P44162, T5480 BF, NDMF Agrostis scabra Willd. (rough bent grass); A2485, F6544, M6754, M6784, M7159, M7303 PB

*Agrostis stolonifera Willd. (creeping bent grass); H2811 W-NF WS Alopecurus aequalis Sobol. (short-awned foxtail); T5533 W-NF *Alopecurus pratensis L. (meadow foxtail); M6997, S53 NHS WS Ammophila breviligulata Fernald (beach grass); Castle 92-18 LSS

(Continued on next page)

Page  88 APPENDIX 1. (Continued)

Taxon Habitat Status

Andropogon gerardii Vitman (big bluestem); M6782, M7219, P43888, T5072 PB, NDF

*Anthoxanthum odoratum L. (sweet vernal grass); H4290, Clark 1260 D Aristida basiramea Vasey (fork-tipped three-awn grass); M6802, P43890 PB Brachyelytrum aristosum (Michx.) Branner & Coville (long-awned wood grass); F6501, M6703, BF, NDMF, NMF WS

M6883, M7037, M7093 Bromus ciliatus L. (fringed brome); F6042, F6088, F6488, M6708, M6886, M7106, M7220, P44104, BF, NDMF, NMF P44151, P44180

*Bromus inermis Leyss. (smooth brome); S137, S279, T5217 D Bromus kalmii A. Gray (prairie brome); A2490, M6739, M6787, M7218, P44012 PB, NDF, NDMF Bromus latiglumis (Shear) Hitchc. (hairy wood brome); H3674 NDF CR Bromus pubescens Spreng. (Canadian brome); H3371, H3720 BF, LNHS, NMF Calamagrostis canadensis (Michx.) P. Beauv. (blue-joint grass); F6091, F6149, F6388, F6417, F6426, BF, W-NF, NHS

F6592, M6841, M6879, M6966, M7285, P44083 Cinna arundinacea L. (common wood reed); F6491, H3611 NDMF, NHS CR Cinna latifolia (Goepp.) Griseb. (wood reed grass); F6103, M6821, M6871 BF, NMF, NHS

*Dactylis glomerata L. (orchard grass); S116, T5267 NMF Danthonia spicata (L.) Roem. & Schult. (poverty oat grass); M6731, M6741, M6742, M6815, M7097, PB, NDF, NDMF P44007, T5412, T5532, T5536 Dichanthelium acuminatum (Sw.) Gould & C. A. Clark (hairy panic grass); H4005, M7146, PB, NDF, NDMF

P44040, P44170 Dichanthelium columbianum (Scribn.) Freckmann (hemlock panic grass); T5409 PB Dichanthelium depauperatum (Muhl.) Gould (starved panic grass); M6755, P44021, P44056, PB, NDF

T5216, T5529 Dichanthelium linearifolium (Scrib.) Gould (linear-leaved panic grass); H4028, M7147 PB CR Dichanthelium meridionale (Ashe) Freckmann (mat panic grass); H4006, M6748 PB WS Dichanthelium oligosanthes (Schult.) Gould (red dot panic grass); H3669 NMF CR Dichanthelium xanthophysum (A. Gray) Freckmann (slender rosette grass); A2605, M6807, M7154, PB, NDF, NDMF

P43852, P44033, P44166, T5553 *Digitaria ischaemum (Schreb.) Muhl. (smooth crabgrass); P43892 PB WS *Echinochloa crusgalli (L.) P. Beauv. (barnyard grass); T5534 PB

Echinochloa muricata (P. Beauv.) Fernald (barnyard grass); H3483, T5103, T5534 PB Elymus canadensis L. (Canada wild rye); S256 LSS Elymus hystrix L. (bottlebrush grass); F6014, F6057, F6114, F6493, P43880, P44173, T5276 BF, NDF, NMF



