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ADDITIONS TO THE VASCULAR FLORA,AND NOTES ON THE PHYTOGEOGRAPHY, OF LAKE COUNTY, MICHIGAN

Bradford S. Slaughter Orbis Environmental Consulting P.O. Box 10235 South Bend, IN 46680 bslaughter@orbisec.com

Amanda K. Klain University of Michigan Herbarium Research Museums Center 3600 Varsity Drive Ann Arbor, MI 48108-2228 aklain@umich.edu

ABSTRACT

The vascular flora of interior northern Lower Michigan is poorly represented in herbaria, with several counties ranking among the most sparsely collected in the Upper Great Lakes region. We initiated collecting in one of these counties, Lake County, in 2013. At the time, we were aware of 658 collections from the county, representing 437 vascular plant species. From 2013 through 2018, we added 698 specimens, representing 631 species, 559 of which were previously undocumented. As of early 2019, a total of 1,006 species for the county are confirmed, placing Lake County among the best-collected counties in northern Lower Michigan. Among the most noteworthy collections were 12 state-listed (Endangered, Threatened, and Special Concern) species and 30 species at their statewide or Lower Peninsula range limits. Additional collecting in Lake County and surrounding counties is recommended to catalog regional diversity and to substantiate or amend species distributions.

KEYWORDS: Lake County, Michigan, vascular flora, plant species, new records, phytogeography, tension zone

INTRODUCTION

Nearly 2,900 vascular plant species1 growing outside of cultivation have been collected from Michigan, including approximately 1,800 native species and 1,100 non-native species (MICHIGAN FLORA ONLINE 2011). Floristic richness in Michigan follows a well-documented latitudinal gradient, declining from approximately 1,300–1,700 species per county in the southernmost counties to 1,000 species or fewer in the most depauperate northern counties (MICHIGAN FLORA ONLINE 2011). Although this diversity gradient is broadly supported

1 MICHIGAN FLORA ONLINE (2011) and its predecessors treat species as the basic taxonomic unit, with very few exceptions (e.g., instances in which both native and non-native subspecies or varieties are present, as is the case for Phragmites australis and Veronica beccabunga). In this paper, “species†refers to searchable taxa on MICHIGAN FLORA ONLINE, with the understanding that associated statistics may include one or more infraspecific taxa.

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by specimen records, many counties are poorly collected due to a variety of biological and cultural factors, such as the absence or presumed absence of unusual or attractive habitats, low road density, or distance from botanically active academic institutions and biological field stations (Fritsch 1993; MICHIGAN FLORA ONLINE 2011). Several counties in interior northern Lower Michigan are among the most sparsely collected in the Upper Great Lakes region (MICHIGAN FLORA ONLINE 2011; Kartesz 2015). Additional collecting in these counties is necessary for the enumeration of regional plant richness, resolution of species range limits, and detection of changes in the abundance and distribution of plant species, particularly those expected to be impacted by climate change. This paper documents our collecting efforts in Lake County from 2013 through 2018.

Description of the Study Area

Lake County covers 1,490 km2 (38.6 ¥ 38.6 km), centered at approximately 43.995°N, 85.811°W in northern Lower Michigan (Figure 1). Baldwin, the county seat, is 105 km (65 mi) north of Grand Rapids and 48 km (30 mi) east of Lake Michigan. The county is sparsely populated, with fewer than 12,000 permanent residents clustered in a few population centers, especially Baldwin,

FIGURE 1. Location of Lake County in northern Lower Michigan, USA, and the location of the townships. (Basemap: State of Michigan–Michigan GIS Open Data.)

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Idlewild, Chase, Luther, and the shores of several of the larger lakes (Lake County Planning Commission 2012; United States Census Bureau 2018). Approximately 7% of the land area is utilized for agriculture, much of it as forage- land, with limited areas of row crops (Lake County Planning Commission 2012; United States Department of Agriculture 2014). Forest cover exceeds 80%, much of it on state and federal land within the Pere Marquette State Forest and Manistee National Forest (Pugh et al. 2012).

Climate

Lake County has a warm-summer humid continental climate (i.e., Köppen– Geiger Classification Dfb), characterized by cold winters, warm summers, and no dry season (Kottek et al. 2006). Monthly average temperature (1981–2010 normals) at Baldwin ranges from –5.9°C (21.4°F) in January to 20.9°C (69.6°F) in July, with an annual average of 7.8°C (46.0°F), growing season (May– October) average of 16.0°C (60.8°F), and an average freeze-free period of 127 days (Arguez et al. 2010). Average liquid-equivalent annual precipitation at Baldwin is 87.9 cm (34.6 in) (Arguez et al. 2010). Average annual snowfall ranges from 183 cm (72 in) to 213 cm (84 in), with slightly lower amounts in the far southeastern part of the county and slightly higher amounts in the northern townships (Midwestern Regional Climate Center 2018a). Regional climate is trending warmer and wetter. Average annual temperature in the region has increased at a rate of 0.1°C/decade (0.2°F/decade) since 1895 and is projected to exceed 11°C (52°F) at Baldwin by 2100 (Girvetz et al. 2009; Midwestern Regional Climate Center 2018b). Average annual precipitation has increased at a rate of 1.6 cm/decade (0.6 in/decade) since 1895 and is projected to exceed 95 cm (37 in) at Baldwin by 2100 (Girvetz et al. 2009; Midwestern Regional Climate Center 2018b).

Landforms

Lake County is mantled by thick Wisconsin-aged glacial sediment overlying Mississippian (358.9–323.2 Ma) and Pennsylvanian (323.2–298.9 Ma) sandstone, shale, and limestone bedrock (Nicholson et al. 2004). The western townships consist of gently sloped outwash plains surrounding several hilly tracts associated with the Lake Border Moraine (Leverett and Taylor 1915) (Figure 2). Elevations range from approximately 200 m (650 ft) along the Pere Marquette River to 365 m (1,200 ft) on some of the moraine summits (United States Geological Survey 2017). The eastern townships comprise a portion of the Saginaw– Lake Michigan interlobate, a region marked by hummocky terrain on sandy glacial deposits exceeding 200 m (650 ft) in depth (Blewett et al. 2009; Schaetzl et al. 2013). Elevation generally exceeds 320 m (1,050 ft), with a high point of approximately 426 m (1,400 ft) in rural Pinora Township (United States Geological Survey 2017). These two primary physiographic regions are separated by a north and south trending escarpment traversing Newkirk, Cherry Valley, and Yates townships (Schaetzl et al. 2013).

Despite its namesake, fewer than 450 lakes or ponds of 0.04 ha (0.1 ac) or larger occur within Lake County, collectively covering less than 1% of its surface

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FIGURE 2. Topographic contour map of Lake County with 10-m contours. A prominent north and south trending escarpment (see arrow) separates the outwash plains and embedded hilly regions of western and central Lake County from the hummocky, high-elevation terrain of the eastern townships. (Basemap: State of Michigan–Michigan GIS Open Data; Topographic contours: U.S. Geological Survey–The National Map.)

area (Annable et al. 1991; Wolfson 2009). Only four of these lakes (Big Star, Wolf, Bass, and Syers) exceed 40 ha (100 ac) (Michigan Department of Natural Resources and Environment 2010). Lakes and drier kettles supporting wetland or upland vegetation are clustered in three areas: near Big Star Lake in the southwest; near Baldwin and Idlewild in Webber, Pleasant Plains, and Yates townships; and in the central portions of Elk and Sauble townships in the northwest.

Nine watersheds occur within the county (USDA, NRCS 2018). Major drainage networks are the Pine, Little Manistee, and Pere Marquette rivers and their tributaries. Lake County valley segments of all three rivers and their tributaries are characterized by cold water (mean July temperature <17.5°C/63.5°F), and all are designated trout streams, supporting noteworthy brown trout, rainbow trout, and Chinook salmon fisheries (Wang et al. 2012; O’Neal and Kolb 2015; Michigan Department of Natural Resources [MDNR], Institute for Fisheries Research 2017). Springs and headwater streams are frequent. Small portions of the northwestern and far eastern townships are drained by tributaries of the Big Sable and Muskegon rivers, respectively.

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Soils

The two dominant soil orders in Lake County are Entisols and Spodosols (USDA, NRCS 1998). Entisols are poorly differentiated from parent sediments and lack diagnostic horizons (USDA, NRCS 2015). They are characteristic of young landscapes where soil development has taken place over a relatively short period of time, where the parent material is abrasion-resistant (e.g., silica), or where organic litter (e.g., pine needles) is absent or recalcitrant (Schaetzl 2009; USDA, NRCS 2015). In the Great Lakes region, these conditions occur on sand dunes and dry outwash plains (Schaetzl 2009). In Lake County, outwash plains are principally distinguished by Entisols such as Plainfield and Coloma sands (USDA, NRCS 2017) (see Figure 3 for a generalized map of surficial soil textures). These soils are droughty, nutrient-poor, and acidic and have negligible silt and clay content.

Like Entisols, Spodosols are characteristic of young landscapes and sandy parent materials, but they support distinctive horizons. Spodosols are the dominant soil order of northern Michigan, forming under mixed forest cover where seasonal water infiltration, often associated with a winter snowpack, leaches organic matter, aluminum, and iron from surface horizons to a distinctive subsurface known as a spodic horizon (Schaetzl 2009; USDA, NRCS 2015). In Lake

FIGURE 3. Generalized soil surface textures in Lake County. Note the preponderance of sandy textures and the restricted distribution of loamy textures. (Basemap: State of Michigan–Michigan GIS Open Data; Soil data: U. S. Department of Agriculture–Natural Resources Conservation Service.)

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County, Spodosols are found on sandy moraines, where common map units include Montcalm-Graycalm complex, Grattan sand, and Rubicon sand (USDA, NRCS 2017). These soils share many physical characteristics with the Entisols, but some of the map units have higher clay and silt fractions and lower acidity (USDA, NRCS 2017).

Two minor soil orders comprise the remainder of Lake County soils: Alfisols and Histosols. Alfisols are characterized by the accumulation of clay in a subsurface layer known as an argillic horizon (USDA, NRCS 2015). They tend to form on loamy glacial tills and under deciduous forest canopies and to have higher nutrient content and pH values relative to the sandier Entisols and Spodosols (Schaetzl 2009; USDA, NRCS 2015). Alfisols are the dominant soil order in southern Lower Michigan, but also occur sporadically in northern Michigan. In Lake County, Alfisols occur locally at the western and eastern margins of the county, with the largest area associated with a ground moraine east and south of Chase (USDA, NRCS 2015). Typical map units include Emmet-Montcalm complex, Kawkawlin loam, and Nester sandy loam (USDA, NRCS 2017).

Histosols are deep (>40 cm) accumulations of partially decayed organic matter, typically forming where soil saturation limits aerobic decomposition of plant materials (USDA, NRCS 2015). Histosols comprise approximately 12% of the land surface in Michigan (Schaetzl 2009). They are scattered throughout Lake County and occur in patches of up to several thousand hectares (e.g., in the Bald- win-Luther, Baylor, and Bear swamps). Typical soil units include Tawas- Roscommon association (shallow, circumneutral muck over sand), Lupton muck (deep, slightly alkaline muck), and Loxley peat (deep, acidic peat, as in bogs) (USDA, NRCS 2017).

Vegetation

The combination of poor soils and frequent fire historically favored the development of oak–pine forests and barrens on sandy outwash plains (Harvey 1920, 1922; Albert 1995; Albert and Comer 2008) (Figure 4). Sandy moraines supported similar communities, although topographic amelioration of the fire regime and other factors resulted in heavier forest cover. Loamier moraines supported white pine–mixed hardwood forest and, where fires were especially infrequent, beech–sugar maple–hemlock forest (Albert and Comer 2008). A variety of forested and non-forested wetlands occurred on wet mineral and organic soils throughout the county (Figure 5). Vegetation cover types mapped for the county by Comer et al. (1995) and their equivalent natural community types (Cohen et al. 2014) are listed in Table 1; generalized land cover circa 1800 is depicted in Figures 6 and 7.

Logging, fires, and land-clearing that took place during the timber boom of the late 1800s irrevocably altered the structure and, to a lesser degree, the composition of plant communities throughout Lake County (Harvey 1920, 1922; Albert 1995). Modern timber and wildlife habitat management, clearing and cultivation, drainage, development, fire suppression, and the introduction of pests, pathogens, and non-native species have all since impacted the majority of the vegetation in the county. Among the most significant changes to undeveloped

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FIGURE 4. Oak–pine forests and openings (both natural and resulting from human activity) are emblematic of the sandy outwash plains in Lake County. Photo by A. K. Klain.

FIGURE 5. Rich conifer swamp dominated by Thuja occidentalis (northern white-cedar) along the Little Manistee River, Peacock Twp. Photo by A. K. Klain.

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TABLE 1. Lake County circa 1800 vegetation types (Comer et al. 1995) and equivalent MNFI natural community types (Cohen et al. 2014) organized by characteristic physiographic settings. Some natural community types not listed here, such as dry sand prairie, occur or may occur in Lake County (Cohen et al. 2014) but were not mapped by Comer et al. (1995) due to their small size or other factors.

Landform/Soil circa 1800 Vegetation Type Natural Community Type Outwash Plain Entisols; Spodosols Jack Pine–Red Pine Forest White Pine–White Oak Forest White Pine–Red Pine Forest Mixed Pine–Oak Forest Dry Northern Forest Dry-mesic Northern Forest Pine Barrens Pine Barrens Oak–Pine Barrens Oak–Pine Barrens End Moraine; Interlobate Moraine Spodosols; Alfisols Beech–Sugar Maple–Hemlock Forest Hemlock–White Pine Forest White Pine–Mixed Hardwood Forest Mesic Northern Forest Mesic Northern Forest Dry-mesic Northern Forest Mixed Pine–Oak Forest White Pine–White Oak Forest Dry-mesic Northern Forest Ground Moraine Alfisols; Spodosols Beech–Sugar Maple–Hemlock Forest White Pine–Mixed Hardwood Forest Mesic Northern Forest Mesic Northern Forest Dry-mesic Northern Forest Depression Histosols; Alfisols; Spodosols; Entisols Cedar Swamp Mixed Conifer Swamp Rich Conifer Swamp Rich Conifer Swamp Hardwood–Conifer Swamp Poor Conifer Swamp Bog Mixed Hardwood Swamp Black Ash Swamp Northern Hardwood Swamp Shrub Swamp/Emergent Marsh Northern Shrub Thicket Northern Wet Meadow Emergent Marsh Submergent Marsh Intermittent Wetland

vegetation are altered forest canopy dominance patterns (e.g., from pine to oak and aspen or stump prairies on uplands and from northern white-cedar to red maple, tag alder, and cat-tail in wetlands), woody encroachment and reduction of herbaceous communities in remnant barrens, and degradation and reduction of shoreline wetlands due to water level manipulations and beaver activity (Albert 1995; Stearns 1997; Haxby et al. 2013a, 2013b).

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FIGURE 6. Circa 1800 upland vegetation, generalized from Comer et al. (1995). Dry and dry-mesic forests dominated by pines and oaks with embedded pockets of barrens characterized most of the county. Mesic forests were concentrated on finer-textured soils in the eastern townships. (Basemap and Data layer: State of Michigan–Michigan GIS Open Data.)

To date, Michigan Natural Features Inventory (MNFI) has identified 15 element occurrences, or high-quality examples, of natural communities within Lake County (MNFI 2018). Ten natural community types are represented, including four intermittent wetlands, three bogs, and one each of dry sand prairie, floodplain forest, northern hardwood swamp, oak–pine barrens, pine barrens, poor conifer swamp, rich conifer swamp, and wet-mesic sand prairie. Collectively, these 15 sites encompass approximately 595 ha (1,470 ac), or 0.3% of the county’s area (MNFI 2018). While Lake County has not been thoroughly inventoried by MNFI, these figures nevertheless indicate the extent and severity of human impacts to the vegetation and the resulting scarcity of sites resembling circa 1800 conditions.

Previous Floristic Work

The oldest known specimens from Lake County were collected between 1888 and 1891 by W. J. Beal, Professor of Botany at Michigan Agricultural College (MAC) (MICHIGAN FLORA ONLINE 2011) (one rare species probably collected by Beal is shown in Figure 8). Beal’s collections coincided with his work establishing and conducting research at the Baldwin Agricultural Experiment

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FIGURE 7. Circa 1800 wetland vegetation, generalized from Comer et al. (1995). Several large swamp forests occurred (and persist) in Lake County, especially at the base of the escarpment between Baldwin and Luther. Other wetland communities occurred mostly as small patches. (Basemap and Data layer: State of Michigan–Michigan GIS Open Data.)

Station, one of several small plots of land in northern Lower Michigan rented by MAC for the purposes of agricultural experimentation on jack pine plains (Michigan State Board of Agriculture 1888). After Beal ceded his role with the Experiment Stations in 1891, three decades passed with no new collections. From 1922 to 1938, 51 specimens were collected, many of these by the prominent botanist E. J. Palmer of the Arnold Arboretum at Harvard University (MICHIGAN FLORA ONLINE 2011).

Over half of the specimen records for Lake County date to the period from 1939 to 1976, coinciding with activity of its three most prolific collectors, C. W. Bazuin (130 specimens, 1939–1962), E. G. Voss (165 specimens, 1955–1973), and G. P. Stegmier (48 specimens, 1976) (MICHIGAN FLORA ONLINE 2011). Collecting slowed after 1976, with 38 specimens recorded from the period 1978– 2007. In sum, a total of 658 collections, representing 437 vascular plant species, were mapped for Lake County at the initiation of our study (MICHIGAN FLORA ONLINE 2011). This total included 410 seed-bearing taxa mapped in Field Manual of Michigan Flora (Voss and Reznicek 2012), 14 taxa subsequently added to MICHIGAN FLORA ONLINE, and 13 ferns and other spore- bearing taxa not treated in the field manual or its predecessors (Voss 1972, 1985,

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FIGURE 8. Cirsium hillii (Hill’s thistle) was collected from the vicinity of Baldwin in July 1888, presumably by

W. J. Beal. It remains locally frequent in the area. Newaygo Co., Michigan. Photo by B. S. Slaughter. 1996). Specimens collected just prior to the publication of Voss and Reznicek (2012) have since been added to MICHIGAN FLORA ONLINE (2011), increasing the total number of individually checked and confirmed2 specimens collected prior to our work to 681 (MICHIGAN FLORA ONLINE 2011).

Early collections (1888–1930s) were made primarily from Baldwin and environs and consist nearly exclusively of terrestrial species. Collection of aquatic plants began in earnest in the late 1930s at Big Star Lake under the auspices of the Institute for Fisheries Research of the Michigan Department of Conservation (Voss 1972; MICHIGAN FLORA ONLINE 2011). Lake regions remained favored collecting locales through the 1970s. In addition to Big Star Lake, important stations included Bowman and Roby lakes in Lake Township; Cashion, Government, Hamlin, and Mench lakes near Baldwin; Syers and Little Syers lakes in Peacock and Eden townships; Totten Lake in Ellsworth Township; and Olga Lake

2 From MICHIGAN FLORA ONLINE (2011): “On this site nothing is added to the flora or to any map without the supporting specimens having been individually checked and confirmed, both as to their identity and their status as wild plants collected in Michigan. We note this explicitly because the increased online availability of herbarium specimen label data . . . means that misidentifications that once slumbered anonymously in folders in herbaria are now enshrined in credible websites for all to see.â€

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in Dover Township. More recent collections were obtained from the Manistee National Forest and a few other scattered locations (MICHIGAN FLORA ONLINE 2011).

METHODS AND MATERIALS

Site Selection

Initial collections in 2013 were gathered from the authors’ field project sites on the Manistee National Forest (HMNF) and private land enrolled in the Michigan Department of Natural Resources (MDNR) Landowner Incentive Program. Additional collection sites were identified through a combination of aerial photographic interpretation, field reconnaissance, drive-by inspections, site leads derived from MNFI (2018), herbarium records, and suggestions and field data from biologists and foresters with the Baldwin Field Office of MDNR and the Baldwin/White Cloud Ranger District of the Manistee National Forest. Lists of target species were generated from these leads and revised during and after each field season.

Collections

Voucher specimens were collected during the period from May 2013 to September 2018. GPS coordinates and notes on abundance, habitat, and associated species were recorded for most collections. Determinations were made by the authors, and specimens were examined and redetermined, where necessary, by A. A. Reznicek at the University of Michigan Herbarium (MICH). All specimens were mounted, databased, and deposited at MICH. Specimen data are searchable on MICHIGAN FLORA ONLINE (2011), coincident with periodic updates of the maps and specimen records.

RESULTS

A total of 698 specimens were collected, representing 631 species. Among these were 559 species newly vouchered3 from Lake County, including 409 native species and 150 non-native species (MICHIGAN FLORA ONLINE 2011) (Table 2; Appendix 1). Twelve species listed as Endangered, Threatened, or Special Concern in Michigan (MNFI 2009) were collected, including eight species not previously collected from Lake County, two species previously collected from other locations within the county, and two species that had been reported but not substantiated with collections (MICHIGAN FLORA ONLINE 2011; MNFI 2018) (Table 3; Appendix 1).