*Elymus repens (L.) Gould (quack grass); H3655, T5526 PB, NDMF Elymus trachycaulus (Link) Gould (slender wheat grass); F6058, H3735, M6811, M7136, S153, S257 BF, PB, NMF Elymus virginicus L. (Virginia wild rye); H3970, F6106, P43879 BF, NHS Elymus wiegandii Fernald (Wiegand's wild rye); F6577, P43878 BF CR Eragrostis hypnoides (Lam.) Britton, Sterns & Poggenb. (creeping love grass); T5344 LSS Festuca saximontana Rydb. (Rocky Mountain fescue); H4058, M6692 PB, NDMF WS Festuca subverticillata (Pers.) E.B.Alexeev (nodding fescue); M7137 NMF WS

*Festuca trachyphylla (Hack.) Krajina (hard fescue); H3680 PB, NDF, NHS WS Glyceria borealis (Nash) Batch. (northern manna grass); F6385, F6533, H3793, M7283 NHS, AP Glyceria canadensis (Michx.) Trin. (rattlesnake grass); F6457, M7164, M7234, M7250, P44076, NHS, BSS

P44134, P44209 Glyceria grandis S. Watson (reed manna grass); H3625, T5481 NHS, W-NF Glyceria striata (Lam.) Hitchc. (fowl manna grass); F6021, F6504, F6518, M6959, M7055 BF, WC, W-NF Koeleria macrantha (Ledeb.) Schult. (June grass); H4030, P44010, T5528 PB, NDF Leersia oryzoides (L.) Sw. (rice cut grass); F6548, H3860, M7282, T5487 NHS, AP Milium effusum L. (wood millet); F6031, F6099, M7051, F6505, T5534 BF, NMF Muhlenbergia glomerata (Willd.) Trin. (marsh muhly); M7215, P43853, P43913 PB, NDMF WS Muhlenbergia mexicana (L.) Trin. (leafy satin-grass); M6779, P43882, P44177 PB, NDF, NDMF WS Oryzopsis asperifolia Michx. (rough-leaved rice grass); F6318, M6688, P43771, T5224, T5225 BF, NDMF, NMF

*Phalaris arundinacea L. (reed canary grass); F6564 W-NF, NHS, NMF

*Phleum pratense L. (Timothy); H4025, T5525 BF, NDF, PB Piptatherum pungens (Spreng.) Dorn (mountain rice grass); H4036, M6694 PB WS Poa alsodes A. Gray (woodland bluegrass); H4037, M7049 NHS WS

*Poa annua L. (annual bluegrass); Gilbert s.n. BF, NDF *Poa compressa L. (Canada bluegrass); H3681, S147 BF, PB, NDMF WS *Poa nemoralis L. (wood bluegrass); F6036, F6052, F6329, P43849 BF, NDF, NDMF WS

Poa palustris L. (marsh bluegrass); F6030, F6498, P44048, S70, T5527, T5535 WC, NHS

*Poa pratensis L. (Kentucky bluegrass); F6316, F6341, H4027, M6690 BF, NHS, NMF WS Poa saltuensis Fernald & Wiegand (old pasture bluegrass); F6326, F6329, H4013, M7015, M7065 WC, NMF Schizachne purpurascens (Torr.) Swallen (false melic grass); F6317, H3405, M7067, P44005, T5506 BF, PB, NDMF Schizachyrium scoparium (Michx.) Nash (little bluestem); M6786, P43896, T5073 PB

*Setaria pumila (Poir.) Roem. & Schult. (yellow foxtail); H3485 D WS Sorghastrum nutans (L.) Nash (Indian grass); P43922 PB Spartina pectinata Link (prairie cord grass); P43886 BF WS

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Page  90 APPENDIX 1. (Continued)

Taxon Habitat Status

Sphenopholis intermedia (Rydb.) Rydb. (slender wedge grass); A2600, M6724 BF, NMF WS Torreyochloa pallida (Torrey) Church (pale false manna grass); P44078 W-NF CR Zizania palustris L. (northern wild rice); S253 AP

PONTEDERIACEAE (Pickerel-weed Family)

Heteranthera dubia (Jacq.) Macmill. (water star grass); H3643, M7313, M7318, S224 AP WS