Specimens were collected from 14 of the 15 townships in the county (Figure 9). Among the 698 specimens, 512 (73%) were collected from southern Lake County, and 186 (27%) were collected from northern Lake County. Important collecting locales included the floodplains of the Pere Marquette (Pleasant Plains, Lake, and Sweetwater townships) and Pine (Dover and Newkirk townships) rivers; the Village of Baldwin (Pleasant Plains and Webber townships); Baldwin-Luther Swamp and vicinity (Cherry Valley Township); “Idlewild Bar

3 This figure includes only those specimens individually checked and confirmed for inclusion on MICHIGAN FLORA ONLINE. Unverified specimens from Lake County exist in several herbaria and are searchable on the Consortium of Midwest Herbaria (2019) website. Some of these may represent taxa we list here as “newly vouchered,†but they are omitted here in concordance with the website’s policy quoted above in footnote 2.

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TABLE 2. Taxonomic summary of species newly vouchered for Lake County. Native Species include those native to Michigan, but not necessarily to Lake County (e.g., Gleditsia triacanthos).

Group Families Genera Native Species Adventive Species Total Species Pteridophytes Gymnosperms Monocots Dicots 6 2 17 73 16 4 73 208 23 4 159 223 0 2 31 117 23 6 190 340 TOTALS 98 301 409 150 559

rens†(Yates Township); and Elbow Lake and Duck Marsh in Elk Township (Figure 9). Eden, Ellsworth, and Sauble townships received the least attention, with one collection from each.

DISCUSSION

Status of the Flora

Prior to our work, only 437 vascular plant species were vouchered from Lake County, ranking 78th among the 83 Michigan counties. With the addition of our recent collections and several independent collections from the same period, a total of 1,006 species have now been vouchered from Lake County, which now ranks 28th (Table 4) (MICHIGAN FLORA ONLINE 2011). This includes 807 native species (28th) and 199 non-native species (30th). Among the 30 counties in

TABLE 3. State-listed species vouchered from Lake County*. E = Endangered; T = Threatened; SC = Special Concern. First collection dates based on specimens examined for MICHIGAN FLORA ONLINE (2011).

Species Common Name State Status First Collection Asclepias ovalifolia dwarf milkweed E 2016 Berula erecta water-parsnip T 2017 Cirsium hillii Hill’s thistle SC 1888 Eleocharis engelmannii Engelmann’s spike-rush SC 2014 Eleocharis melanocarpa black-fruited spike-rush SC 1995 Geum triflorum prairie-smoke T 2007 Helianthus hirsutus hairy sunflower SC 2017 Lipocarpha micrantha dwarf-bulrush SC 1946 Poa paludigena bog bluegrass T 1890 Prunus umbellata Alleghany plum SC 1925 Pycnanthemum verticillatum Whorled mountain mint SC 2018 Rhynchospora macrostachya tall beak-rush SC 2014 Scleria pauciflora few-flowered nut-rush E 2014 Scleria triglomerata tall nut-rush SC 2014 Sisyrinchium strictum blue-eyed-grass SC 2014 Wolffia brasiliensis pointed water meal T 2012

*Brickellia eupatorioides (false boneset, SC) was recently collected from a grassland planting where it was presumably introduced. Boechera missouriensis (Missouri rock cress, SC) is reported by Michigan Natural Features Inventory (MNFI 2018) but not substantiated with specimens.

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FIGURE 9. Collection locations of all 698 collections made in this study. (Basemap: State of Michigan–Michigan GIS Open Data.)

northern Lower Michigan, Lake County ranks sixth (fifth for native species). Based on our observations, HMNF stand data (P. R. McGahn, pers. comm.), and species present in adjacent counties but not yet documented from Lake County, actual vascular plant richness likely exceeds 1,200 species.

We collected more specimens in 2013–2018 than had been collected over the preceding 125 years, increasing the total number of confirmed records for Lake

TABLE 4. Collection statistics for Lake County, derived from MICHIGAN FLORA ONLINE (2011), as of January 2019. Rank: Rank for each category among counties (1 = highest, 83 = lowest). Range: Lowest value (County)–highest value (County).

Category Number Rank Range Vouchered Species 1006 28 346 (Gladwin)–1750 (Washtenaw) Vouchered Native Species% of Total Flora, Native Species 807 80.2% 28 45 278 (Gladwin)–1207 (Kalamazoo) 67.9 (Wayne)–88.8 (Wexford) Vouchered Non-native Species 199 % of Total Flora, Non-native Species 19.8% 30 39 41 (Osceola)–558 (Washtenaw) 11.2 (Wexford)–32.1 (Wayne) Total Collections 1381 51 410 (Gladwin)–13878 (Washtenaw) Avg. No. Collections per Species 1.4 79 1.2 (Gladwin)–9.8 (Keweenaw)

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County to 1,381 and improving its standing among Michigan counties from 79th to 51st (Table 4). Because we targeted new county records, most species are represented by only one collection, which is reflected in the low ratio of 1.4 specimens per species (Table 4). This ratio is below 2.0 for only three other counties with 1,000 or more confirmed species, all of which have been the focus of targeted collecting in recent decades (Fritsch 1993; MICHIGAN FLORA ONLINE 2011; W. S. Martinus and R. G. Schipper, pers. comm.). Additional collecting of species substantiated by only one specimen is recommended to support a more thorough treatment of the Lake County flora in its entirety.

Rare Species

Sixteen naturally occurring state-listed (Endangered, Threatened, and Special Concern) species have now been collected from Lake County (Table 3). Most of these occur in two natural communities—intermittent wetland (seven species) and oak–pine barrens (five species) (Table 3; see also the Species at the Edge of Their Range section below). The remainder occur in cold seeps (two species), beaver ponds (one species), and old fields (one species). Three additional listed species have been reported but are omitted from Table 3. Boechera missouriensis (Missouri rock-cress, SC) is reported from two localities by MNFI (2018) but is not substantiated with specimens. Brickellia eupatorioides (false boneset, SC) was collected from a grassland planting where it was likely introduced. Artemisia ludoviciana (western mugwort, ST) is presumed to be native along the Menominee River in Menominee County but occurs in Lake County (and elsewhere in Michigan) as an introduction (MICHIGAN FLORA ONLINE 2011).

Statewide, rare species richness tends to decline with increasing latitude and distance from the Great Lakes shoreline4 (Figure 10). The southernmost counties are particularly rich in rare species due to proximity to the state line (and hence the presence of edge-of-range taxa that barely enter the state), presence of uncommon habitats, and the scarcity of relatively undisturbed remnants of historically common habitats. Berrien County in the southwestern corner of the state has the richest assemblage of rare species (119), which comprise 10% of its native flora (MICHIGAN FLORA ONLINE 2011). Most counties in interior northern Lower Michigan, in comparison, support fewer than 10 vouchered rare taxa (Figure 10). The somewhat higher figure in Lake County is due to the presence of intermittent wetlands, which otherwise occur primarily to its southwest (see the Species at the Edge of Their Range section below). Among neighboring counties, Newaygo County shares 12 (75%) of the 16 rare species found in Lake County, followed by Mason County, with eight (50%), and Oceana County, with four (25%). This is unsurprising, as contiguous portions of these counties share similar landforms, soils, and vegetation (Albert 1995; Schaetzl et al. 2013; United States Environmental Protection Agency 2013) (Figure 10).

4 In the Upper Peninsula, rare species richness is higher than in northern Lower Michigan due to the presence of an assortment of primarily western, boreal, and arctic-alpine species, especially on Isle Royale (Keweenaw County). Of the 127 species known in Michigan only from the Upper Peninsula, 108 (83%) are state-listed (MNFI 2009; MICHIGAN FLORA ONLINE 2011).

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FIGURE 10. The number of vouchered, naturally occurring state-listed species (endangered, threatened, special concern) per Michigan county, derived from MNFI (2009) and MICHIGAN FLORA ONLINE (2011). (Basemap: State of Michigan–Michigan GIS Open Data.)

Species at the Edge of Their Range

The presence of a vegetation tension zone extending approximately east to west across the center of Michigan’s Lower Peninsula was documented by the beginning of the 20th Century (McCann 1979; Andersen 2005). The tension zone marks a relatively abrupt transition in the dominant vegetation from the hardwood- dominated forests of southern Lower Michigan to the mixed hardwood- conifer forests emblematic of northern Lower Michigan, corresponding to gradients in regional climate, physiography, soils, and hydrology (Albert 1995). The tension zone is typically mapped as extending from Muskegon County northeastward to Bay County and continuing northeastward across Saginaw Bay and the Thumb or angling southeast to Port Huron (McCann 1979; Andersen 2005). Lake County is approximately 40 to 80 km north of the transition zone, and its

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dominant plant communities (e.g., dry northern forest, dry-mesic northern forest) are typical of northern Michigan (Cohen et al. 2014).

In addition to dividing broad vegetation zones, the tension zone corresponds with the northern range limits of a number of southern plant species. Range limits of northern plants, on the other hand, are not concentrated in the same region (McCann 1979). Our findings reflect this pattern. Among the species newly vouchered from Lake County, 27 are at their northern (latitudinal) range limits within the state and only three are at their southern statewide range limits. The county flora in its entirety includes 35 native species at their northern statewide or Lower Peninsula range limits and only six species at their southern range limits (MICHIGAN FLORA ONLINE 2011) (Tables 5–7).

Species at their northern range limits in Lake County cluster in four principal habitats. Eight of the species were collected from shallow, seasonally and inter- annually desiccated kettle depressions known as intermittent wetlands (Tables 5, 6). Several of these species are disjunct from, or have ranges centered on, the Atlantic and Gulf coastal plains and occur in Michigan primarily in the southwestern counties, where their presence typifies a variant of intermittent wetland known as coastal plain marsh (Reznicek 1994; Kost and Penskar 2000; Cohen and Kost 2007; Cohen et al. 2014). A notable assemblage of these species occurs at and near Duck Marsh in Elk Township and is discussed further by Slaughter

TABLE 5. Species at their northernmost statewide range limits in Lake County and the natural communities associated with each, indicated by “xâ€. Natural community classification follows Cohen et al. (2014). BAR = oak–pine or pine barrens; BOG = bog or poor conifer swamp; CS = rich conifer swamp or hardwood–conifer swamp; DMF = dry-mesic northern forest; FF = floodplain forest; IW = intermittent wetland; MF = mesic northern forest; OTH = other.