Potamogeton alpinus Balb. (alpine pondweed); H3768, H3892, H3956, S231, T5088, T5262, T5280, T5347 AP Potamogeton amplifolius Tuck. (large-leaved pondweed); S220 AP Potamogeton berchtoldii Fieber (slender pondweed); H3923 AP Potamogeton epihydrus Raf. (ribbon leaf pondweed); F6545, F6588, H3694, H3698, H3851 AP Potamogeton foliosus Raf. (leafy pondweed); H3693, H3637, M7293, S230, S246, T5297, T5486 AP Potamogeton friesii Rupr. (Fries's pondweed); F6519 AP Potamogeton gramineus L. (variable leaf pondweed); F6545, H3785, H3647, S227, T5346, T5234 AP Potamogeton illinoensis Morong (Illinois pondweed); H4066, M7274 AP WS Potamogeton natans L. (floating-leaf pondweed); F6524, H3634, H3696, M7310, T5279, T5361 AP Potamogeton nodosus Poir. (long-leaf pondweed); H3788 AP CR Potamogeton oakesianus J.W. Robbins (Oakes' pondweed); P44198, P44218 AP CR Potamogeton obtusifolius Mert. & W.D.J.Koch (blunt-leaf pondweed); M7273, Ts.n. AP Potamogeton praelongus Wulfen (white-stemmed pondweed); H3636, H4294, T5164, T5270 AP Potamogeton pusillus L. (small pondweed); F6539, H4293, P44217, T5278 AP WS Potamogeton richardsonii (A. Benn.) Rydb. (Richardson's pondweed); F6558, F6589, F6016, M7290, AP

T5167, T5168, T5276 Potamogeton robbinsii Oakes (Robbin's pondweed); H3574, H3787 AP Potamogeton spirillus Tuck. (spiral pondweed); S225, S228, S229, S235, S244 AP Potamogeton strictifolius A. Benn. (narrow-leaved pondweed); H3915, T5296 AP Potamogeton zosteriformis Fernald (flat-stemmed pondweed); H3762, H3914, F6529, T5274, T5299 AP Stuckenia filiformis (Pers.) Börner (narrow-leaved pondweed); Alverson 1803a AP Stuckenia pectinata (L.) Börner (sago pondweed); H3767, H3960, F6590, F6591, T5271, T5172, T5175, AP

T5277, T5285, T5511


Scheuchzeria palustris L. (pod-grass); F6436, M7184, P44074, P44120 WC, W-NF WS



SMILACACEAE (Carrion Flower Family) Smilax illinoensis Mangaly (Illinois carrion-flower); P43775.2 BF CR Smilax lasioneura L. (bristly greenbrier); H3840 BF WS

TRILLIACEAE (Trillium Family) Trillium cernuum L. (nodding trillium); F5639, F6096, F6320, M6718, M6875, M6984, P43770, T5420, T5562 BF, NDMF, NMF Trillium grandiflorum (Michx.) Salisb. (big white trillium); S64 BF, WC WS

TYPHACEAE (Cat-tail Family) Sparganium americanum Nutt. (American bur-reed); F6557 AP Sparganium angustifolium Michx. (narrow-leaved bur-reed); S238, Ts.n. AP Sparganium emersum Rehm. (green-fruited bur-reed); S237, S251, T5490, T5512 AP Sparganium eurycarpum Engelm. (common bur-reed); H4152, T5493 AP Sparganium fluctuans (Morong) B. L. Rob. (floating bur-reed); H3644, H3786, H3930, S241, T5489 AP *Typha angustifolia L. (narrow-leaved cat-tail); F6528, H3775, M7316 AP WS Typha latifolia L. (common cat-tail); M7317, T5352 AP *Typha × glauca Godr. (hybrid cat-tail); H3920 AP WS


APPENDIX 2. Prevalent ground layer species in each forest community type. Frequency of occurrence is the percentage of sites within the forest community type in which the species occurs. Frequency of common occurrence is the percentage of sites within the forest community type in which the species is widely distributed.