Species BAR BOG CS DMF FF IW MF OTH Agrimonia pubescens x Aristida purpurascens x Bidens discoidea x Carex laxiculmis var. copulata x Cornus florida x Crataegus calpodendron x Desmodium illinoense x Desmodium rotundifolium x x Dichanthelium commutatum subsp. ashei x Dichanthelium dichotomum x Eleocharis engelmannii x Eleocharis melanocarpa x Enemion biternatum x Lespedeza violacea x x Muhlenbergia schreberi x Pycnanthemum verticillatum x Rhynchospora macrostachya x Rotala ramosior x Samolus parviflorus x Scleria pauciflora x Scleria triglomerata x Wolffia brasiliensis x x

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TABLE 6. Species at their northernmost Lower Peninsula range limits in Lake County and the natural communities associated with each, indicated by “xâ€. Natural community classification follows Cohen et al. (2014). BAR = oak–pine or pine barrens; BOG = bog or poor conifer swamp; CS = rich conifer swamp or hardwood–conifer swamp; DMF = dry-mesic northern forest; FF = floodplain forest; IW = intermittent wetland; MF = mesic northern forest; OTH = other.

Species BAR BOG CS DMF FF IW MF OTH

Acalypha rhomboidea xx Allium canadense x Cyperus strigosus x Dichanthelium clandestinum x Dichanthelium praecocius x Geum triflorum x Helianthus hirsutus x Leersia virginica x Micranthes pensylvanica x Muhlenbergia frondosa xx Podophyllum peltatum x Ranunculus fascicularis x Symphyotrichum oolentangiense xx

TABLE 7. Species at their southernmost statewide range limits in Lake County and the natural communities associated with each, indicated by “xâ€. Natural community classification follows Cohen et al. (2014). BAR = oak–pine or pine barrens; BOG = bog or poor conifer swamp; CS = rich conifer swamp or hardwood–conifer swamp; DMF = dry-mesic northern forest; FF = floodplain forest; IW = intermittent wetland; MF = mesic northern forest; OTH = other.

Species BAR BOG CS DMF FF IW MF OTH

Arceuthobium pusillum x Asclepias ovalifolia x Carex adusta x Oxalis acetosella x Sibbaldiopsis tridentata x Sorbus americana x

(2016) (an example of these species is shown in Figure 11). A larger cluster of intermittent wetlands in the vicinity of Big Star Lake in Lake Township merits additional exploration.

Riparian forests were lucrative collecting grounds, supporting both northern and southern floristic elements. Among the latter are eight species at their northern range limits, all collected from the wooded floodplain of the Pere Marquette River (Tables 5, 6). The extra-latitudinal presence of “southern†plant species in riparian habitats is well-documented and is thought to be related to microclimatic amelioration (Tepley et al. 2004; MICHIGAN FLORA ONLINE 2011). However, empirical evidence for the presence of distinct riparian microclimates in eastern North America is scarce (Brooks and Kyker-Snowman 2009), and other variables such as soil characteristics and post-glacial dispersal may be responsible for the presence of these range-limited species. In general, species richness within Michigan floodplain forests declines from south to north (Tepley

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FIGURE 11. Rhynchospora macrostachya (tall beak-rush), which is state-listed as a Special Concern species, is a locally frequent sedge of intermittent wetlands occurring at its northern statewide range limit in northwestern Lake County and northeastern Mason County. Photo by B. S. Slaughter, Van Buren Co., Michigan.

et al. 2004) but attempts to make inferences at finer scales are hampered by an absence of data.

Among upland habitats, species at their northern range limits occurred primarily in dry-mesic northern forest and oak–pine barrens. Noteworthy are several legumes, including Desmodium illinoense (prairie tick-trefoil), D. rotundifolium (round-leaved tick-trefoil), and Lespedeza violacea (bush-clover). These and several other species prevalent in southern Michigan prairies and savannas, such as Lespedeza hirta (hairy bush-clover), Lupinus perennis (wild lupine), and Ranunculus fascicularis (early buttercup), occur locally within Lake County (mostly in the southern townships) but are conspicuously absent from the jack pine plains of north-central and northeastern Lower Michigan5, perhaps due to inhospitable climate, dispersal limitations, soil characteristics, absence of nitrogen- fixing symbionts, or other factors (Bordeleau and Prévost 1994; Comer 1996; Chapman and Brewer 2008; MICHIGAN FLORA ONLINE 2011; Simon- sen et al. 2017).

Although only three species at their northern range limits were collected from mesic northern forest (Table 6), several species known to occur north of Lake

5 Of the jack pine plains flora of northern Lower Michigan, W. J. Beal remarked that the absence of Fabaceae was “most remarkable of all†(Michigan State Board of Agriculture 1888).

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FIGURE 12. The showy biennial Hydrophyllum appendiculatum (great waterleaf) occurs very locally in rich forests in the southeastern part of the county. Photo by B. S. Slaughter, Cass Co., Michigan.

County only in lakeshore counties or along rivers were collected, including Hydrophyllum appendiculatum (great waterleaf), H. canadense (Canada waterleaf) (Figure 12), Persicaria virginiana (jumpseed), and Symphyotrichum cordifolium (heart-leaved aster) (MICHIGAN FLORA ONLINE 2011). These species were concentrated in Chase Township in the southeastern corner of the county, where a lobe of nutrient-rich, loamy glacial till supports a few relatively intact woodlots dominated by Acer saccharum (sugar maple), Fagus grandifolia (American beech), and Tilia americana (basswood). Such forests occur with greater prevalence just east and north of Lake County (Burger and Kotar 2003; USDA, NRCS 2015). Because they are agriculturally productive, these lands are primarily in private ownership and are relatively inaccessible to botanists. Exploration of mesic woodlots in northwestern Lower Michigan is essential for the demarcation of range limits of many principally southern forest herbs.

Species at their southern statewide range limits are few and apparently localized (Table 7). The most remarkable find was a small disjunct population of Asclepias ovalifolia (dwarf milkweed) (Figure 13) discovered by HMNF surveyors in Yates Township (D. Gaebel and M. Yageman 1, MICH). This species was previously known in Michigan only from Menominee County in the Upper Peninsula, but it occurs as far south as southern Iowa in its core Great Plains range (Kartesz 2015). The principally northern Arceuthobium pusillum (dwarf mistletoe) and northern (and/or high-elevation) Oxalis acetosella (northern wood-sorrel) both occur locally in cold conifer-dominated swamps in the

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FIGURE 13. The state-endangered Asclepias ovalifolia (dwarf milkweed), thought to be restricted to Menominee County in the Upper Peninsula until its recent discovery in Yates Township in Lake County. Photo by B. S. Slaughter, Menominee Co., Michigan.

northern townships (Table 7). Several other species at or near their extant southern range limits were encountered on cool riverbanks and wooded slopes adjacent to swamps. Among these were Cynoglossum boreale (northern wild comfrey), Equisetum pratense (meadow horsetail), Lonicera hirsuta (hairy honeysuckle), and Petasites frigidus (sweet-coltsfoot). The northern conifer Abies balsamea (balsam fir) appears to be scarce outside of a small area where Lake, Osceola, and Wexford counties adjoin6. Further surveys are needed to clarify the abundance and distribution of these and other principally northern species within the county.

Non-native and Invasive Species

When we initiated our study in 2013, approximately 10% of the vascular flora vouchered from Lake County was comprised of non-native species, ranking lowest among Michigan counties in that respect (MICHIGAN FLORA ONLINE

6 Abies balsamea is almost entirely restricted to northern Michigan, with one outlying collection from a bog in Ingham County (MICHIGAN FLORA ONLINE 2011). Just east and north of Lake County, A. balsamea and Picea mariana (black spruce), the latter nearly restricted to acidic bogs in Lake County, mingle with Thuja occidentalis (northern white-cedar) over extensive tracts. These cold, dark, moss-carpeted swamps beckon and charm seekers of botanical treasures but are unfortunately themselves elusive in Lake County, presumably due to the relatively warm climate and perhaps human disturbance.

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2011; Voss and Reznicek 2012). Our collections markedly closed that collection gap, increasing the relative proportion of non-native flora to 20% (Table 4)7. A few non-native invasive species are particularly abundant within Lake County on dry, open or semi-shaded sandy soils. From a stewardship perspective, the most troublesome examples include Centaurea stoebe (spotted knapweed), Elaeagnus umbellata (autumn-olive), Euphorbia virgata (leafy spurge), Lonicera ï‚´bella (hybrid honeysuckle), L. morrowii (Morrow honeysuckle), Melilotus albus (white sweet-clover), M. officinalis (yellow sweet-clover), and Securigera varia (crown-vetch). Species typical of cultivated ground or urban areas, such as Abutilon theophrasti (velvet-leaf), Brassica rapa (field mustard), Datura stramonium (jimson-weed), Draba verna (whitlow-grass), Eragrostis cilianensis (stink grass), Lamium purpureum (purple dead-nettle), and Thlaspi arvense (penny cress) occurred more locally, usually on heavier soils (e.g., in Chase Township) and in waste areas and near structures, as at Baldwin.

Many of Michigan’s most destructive invasive plants are uncommon within Lake County. Examples among our collections include Celastrus orbiculatus (Oriental bittersweet), Phragmites australis subsp. australis (reed), Rhamnus cathartica (common buckthorn), Rosa multiflora (multiflora rose), and Typha angustifolia (narrow-leaved cat-tail). We observed only a few localized populations of Alliaria petiolata (garlic mustard), but dozens of stations are reported on the Midwest Invasive Species Information Network (MISIN 2019) and Early Detection & Distribution Mapping System (EDDMapS 2019). Frangula alnus (glossy buckthorn) was not observed and has not been reported from Lake County as of early 2019 on either of these websites, but its presence in adjacent counties suggests its furtive residence.

CONCLUSIONS

Prior to the unveiling of MICHIGAN FLORA ONLINE (2011), compiling county-level species lists and identifying unvouchered targets involved manual examination of dated county maps in Michigan Flora (Voss 1972, 1985, 1996). Now, botanists have access to regularly updated species treatments, maps, and specimen data that greatly enhance our ability to address collection gaps. While our examination of the Lake County flora is incomplete, we were able to add hundreds of new species records that considerably improve our understanding of its vegetation and flora. Additional exploration and collecting focused on new records and resolution of species ranges and abundances within the county is recommended, as is coordination among herbaria to ensure that existing but unverified specimens are examined for inclusion in MICHIGAN FLORA ON

7 For perspective, non-native species comprise approximately 37.5% of the Michigan Flora (MICHIGAN FLORA ONLINE 2011). At the county level, the proportion of non-native species ranges from 11% (Wexford County) to 32% (Washtenaw and Wayne counties), indicative of both collection bias and the large number of non-native species represented by one or few populations, these often transient.