Frequency of Frequency of Species occurrence common occurrence

Boreal Forest

Eurybia macrophylla Rubus parviflorus Maianthemum canadense Pteridium aquilinum Aralia nudicaulis Cornus canadensis Carex gracillima Athyrium filix-femina Cornus sericea Equisetum arvense Anemone quinquefolia Calamagrostis canadensis

Northern Wet-Mesic Forest

Rubus pubescens Coptis trifolia Maianthemum canadense Trientalis borealis Cornus canadensis Clintonia borealis Gaultheria hispidula Mitella nuda Osmunda cinnamomea Aralia nudicaulis Orthilia secunda Carex disperma

Pine Barrens

Comptonia peregrina Corylus americana Prunus pumila Rubus flagellaris Carex pensylvanica Vaccinium angustifolium Quercus macrocarpa Quercus ellipsoidalis Andropogon gerardii Danthonia spicata Monarda fistulosa Hieracium aurantiacum Solidago nemoralis Schizachyrium scoparium Arctostaphylos uva-ursi Bromus kalmii

100 100 100 100 80 100 100 90 90 90 80 80

100 100 100 100 100 100 100 100 100 91 91 82

100 100 100 100 100 100 100 100 83 83 83 100 100 100 83 83

100 100 90 80 100 50 50 67 56 56 63 50

100 100 91 82 82 64 64 55 55 50 50 56

100 100 100 100 100 100 83 83 100 100 80 50 50 50 60 60

(Continued on next page)


APPENDIX 2. (Continued)

Frequency of Frequency of Species occurrence common occurrence

Northern Dry Forest Maianthemum canadense Pteridium aquilinum Oryzopsis asperifolia Carex pensylvanica Eurybia macrophylla Rubus allegheniensis Uvularia sessilifolia

Northern Dry Mesic Forest Maianthemum canadense Pteridium aquilinum Aralia nudicaulis Eurybia macrophylla Oryzopsis asperifolia Clintonia borealis Lonicera canadensis

Northern Hardwood Swamp Carex stipata Rubus pubescens Glyceria striata Carex intumescens Carex gracillima Onoclea sensibilis Carex projecta

Northern Mesic Forest Acer saccharum (seedlings) Maianthemum canadense Clintonia borealis Carex pensylvanica Aralia nudicaulis Quercus rubra (seedlings)

Northern Wet Forest Rhododendron groenlandicum Maianthemum trifolium Carex trisperma Vaccinium angustifolium

100 100 80 100 100 100 100

100 100 100 100 100 100 100

100 100 86 100 100 100 86

100 100 89 100 100 89

100 100 100 100

100 100 80 60 60 60 60

100 100 83 83 67 50 50

100 86 86 71 57 56 71

89 89 88 67 56 78

100 83 66 50


APPENDIX 3. Summary of data for the eight forest community types. Major tree dominants are derived from importance values in the Forest stand changes in the Bois Brule River 1968 to 2016 (Hlina et. al. 2020). Leading families are the percentage of all species in the forest community type that are in that family.