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LINE. Trowels, field presses, and their human assistants should also be directed to other poorly-collected counties, especially the contiguous three-county area of Osceola, Missaukee, and Wexford. The paucity of specimens from interior northern Lower Michigan is, reciprocally, an opportunity for discovery.

ACKNOWLEDGMENTS

We appreciate the support of our former and current colleagues at MNFI (Michigan State University Extension), HMNF, Orbis Environmental Consulting, and MICH. We thank Erin Victory, Cheryl Nelson, Mike Lesinski, and Gary Mees (MDNR, Baldwin Field Office) for site leads on state land, and Pat Ruta McGhan (USFS, Baldwin/White Cloud Ranger District, HMNF) for providing substantial assistance in the form of habitat descriptions, forest stand data, locations of individual species, and other site leads. Several MICH staff provided critical assistance; we thank Tony Reznicek for examining and verifying (or redetermining) identifications, Bev Walters and assistants for specimen preparation, data entry, and map updates, and Rich Rabeler for data entry and assistance locating specimens. We also thank Greg Schmidt (USDA, NRCS), Trevor Hobbs (Contour Geographic), and Kevin Kincare (USGS) for providing insight and guidance on the abiotic and biotic context of the flora. Last, we thank Michael Huft, Mike Penskar, and one anonymous reviewer for suggesting edits that improved the manuscript.

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APPENDIX 1. CHECKLIST OF VASCULAR PLANTS NEWLY CONFIRMED FOR LAKE COUNTY

The following checklist is arranged phylogenetically by major clade or group and then alphabetically by family, genus and species. Classes, subclasses, orders, and family circumscriptions for pteridophytes follow PPG I (2016); conifer subclass and family circumscriptions follow Chase and Reveal (2009) and MICHIGAN FLORA ONLINE (2011), respectively; and major angiosperm clades and grades and family circumscriptions follow APG IV (2016). Families treated in MICHIGAN FLORA ONLINE (2011) that are not treated in APG IV are included and cross-referenced in the checklist. Genera and species follow MICHIGAN FLORA ONLINE (2011). This checklist is limited to species that are newly confirmed for Lake County and substantiated with our specimens. The full list of vouchered taxa known from the county is searchable on MICHIGAN FLORA ONLINE (2011).

Each checklist entry includes the scientific name in italics, followed by the authority for the name, the common name, and, in parentheses, collector initial (K =Amanda K. Klain; S = Bradford

S. Slaughter), and collection number. Introduced species are indicated with an asterisk (*) preceding the scientific name. Species lacking an asterisk are considered native to Michigan (MICHIGAN FLORAONLINE 2011) but are not necessarily native to Lake County (e.g., Gymnocladus dioicus). Species listed as Endangered, Threatened, or Special Concern in Michigan (following MNFI 2009) are denoted by the listing category in BOLD CAPITALS following the collection numbers. All collections were deposited at The University of Michigan Herbarium (MICH). Location, population, and habitat data are not provided in the checklist but are searchable (or will become searchable with future updates) on MICHIGAN FLORAONLINE (2011). LYCOPODIALES (Lycopods)

LYCOPODIACEAE Dendrolycopodium hickeyi (W. H. Wagner, Beitel & R. C. Moran) A. Haines (S2005) Diphasiastrum digitatum (A. Braun) Holub, ground-cedar (S1334)

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Lycopodium clavatum L., running ground-pine (S1336) Spinulum annotinum (L.) A. Haines, stiff clubmoss (S1644)

EQUISETIDAE (Horsetails)

EQUISETACEAE Equisetum arvense L., common horsetail (K177, S1263) Equisetum fluviatile L., water horsetail (S1793) Equisetum laevigatum A. Braun, smooth scouring rush (S1654) Equisetum pratense Ehrh., meadow horsetail (S1260) Equisetum scirpoides Michx., dwarf scouring rush (S1400) Equisetum sylvaticum L., woodland horsetail (S1127)

POLYPODIIDAE (Leptosporangiate Ferns)

ASPLENIACEAE Asplenium platyneuron (L.) D. C. Eaton, ebony spleenwort (S1621)

ATHYRIACEAE Athyrium filix-femina (L.) Roth, lady fern (S1495) Deparia acrostichoides (Michx.) Desv., silvery spleenwort (S1953)

CYSTOPTERIDACEAE Cystopteris bulbifera (L.) Bernh., bulblet fern (S1254) Gymnocarpium dryopteris (L.) Newm., oak fern (S1241)

DRYOPTERIDACEAE Dryopteris cristata (L.) A. Gray, crested shield fern (S1463) Polystichum acrostichoides (Michx.) Schott., Christmas fern (S1200)

ONOCLEACEAE Matteuccia struthiopteris (L.) Todaro, ostrich fern (S1415) Onoclea sensibilis L., sensitive fern (S1267)

OSMUNDACEAE Osmunda claytoniana L., interrupted fern (S1424) Osmunda regalis L., royal fern (S1476)

THELYPTERIDACEAE Thelypteris noveboracensis (L.) Nieuwl., New York fern (S1692) Thelypteris palustris Schott, marsh fern (S1690)

PINIDAE (Conifers)

CUPRESSACEAE Juniperus communis L., common juniper (S1935) Juniperus virginiana L., red-cedar (K404)

PINACEAE Abies balsamea (L.) Mill., balsam fir (S1797) *Picea abies (L.) H. Karst., Norway spruce (S2284)

Pinus strobus L., white pine (K218)

*Pinus sylvestris L., Scots pine (S1623)

BASAL ANGIOSPERMS

CABOMBACEAE Brasenia schreberi J. F. Gmel., water-shield (S1706)

MAGNOLIIDAE (Magnoliids)

ARISTOLOCHIACEAE Asarum canadense L., wild-ginger (S1399)

LAURACEAE Lindera benzoin (L.) Blume, spicebush (S1800)

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MONOCOTS

ALISMATACEAE Alisma triviale Pursh, northern water-plantain (S1911, 1961) Sagittaria latifolia Willd., wapato (S1729)

AMARYLLIDACEAE Allium canadense L., wild garlic (S1138, 1192) *Narcissus pseudonarcissus L., daffodil (S2004)

ARACEAE Calla palustris L., wild calla (S1489) Lemna minor L., common duckweed (S1901) Lemna turionifera Landolt, red duckweed (S1642) Symplocarpus foetidus (L.) Nutt., skunk-cabbage (S1384) Wolffia columbiana H. Karst., common water meal (S2286)

ASPARAGACEAE *Asparagus officinalis L., garden asparagus (S1490) *Convallaria majalis L., lily-of-the-valley (S1790)

Maianthemum canadense Desf., Canada mayflower (S1142) Maianthemum racemosum (L.) Link, false spikenard (S1166, 1266) Maianthemum stellatum (L.) Link, starry false Solomon-seal (S1137) Maianthemum trifolium (L.) Sloboda, false mayflower (S2285)

*Muscari neglectum Tenore, grape-hyacinth (S1386, 1769) Uvularia grandiflora Sm., bellwort (S1261)

ASPHODELACEAE *Hemerocallis fulva (L.) L., orange day-lily (S1876) *Hemerocallis lilio-asphodelus L., yellow day-lily (S1820)

CONVALLARIACEAE—SEE ASPARAGACEAE

CYPERACEAE

Carex adusta Boott, sedge (S1666) Carex albicans Spreng. var. emmonsii (Torr.) Rettig, sedge (S1633) Carex albursina E. Sheld., sedge (S1613) Carex alopecoidea Tuck., sedge (S2235) Carex arctata Boott, sedge (S1410) Carex argyrantha Tuck., sedge (S1997) Carex aurea Nutt., sedge (S1242) Carex blanda Dewey, sedge (S2210) Carex brevior (Dewey) Mack., sedge (S1867) Carex bromoides Willd., sedge (S1194) Carex brunnescens (Pers.) Poir., sedge (S1643) Carex buxbaumii Wahlenb., sedge (S1669) Carex canescens L., sedge (S1635) Carex castanea Wahlenb., sedge (S1630, 1814) Carex cephaloidea (Dewey) Dewey, sedge (S1431) Carex comosa Boott, sedge (S1686) Carex crinita Lam., sedge (S1190, 1634) Carex cristatella Britton, sedge (S1196) Carex debilis Michx. var. rudgei L. H. Bailey, swamp sedge (S1631) Carex deweyana Schwein., sedge (S1201) Carex diandra Schrank, sedge (S1652) Carex disperma Dewey, sedge (S1234) Carex flava L., sedge (S1236) Carex gracillima Schwein., sedge (S1188) Carex granularis Willd., sedge (S1649) Carex grisea Wahlenb., sedge (S1191) Carex hirtifolia Mack., sedge (S1763)

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Carex hitchcockiana Dewey, sedge (S1612) Carex hystericina Willd., sedge (S1233) Carex interior L. H. Bailey, sedge (S1129) Carex lacustris Willd., sedge (S1601) Carex laevivaginata (Kük.) Mack., sedge (S1817) Carex laxiculmis Schwein. var. copulata (L. H. Bailey) Fernald, sedge (S1202) Carex laxiflora Lam., sedge (S1404) Carex leptalea Wahlenb., sedge (S1235) Carex leptonervia (Fernald) Fernald, sedge (S1620) Carex lucorum Link, sedge (S1834) Carex lurida Wahlenb., sedge (S2290) Carex pallescens L., pale sedge (S1826) Carex peckii Howe, sedge (S1414) Carex pedunculata Willd., sedge (S1398) Carex pensylvanica Lam., sedge (S1099, 1104, 1629) Carex plantaginea Lam., sedge (S1427) Carex prairea Dewey, sedge (S1239) Carex prasina Wahlenb., sedge (S1794) Carex projecta Mack., sedge (S1657, 1661, 1700, 1887) Carex pseudocyperus L., sedge (S1482) Carex radiata (Wahlenb.) Small, straight-styled wood sedge (S1193, 1798) Carex retrorsa Schwein., sedge (S1659) Carex scabrata Schwein., sedge (S1244) Carex schweinitzii Schwein., sedge (S1815) Carex scoparia Willd., sedge (S1670, 1672) Carex siccata Dewey, sedge (S1591) Carex sparganioides Willd., sedge (S1199) Carex sprengelii Spreng., sedge (S1402) Carex stipata Willd., sedge (S1198) Carex stricta Lam., sedge (S1268, 1606) Carex tenera Dewey, sedge (S1671, 1835) Carex tuckermanii Dewey, sedge (S1253) Carex utriculata Boott, sedge (S1811) Carex woodii Dewey, sedge (S1796) Cladium mariscoides (Muhl.) Torr., twig-rush (S1226) Cyperus schweinitzii Torr., rough sand sedge (K402, S1923) Cyperus strigosus L., long scaled nut sedge (S1956) Eleocharis elliptica Kunth, golden-seeded spike-rush (S1339) Eleocharis engelmannii Steud., Engelmann’s spike-rush (S1382). SPECIALCONCERN Eleocharis erythropoda Steud., spike-rush (S1728) Eleocharis intermedia Schult., spike-rush (S2549) Eleocharis obtusa (Willd.) Schult., spike-rush (S1885) Eleocharis robbinsii Oakes, spike-rush (S1931) Eleocharis rostellata Torr., beaked spike-rush (S1331) Eriophorum gracile W. D. J. Koch, slender cotton-grass (S1603) Rhynchospora fusca (L.) W. T. Aiton, beak-rush (S1608) Rhynchospora macrostachya A. Gray, tall beak-rush (S1315). SPECIALCONCERN Schoenoplectus pungens (Vahl) Palla, three-square (S1328) Scirpus atrovirens Willd., bulrush (S1487) Scirpus pendulus Muhl., bulrush (S1645) Scleria pauciflora Willd. var. pauciflora, few-flowered nut-rush (S1312). ENDANGERED Scleria triglomerata Michx., tall nut-rush (S1311). SPECIALCONCERN

DIOSCOREACEAE Dioscorea villosa L., wild yam (S1262)

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HYACINTHACEAE—SEE ASPARAGACEAE

HYDROCHARITACEAE Elodea canadensis Michx., common waterweed (S1387)

IRIDACEAE

*Iris germanica L., flag (S1627) Iris versicolor L., wild blue flag (S1650) Sisyrinchium montanum Greene var. crebrum Fernald, mountain blue-eyed-grass (S1830) Sisyrinchium mucronatum Michx., slender blue-eyed-grass (S1647) Sisyrinchium strictum E. P. Bicknell, blue-eyed-grass (S1154, 1223, 1596). SPECIAL

CONCERN

JUNCACEAE Juncus articulatus L., jointed rush (S1926) Juncus balticus Willd. subsp. littoralis (Engelm.) Snogerup., Baltic rush (S1222) Juncus bufonius L. var. bufonius, toad rush (S1914) Juncus nodosus L., knotted rush (S1240) Juncus pylaei Laharpe, Pylaei’s soft rush (S1460, 1730, 1869) Juncus tenuis Willd., path rush (K142) Juncus torreyi Coville, Torrey’s rush (S1973) Luzula multiflora (Ehrh.) Lej., common wood rush (S1825)

LILIACEAE Lilium michiganense Farw., Michigan lily (S1256) Medeola virginiana L., Indian cucumber-root (S1632)

MELANTHIACEAE Aletris farinosa L., colic root (S2403) Trillium cernuum L., nodding trillium (S1405)

ORCHIDACEAE Arethusa bulbosa L., dragon’s mouth (S1605) Corallorhiza odontorhiza (Willd.) Nutt., fall coral-root (S1337) Cypripedium reginae Walter, showy lady-slipper (S1187)

*Epipactis helleborine (L.) Crantz, helleborine (S1272) Liparis loeselii (L.) Rich., Loesel’s twayblade (S1898)

POACEAE

*Agrostis gigantea Roth, redtop (S1473) Agrostis perennans (Walter) Tuck., autumn bent (S1977) Agrostis scabra Willd., ticklegrass (S1045) Alopecurus aequalis Sobol., short-awned foxtail (S1641) Andropogon virginicus L., broom-sedge (S2001)

*Anthoxanthum odoratum L., sweet vernal grass (S1818) Aristida basiramea Vasey, fork-tipped three-awned grass (S1044) Aristida purpurascens Poir., three-awned grass (S2000)

*Bromus japonicus Murray, Japanese brome (K158, 391; S1247) Bromus latiglumis (Shear) Hitchc., ear-leaved brome (S1494) Bromus nottowayanus Fernald, satin brome (S1465)

*Bromus squarrosus L., brome (S1889)

*Bromus tectorum L., soft chess (S1805) Cinna arundinacea L., wood reedgrass (S1732) Cinna latifolia (Goepp.) Griseb., wood reedgrass (S1504)

*Dactylis glomerata L., orchard grass (S1445) Dichanthelium clandestinum (L.) Gould, panic grass (S2240) Dichanthelium commutatum (Schult.) Gould subsp. ashei (Ashe) Fernald, panic grass (S1930) Dichanthelium dichotomum (L.) Gould, panic grass (S1938) Dichanthelium spretum (Schult.) Freckmann, Eaton’s rosette grass (S1316) Digitaria cognata (Schult.) Pilg., fall witch grass (S1975)

*Digitaria ischaemum (Schreb.) Muhl., smooth crab grass (S1734)

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Echinochloa muricata (P. Beauv.) Fernald var. microstachya Wiegand, barnyard grass (S1989)

*Elymus repens (L.) Gould, quack grass (S1656) Elymus riparius Wiegand, riverbank wild-rye (S1496) Elymus virginicus L., Virginia wild-rye (S1493)

*Eragrostis cilianensis (All.) Janch., stink grass (S1992) Eragrostis hypnoides (Lam.) Britton, Sterns & Poggenb., creeping love grass (S2548)

*Eragrostis minor Host, low love grass (S1846) Eragrostis pectinacea (Michx.) Nees, love grass (S1048) Eragrostis spectabilis (Pursh) Steud., purple love grass (S1047)

*Festuca rubra L., red fescue (S2236) Festuca subverticillata (Pers.) E. B. Alexeev, nodding fescue (S1702)

*Festuca trachyphylla (Hack.) Krajina, sheep fescue (S1446) Glyceria grandis S. Watson, reed manna grass (S1471) Glyceria septentrionalis Hitchc., floating manna grass (S1884) Glyceria striata (Lam.) Hitchc., fowl manna grass (S1189) Leersia virginica Willd., white grass (S1736)

*Lolium perenne L., ryegrass (S1688) Muhlenbergia frondosa (Poir.) Fernald, common satin grass (S2546) Muhlenbergia glomerata (Willd.) Trin., marsh wild timothy (S1043, 1330) Muhlenbergia schreberi J. F. Gmel., nimblewill (S1947) Muhlenbergia sylvatica Torr., woodland satin grass (S2547) Oryzopsis asperifolia Michx., rough-leaved rice-grass (S1388) Panicum dichotomiflorum Michx., panic grass (S2545) Panicum virgatum L., switch grass (S1737)

*Phleum pratense L., timothy (S1655) Phragmites australis (Cav.) Steud. subsp. americanus Saltonst., P. M. Peterson & Soreng, reed (S1969) *Phragmites australis (Cav.) Steud. subsp. australis, reed (S1951)

Poa alsodes A. Gray, grove bluegrass (S1144, 1637, 1810) *Poa annua L., annual bluegrass (S1107) *Poa bulbosa L., bluegrass (S1773) *Poa nemoralis L., bluegrass (S1703)

Poa saltuensis Fernald & Wiegand, bluegrass (S1799, 1809) *Schedonorus arundinaceus (Schreb.) Dumort., tall fescue (K141) *Secale cereale L., rye (S1648) *Setaria faberi Herrm., giant foxtail (S1979) *Sorghum bicolor (L.) Moench var. bicolor, sorghum (S1988)

Sphenopholis intermedia (Rydb.) Rydb., slender wedgegrass (S1701) Torreyochloa fernaldii (Hitchc.) G. L. Church, Fernald’s false mannagrass (S2289) Tridens flavus (L.) Hitchc., purpletop (S2544)

POTAMOGETONACEAE Potamogeton berchtoldii Fieber, pondweed (S2288) Stuckenia pectinata (L.) Börner, sago pondweed (K400)

SMILACACEAE Smilax ecirrata (Kunth) S. Watson, upright carrion-flower (S1255)

TRILLIACEAE—SEE MELANTHIACEAE

TYPHACEAE Sparganium americanum Nutt., American bur-reed (S1726) Sparganium natans L., small bur-reed (S1948)

*Typha angustifolia L., narrow-leaved cat-tail (S1329) Typha latifolia L., common cat-tail (S1481)

EUDICOTS

ADOXACEAE Sambucus racemosa L., red-berried elder (S1764)

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Viburnum lentago L., nannyberry (S1425) *Viburnum opulus L., European highbush-cranberry (S1829) Viburnum trilobum Marshall, American highbush-cranberry (K193)

AMARANTHACEAE

Amaranthus albus L., tumbleweed (S1968) *Amaranthus powellii S. Watson, tall amaranth (S1990) *Amaranthus retroflexus L., rough amaranth (S1942) *Chenopodium album L., lambs-quarters (S1374, 1919) *Chenopodium glaucum L., oak-leaved goosefoot (S1943)

Chenopodium simplex Raf., maple-leaved goosefoot (S1983)

ANACARDIACEAE *Cotinus coggygria Scop., smoke-tree (S1710) Rhus typhina L., staghorn sumac (K216)

APIACEAE Berula erecta (Huds.) Coville, water-parsnip (S1902). THREATENED Cicuta bulbifera L., water hemlock (S1322) Cicuta maculata L., water hemlock (S1501) Cryptotaenia canadensis (L.) DC., honewort (S1693) Heracleum maximum Bartram, cow-parsnip (S1436) Osmorhiza claytonii (Michx.) C. B. Clarke, hairy sweet-cicely (S1689)

*Pastinaca sativa L., wild parsnip (S1877) Sium suave Walter, water-parsnip (S1883) *Torilis japonica (Houtt.) DC., hedge-parsley (K195, S1270) Toxicodendron radicans (L.) Kuntze, poison-ivy (S1910)

APOCYNACEAE Apocynum cannabinum L. var. cannabinum, Indian-hemp (S1904) *Vinca minor L., periwinkle (S1767)

ARALIACEAE Aralia hispida Vent., bristly sarsaparilla (S1665) Aralia nudicaulis L., wild sarsaparilla (S1433) Hydrocotyle americana L., water-pennywort (K191)

ASTERACEAE Achillea millefolium L., yarrow (S1440) Ageratina altissima (L.) R. M. King & H. Rob., white snakeroot (S1502) Antennaria parlinii Fernald, smooth pussytoes (S1103)

*Anthemis arvensis L., corn chamomile (S1106) *Arctium minus Bernh., common burdock (S1880) *Artemisia ludoviciana Nutt., western mugwort (S1976)

Bidens discoidea (Torr. & A. Gray) Britton, swamp beggar-ticks (S1998) Bidens frondosa L., common beggar-ticks (K214) Bidens vulgata Greene, tall beggar-ticks (S1959)

*Cirsium arvense (L.) Scop., Canada thistle (S1681) Cirsium discolor (Willd.) Spreng., pasture thistle (S1915) *Cirsium vulgare (Savi) Ten., bull thistle (S1722) Coreopsis tripteris L., tall coreopsis (S1908)

*Crepis tectorum L., hawk’s beard (S1921) Doellingeria umbellata (Mill.) Nees, flat-topped white aster (S1327) Erechtites hieraciifolius (L.) Raf., fireweed (S1318) Erigeron annuus (L.) Pers., daisy fleabane (S1865, 1936) Erigeron philadelphicus L., common fleabane (S1141) Erigeron pulchellus Michx., robin’s-plantain (S1134) Erigeron strigosus Willd., daisy fleabane (S1049) Helianthus giganteus L., tall sunflower (S2551) Helianthus hirsutus Raf., hairy sunflower (S1894). SPECIALCONCERN.