Category Data Boreal Forest Major tree dominants Populus tremuloides, Abies balsamea, Picea glauca, Pinus strobus Most prevalent ground Eurybia macrophylla, Maianthemum canadense, layer species Pteridium aquilinum, Aralia nudicaulis Leading families Cyperaceae (10.1%), Asteraceae (9.6%), Rosaceae (8.0%), Poaceae (7.4%) Ranunculaceae (5.3%) Average species richness 132 Total species richness 362 Ct 4.5 Cn 5.3 Non-native species (%) 14.9% Growth form counts Trees (26), Shrubs (50), Forbs (208), Graminoids (64), Vines (6) Northern Wet-Mesic Forest Major tree dominants Thuja occidentalis, Abies balsamea Most prevalent ground Rubus pubescens, Coptis trifolia, Maianthemum layer species canadense, Trientalis borealis, Cornus canadensis Leading families Cyperaceae (10.8%), Asteraceae (9.9%), Rosaceae (7.9%), Poaceae (5.8%) and Ericaceae (5.3%) Average species richness 98 Total species richness 299 Ct 5.8 Cn 6.1 Non-native species (%) 4.3% Growth form counts Trees (18), Shrubs (48), Forbs (142), Graminoids (51), Vines (1) Pine Barren Major tree dominants Pinus banksiana, Pinus resinosa Most prevalent ground Comptonia peregrina, Corylus americana, Prunus layer species pumila, Rubus flagellaris, Carex pensylvanica, Vaccinium angustifolium, Quercus macrocarpa, Quercus ellipsoidalis, Andropogon gerardii, Danthonia spicata, Monarda fistulosa Leading families Asteraceae (17.8%), Poaceae (12.0%), Rosaceae (11.6%) Average species richness 79 Total species richness 207 Ct 4.1 Cn 4.8 Non-native species (%) 16.4% Growth form counts Trees (12), Shrubs (27), Forbs (118), Graminoids (32), Vines (1)


APPENDIX 3. (Continued)

Category Data Northern Hardwood Swamp Major tree dominants Fraxinus nigra, Thuja occidentalis Most prevalent ground Carex stipata, Alnus incana, Rubus pubescens, layer species Glyceria striata Leading families Cyperaceae (15.9%), Asteraceae (11.4%), Poaceae (10.6%) and Rosaceae (9.3%). Average species richness 92 Total species richness 307 Ct 5.0 Cn 5.6 Non-native species (%) 10.7% Growth form counts Trees (19), Shrubs (43), Forbs (164), Graminoids (75), Vines (6) Northern Dry Forest Major tree dominants Pinus resinosa Most prevalent ground Maianthemum canadense, Pteridium aquilinum, layer species Oryzopsis asperifolia Leading families Asteraceae (11.7%), Rosaceae (11.2%), Poaceae (11.2%), Cyperaceae (6.6%) and Ericaceae (6.1%) Average species richness 75 Total species richness 209 Ct 4.4 Cn 5.2 Non-native species (%) 15.8% Growth form counts Trees (14), Shrubs (42), Forbs (104), Graminoids (33), Vines (4) Northern Dry Mesic Forest Major tree dominants Pinus resinosa, Abies balsamea, Pinus strobus Most prevalent ground Maianthemum canadense, Pteridium aquilinum, layer species Aralia nudicaulis, Eurybia macrophylla Leading families Asteraceae (11.3%), Rosaceae (9.2%), Poaceae (8.3%), Cyperaceae (5.4%) and Ericaceae (5.4%) Average species richness 94 Total species richness 263 Ct 5.5 Cn 4.8 Non-native species (%) 12.9% Growth form counts Trees (27), Shrubs (38), Forbs (133), Graminoids (35), Vines (7) (Continued on next page)


APPENDIX 3. (Continued)

Category Data Northern Mesic Forest Major tree dominants Acer saccharum, Tilia americana, Acer rubrum Most prevalent ground Acer saccharum, Maianthemum canadense, layer species Clintonia borealis Leading families Cyperaceae (10.7%), Poaceae (9.4%), Asteraceae (9.0%), Rosaceae (7.7%), Ranunculaceae (5.6%) and Liliaceae (3.4%) Average species richness 85 Total species richness 242 Ct 5.2 Cn 5.4 Non-native species (%) 10% Growth form count Trees (20), Shrubs (37), Forbs (125), Graminoids (50), Vines (2) Northern Wet Forest Major tree dominants Picea mariana, Larix laricina Most prevalent ground Rhododendron groenlandicum, Maianthemum trifolium layer species Leading families Cyperaceae (19.0%), Ericaceae (13.1%), Asteraceae (6.4%), Rosaceae (6.0%), and Poaceae (4.8%) Average species richness 36 Total species richness 156 Ct 5.7 Cn 6.1 Non-native species (%) 6.4% Growth form count Trees (10), Shrubs (22), Forbs (31), Graminoids (21), Vines (0)