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*Hieracium caespitosum Dumort., king devil (S1592) Petasites frigidus (Aiton) A. Gray var. palmatus (Aiton) Cronquist, sweet-coltsfoot (S1246) Prenanthes alba L., white lettuce (S1505) Rudbeckia laciniata L., tall coneflower (S1723) Solidago altissima L., tall goldenrod (S1751) Solidago caesia L., bluestem goldenrod (S1340) Solidago flexicaulis L., zigzag goldenrod (S1982) Solidago gigantea Aiton, late goldenrod (S1324) Solidago uliginosa Nutt., bog goldenrod (S1326)

*Sonchus arvensis L., field sow-thistle (S1899) Symphyotrichum cordifolium (L.) G. L. Nesom, heart-leaved aster (S1960) Symphyotrichum novae-angliae (L.) G. L. Nesom, New England aster (S1972) Symphyotrichum ontarionis (Wiegand) G. L. Nesom, Lake Ontario aster (S1981) Symphyotrichum pilosum (Willd.) G. L. Nesom var. pilosum, frost aster (S1980) Symphyotrichum puniceum (L.) Ã. Löve & D. Löve, swamp aster (S1727)

*Tanacetum vulgare L., common tansy (S1907) *Taraxacum officinale F. H. Wigg., common dandelion (S1385) *Tragopogon dubius Scop., goat’s beard (S1600) *Tragopogon pratensis L., common goat’s beard (S1599)

BERBERIDACEAE *Berberis thunbergii DC., Japanese barberry (S2006) *Berberis vulgaris L., common barberry (S1813)

Podophyllum peltatum L., may-apple (S1426)

BETULACEAE Betula alleghaniensis Britton, yellow birch (S1795) Carpinus caroliniana Walter, blue-beech (K129) Corylus americana Walter, hazelnut (S1638) Corylus cornuta Marshall, beaked hazelnut (S2209) Ostrya virginiana (Mill.) K. Koch, ironwood (S1705)

BIGNONIACEAE *Catalpa speciosa Warder, northern catalpa (S2397)

BORAGINACEAE Cynoglossum boreale Fernald, northern wild comfrey (S1245) Hackelia virginiana (L.) I. M. Johnst., beggar’s lice (S1879) Hydrophyllum appendiculatum Michx., great waterleaf (S1614) Hydrophyllum canadense L., Canada waterleaf (S1844) Hydrophyllum virginianum L., Virginia waterleaf (S1803) Lithospermum caroliniense (Walter) MacMill., hairy puccoon (S1167)

*Myosotis stricta Link, small-flowered forget-me-not (S1772) *Myosotis sylvatica Hoffm., garden forget-me-not (S1416)

BRASSICACEAE *Alliaria petiolata (M. Bieb.) Cavara & Grande, garlic mustard (S1408) *Alyssum alyssoides (L.) L., pale alyssum (S1112) *Barbarea vulgaris R. Br., yellow rocket (S1131) *Brassica rapa L., field mustard (S1788)

Cardamine bulbosa (Muhl.) Britton, Sterns & Poggenb., spring cress (S1139) Cardamine concatenata (Michx.) O. Schwarz, cut-leaved toothwort (S1389) Cardamine diphylla (Michx.) Alph. Wood, two-leaved toothwort (S1403)

*Cardamine flexuosa With., bitter cress (S1849)

Cardamine pensylvanica Willd., Pennsylvania bitter cress (S1128) *Draba verna L., whitlow-grass (S2003) *Hesperis matronalis L., dame’s rocket (S1432) *Lunaria annua L., money-plant (S2007) *Nasturtium microphyllum Rchb., watercress (S2291)

Nasturtium officinale W. T. Aiton, watercress (S1238)

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*Sisymbrium officinale L., hedge mustard (S2400) *Thlaspi arvense L., penny cress (S1848)

CACTACEAE Opuntia cespitosa Raf., prickly-pear (S2399)

CAMPANULACEAE Lobelia inflata L., indian-tobacco (S1733) Triodanis perfoliata (L.) Nieuwl., Venus’s looking-glass (S1847)

CANNABACEAE Humulus lupulus L., common hops (S1971)

CAPRIFOLIACEAE

Diervilla lonicera Mill., bush-honeysuckle (S1843) *Dipsacus fullonum L., wild teasel (S1891) *Lonicera ï‚´bella Zabel, hybrid honeysuckle (S1126)

Lonicera canadensis Marshall, Canadian fly honeysuckle (S1417)

Lonicera hirsuta Eaton, hairy honeysuckle (S1437) *Lonicera morrowii A. Gray, Morrow honeysuckle (S1801) *Lonicera tatarica L., Tartarian honeysuckle (S1893)

Symphoricarpos albus (L.) S. F. Blake var. albus, snowberry (S1466)

CARYOPHYLLACEAE *Agrostemma githago L., corn-cockle (S1252) *Arenaria serpyllifolia L., thyme-leaved sandwort (K143, S1109)

Cerastium arvense L., field chickweed (K133, S1096, S1770) *Cerastium semidecandrum L., small mouse-ear chickweed (S1108) *Dianthus armeria L., Deptford pink (S1685) *Herniaria glabra L., herniary (S1866) *Myosoton aquaticum (L.) Moench, giant chickweed (S2237) *Petrorhagia saxifraga (L.) Link, pink (S1488) *Scleranthus annuus L., knawel (S1110) *Silene latifolia Poir., white campion (S1441)

Stellaria longifolia Willd., long-leaved chickweed (S1143) *Stellaria media (L.) Vill., common chickweed (S1265)

CELASTRACEAE *Celastrus orbiculatus Thunb., oriental bittersweet (S2396) Celastrus scandens L., climbing bittersweet (S1892)

CISTACEAE Lechea intermedia Britton, intermediate pinweed (S1313)

CLEOMACEAE Polanisia dodecandra (L.) DC. var. dodecandra, clammy-weed (K395, S1918)

CONVOLVULACEAE Calystegia sepium (L.) R. Br., hedge bindweed (S2239) Calystegia spithamea (L.) Pursh, low bindweed (S1662)

*Convolvulus arvensis L., field bindweed (S1598)

CORNACEAE Cornus alternifolia L. f., alternate-leaved dogwood (S1694) Cornus amomum Mill. subsp. obliqua (Raf.) J. S. Wilson, silky dogwood (S1871) Cornus foemina Mill., gray dogwood (K163)

CRASSULACEAE *Sedum acre L., mossy stonecrop (K152, S1156) *Sedum album L., stonecrop (K398)

CUCURBITACEAE Echinocystis lobata (Michx.) Torr. & A. Gray, wild-cucumber (S1492)

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DIERVILLACEAE—SEE CAPRIFOLIACEAE

DIPSACACEAE—SEE CAPRIFOLIACEAE

ELAEAGNACEAE *Elaeagnus umbellata Thunb., autumn-olive (S1420)

ERICACEAE Chimaphila umbellata (L.) Nutt., pipsissewa (S1216) Gaultheria hispidula (L.) Bigelow, creeping-snowberry (S1422) Hypopitys monotropa Crantz, pinesap (S1937) Orthilia secunda (L.) House, one-sided pyrola (S1464) Vaccinium myrtilloides Michx., velvetleaf blueberry (S1421)

EUPHORBIACEAE

Acalypha rhomboidea Raf., three-seeded mercury (S1941, 1955) *Euphorbia cyparissias L., cypress spurge (S1125) *Euphorbia glyptosperma Engelm., ridge-seeded spurge (S1895) *Euphorbia virgata Waldst. & Kit., leafy spurge (S1132)

FABACEAE Apios americana Medik., groundnut (S2553) Desmodium canadense (L.) DC., showy tick-trefoil (S1916) Desmodium illinoense A. Gray, prairie tick-trefoil (S2395) Desmodium paniculatum (L.) DC., panicled tick-trefoil (S1886) Desmodium rotundifolium DC., round-leaved tick-trefoil (S1944) Gleditsia triacanthos L., honey locust (S2554) Hylodesmum glutinosum (Willd.)H. Ohashi&R. R. Mill,clustered-leavedtick-trefoil(S1257) Hylodesmum nudiflorum (L.) H. Ohashi & R. R. Mill, naked tick-trefoil (S1888)

*Lathyrus latifolius L., everlasting pea (S1273) Lathyrus ochroleucus Hook., pale vetchling (S2212) Lathyrus palustris L., marsh pea (S2238)

*Lathyrus sylvestris L., perennial pea (S1462) Lespedeza hirta (L.) Hornem., hairy bush-clover (S1333) Lespedeza violacea (L.) Pers., bush-clover (S1946)

*Lotus corniculatus L., birdfoot trefoil (S1682) *Medicago sativa L., alfalfa (S1687) *Melilotus albus Medik., white sweet-clover (S1480) *Melilotus officinalis (L.) Pall., yellow sweet-clover (S1479) *Robinia hispida L., bristly locust (S1610) *Robinia pseudoacacia L., black locust (S1622) *Securigera varia (L.) Lassen, crown-vetch (S1474) *Trifolium arvense L., rabbitfoot clover (S1626) *Trifolium aureum Pollich, hop clover (K363) *Trifolium pratense L., red clover (S1499) *Trifolium repens L., white clover (K156)

Vicia americana Willd., American vetch (S1155, 1442) *Vicia tetrasperma (L.) Schreb., sparrow vetch (S1639)

FAGACEAE Fagus grandifolia Ehrh., American beech (S1617) Quercus alba L., white oak (S1130) Quercus macrocarpa Michx., bur oak (S1699) Quercus rubra L., red oak (S1713)

GERANIACEAE *Erodium cicutarium (L.) L’Hér, stork’s-bill (S1124) Geranium maculatum L., wild geranium (S1133) *Geranium pusillum L., small geranium (S1806) Geranium robertianum L., herb Robert (S1435)

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GROSSULARIACEAE Ribes americanum Mill., wild black currant (S1406) Ribes triste Pall., swamp red currant (S1418)

HALORAGACEAE Myriophyllum sibiricum Komarov, spiked water-milfoil (S1929) Proserpinaca palustris L., mermaid-weed (S1483)

HYPERICACEAE Hypericum punctatum Lam., spotted St. John’s-wort (S1477)

JUGLANDACEAE Carya cordiformis (Wang.) K. Koch, bitternut hickory (S1673) Juglans nigra L., black walnut (S1684)

LAMIACEAE *Ajuga reptans L., carpet bugle (S2211) *Glechoma hederacea L., ground-ivy (S1412)

Hedeoma hispida Pursh, rough pennyroyal (S1833) *Lamiastrum galeobdolon (L.) Ehrend. & Polatschek, yellow archangel (S1903) *Lamium purpureum L., purple dead-nettle (S1549) *Mentha ×piperita L., peppermint (S1909) *Origanum vulgare L., oregano (S1906)

Pycnanthemum verticillatum (Michx.) Pers., whorled mountain mint (K392). SPECIAL CONCERN

Pycnanthemum virginianum (L.) Durand & Jackson, common mountain mint (S1708)

LENTIBULARIACEAE Utricularia intermedia Hayne, flat-leaved bladderwort (S1335) Utricularia minor L., small bladderwort (S1927) Utricularia purpurea Walter, purple bladderwort (S1707) Utricularia vulgaris L., common bladderwort (S1484)

LYTHRACEAE *Lythrum salicaria L., purple loosestrife (S1945) Rotala ramosior (L.) Koehne, tooth-cup (S1319)

MALVACEAE *Abutilon theophrasti Medik., velvet-leaf (S1939) *Malva neglecta Wallr., common mallow (S1597)

Tilia americana L., basswood (S1203)

MENISPERMACEAE Menispermum canadense L., moonseed (S1434)

MENYANTHACEAE Menyanthes trifoliata L., bogbean (S1604)

MOLLUGINACEAE *Mollugo verticillata L., carpetweed (S1046)

MYRICACEAE Myrica gale L., sweet gale (S1607)

MYRSINACEAE *Lysimachia nummularia L., moneywort (K173, S1204) Lysimachia thyrsiflora L., tufted loosestrife (S1197)

OLEACEAE Fraxinus americana L., white ash (S1819) Fraxinus nigra Marshall, black ash (K204) Fraxinus pennsylvanica Marshall, green ash (S1205, 1711)

*Syringa vulgaris L., common lilac (K361)

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ONAGRACEAE Circaea canadensis (L.) Hill, enchanter’s-nightshade (S1258) *Epilobium hirsutum L., great hairy willow-herb (S1500) Epilobium leptophyllum Raf., fen willow-herb (S1332) *Epilobium parviflorum Schreb., willow-herb (S1900) Ludwigia palustris (L.) Elliott, water-purslane (S1724)

OROHACEAE Agalinis tenuifolia (M. Vahl) Raf., common false foxglove (S1962) Conopholis americana (L.) Wallr., squaw-root (S1832) Epifagus virginiana (L.) Bart., beech-drops (S1957) Pedicularis canadensis L., wood-betony (S1123) Pedicularis lanceolata Michx., swamp-betony (S2552)

OXALIDACEAE Oxalis acetosella L., northern wood-sorrel (S1447) *Oxalis dillenii Jacq., common yellow wood-sorrel (K128)

PAPAVERACEAE *Chelidonium majus L., celandine (S1822) *Papaver orientale L., Oriental poppy (S1827)

PENTHORACEAE Penthorum sedoides L., ditch stonecrop (S1954)

PHRYMACEAE Mimulus ringens L., monkey-flower (S1461) Phryma leptostachya L., lopseed (S1878)

PHYTOLACCACEAE Phytolacca americana L., pokeweed (S1864)

PLANTAGINACEAE

*Chaenorhinum minus (L.) Lange, dwarf-snapdragon (S1920) Gratiola neglecta Torr., clammy hedge hyssop (S1668) Penstemon hirsutus (L.) Willd., hairy beard-tongue (K147a)

*Plantago aristata Michx., bracted plantain (S2398)

Veronica anagallis-aquatica L., water speedwell (S1704) *Veronica arvensis L., field speedwell (S1785) *Veronica officinalis L., common speedwell (K167)

Veronica peregrina L., purslane speedwell (S1787) *Veronica verna L., spring corn speedwell (S1111, 1771)

POLEMONIACEAE Phlox divaricata L., wild blue phlox (S1140) *Phlox subulata L., moss-pink (S1768, 1791)

POLYGONACEAE

Fallopia cilinodis (Michx.) Holub, fringed false buckwheat (S1438) *Fallopia convolvulus (L.) Ã. Löve, black-bindweed (S1868) *Fallopia japonica (Houtt.) Ronse Decr., Japanese knotweed (S1823)

Persicaria hydropiper (L.) Delabare, water-pepper (S1932) Persicaria hydropiperoides (Michx.) Small, mild water-pepper (S1725)

*Persicaria maculosa Gray, lady’s-thumb (S1913) Persicaria sagittata (L.) H. Gross, arrow-leaved tear-thumb (S1731) Persicaria virginiana (L.) Gaertn., jumpseed (S1958)

*Polygonum aviculare L., knotweed (S1343) *Rumex acetosella L., sheep sorrel (K161, S1100) *Rumex crispus L., curly dock (S1863) *Rumex obtusifolius L., bitter dock (S1470)

Rumex orbiculatus A. Gray, great water dock (S1974)

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PORTULACACEAE Portulaca oleracea L., purslane (S1897)

RANUNCULACEAE Actaea pachypoda Elliott, white baneberry (S1845) Actaea rubra (Aiton) Willd., red baneberry (S1498) Anemone canadensis L., Canada anemone (S1269) Enemion biternatum Raf., false rue-anemone (S1762)

*Ranunculus acris L., common buttercup (S1430) Ranunculus fascicularis Bigelow, early buttercup (S1548) Ranunculus flabellaris Raf., yellow water crowfoot (S1812) Ranunculus hispidus Michx. var. caricetorum (Greene) T. Duncan, swamp buttercup (S1135) Ranunculus recurvatus Poir., hooked crowfoot (S1136) Ranunculus sceleratus L., cursed crowfoot (S1999) Thalictrum dioicum L., early meadow-rue (S1391)

RHAMNACEAE *Rhamnus cathartica L., common buckthorn (S1618)

ROSACEAE Agrimonia gryposepala Wallr., tall agrimony (S1491) Agrimonia pubescens Wallr., soft agrimony (S1978) Amelanchier arborea (F. Michx.) Fernald, juneberry (S1541) Amelanchier interior (Pursh) DC., round-leaved serviceberry (S1401) Amelanchier spicata (Lam.) K. Koch, shadbush serviceberry (S1098) Comarum palustre L., marsh cinquefoil (S1224) Crataegus punctata Jacq., dotted hawthorn (K207) Geum aleppicum Jacq., yellow avens (S1237)

*Malus pumila Mill., apple (S1792) Prunus americana Marshall, American wild plum (K208) Prunus nigra Aiton, Canada plum (K130) Prunus virginiana L., choke cherry (S1105) Rosa blanda Aiton, wild rose (S1443)

*Rosa cinnamomea L., cinnamon rose (S1828)

*Rosa multiflora Murray, multiflora rose (S1870) Rubus allegheniensis Porter, common blackberry (S1615) Rubus occidentalis L., black raspberry (S1619) Sibbaldiopsis tridentata (Aiton) Rydb., three-toothed cinquefoil (S1628)

RUBIACEAE Galium circaezans Michx., white wild licorice (S1264) Galium obtusum Bigelow, wild madder (S1698) Galium trifidum L., small bedstraw (S1949)

RUTACEAE Zanthoxylum americanum Mill., prickly-ash (S1259)

SALICACEAE

*Populus alba L., white poplar (S1695) Populus deltoides Marshall, cottonwood (S1824) Populus grandidentata Michx., big-tooth aspen (S1712)

*Populus nigra L., Lombardy poplar (S1714) Salix amygdaloides Andersson, peach-leaved willow (S1653) Salix bebbiana Sarg., beaked willow (S1970) Salix eriocephala Michx., heart-leaved willow (S1468) Salix lucida Muhl., shining willow (S1407, 1469) Salix petiolaris Sm., slender willow (S1485)

SAPINDACEAE Acer negundo L., box-elder (S1411) Acer rubrum L., red maple (S1122)

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Acer saccharinum L., silver maple (S1145) Acer saccharum Marshall, sugar maple (S1611)

SARRACENIACEAE Sarracenia purpurea L., pitcher-plant (S1227)

SAXIFRAGACEAE Chrysosplenium americanum Hook., golden saxifrage (S1423) Micranthes pensylvanica (L.) Haw., swamp saxifrage (S1816) Mitella nuda L., naked miterwort (S1243)

SOLANACEAE *Datura stramonium L., jimson-weed (S1991)

Physalis heterophylla Nees, clammy ground-cherry (S1625) *Solanum carolinense L., horse-nettle (S1933) *Solanum dulcamara L., bittersweet nightshade (S1439)

Solanum ptychanthum Dunal, black nightshade (S1940)

THEOPHRASTACEAE Samolus parviflorus Raf., water-pimpernel (S2550)

ULMACEAE Ulmus americana L., American elm (S1691)

*Ulmus pumila L., Siberian elm (S1804) Ulmus rubra Muhl., slippery elm (S1802) Ulmus thomasii Sarg., rock elm (S1784)

URTICACEAE Boehmeria cylindrica (L.) Sw., false nettle (S1321) Laportea canadensis (L.) Wedd., wood nettle (S1503) Pilea fontana (Lunell) Rydb., bog clearweed (S1928) Pilea pumila (L.) A. Gray, clearweed (S1735) Urtica dioica L., stinging nettle (S1478)

VIOLACEAE Viola canadensis L., Canada violet (S1765) Viola cucullata Aiton, marsh violet (K157)

*Viola odorata L., sweet violet (S2002) Viola rostrata Pursh, long-spurred violet (S1409) Viola sororia Willd., common blue violet (S1390)

VITACEAE Parthenocissus inserta (A. Kern.) Fritsch, thicket creeper (S1683) Parthenocissus quinquefolia (L.) Planch., Virginia creeper (S1881) Vitis aestivalis Michx., summer grape (S1890) Vitis riparia Michx., river-bank grape (S1429)