January-June, 2014

Vol. 53, Nos. 1-2

THE

MICHIGAN BOTANIST

A Journal of Great Lakes Botany

THE MICHIGAN BOTANIST (ISSN 0026-203X) is published four times per year by the Michigan
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Anton A. Reznicek
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Volume 52, Nos. 3-4 was mailed 4-29-2015.

2014

THE MICHIGAN BOTANIST

1

IN THIS ISSUE

Derek Anderson and Julie Fox present the latest in a long line of floristic
studies published in The Michigan Botanist, in this case a flora of Interstate State
Park, Wisconsin’s oldest state park, which is situated in the tension zone along
the St. Croix River in northwestern Wisconsin. The authors take into account
previous collections at the park as well as their own collections to provide a
complete list of the park’s vascular flora as currently known. This is supple¬
mented with a geological and human history of the area, along with a description
of the various plant communities that occur in the park. This paper can be con¬
sidered a companion piece to an earlier flora of Amnicon Falls State Park, which
lies some 150 km to the northeast, near Lake Superior, authored by Paul Hlina,
Derek Anderson, and Donald Davidson (The Michigan Botanist 47: 121-146).
Indeed, the article in this issue provides a comparison of the floras of the two
state parks and discusses how the differences arise from their differing geo¬
graphical settings and floristic regions.

The second article in this issue is a collaborative effort by the members of a
field botany course, co-authored by their teacher, Jordan Marshall. The authors
studied three forest fragments in relatively close proximity to assess how the dif¬
ferences in species composition in canopy, midstory, and understory strata, as
well as in certain factors such as species richness, canopy closure, leaf litter
depth, and incidence of non-native species, are related to the different manage¬
ment histories and levels of protection of the three study sites.

Neil Harriman completes his series of reviews of the field guides written by
Steve Chadde on various regions of the Midwest, including floras of the states of
Wisconsin and Minnesota and of the Upper Peninsula of Michigan, as well as a
guide to the ferns and fem allies of the north central United States. His earlier
survey of six of Chadde’s wetland floras of various states of the Great Lakes re¬
gion, as well as one for the entire region, appeared in the previous issue of The
Michigan Botanist (52: 109-114).

The editor has provided a catalogue of floristic articles from throughout the
whole 53 years of The Michigan Botanist from its inception to date in order to
make this rich resource readily available to current readers and researchers. The
issue is rounded out with two book reviews.

Michael Huft

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THE MICHIGAN BOTANIST

Vol. 53

FLORA OF INTERSTATE STATE PARK,
POLK COUNTY, WISCONSIN

Derek S. Anderson*

St. Croix Falls, WI 54024

Julie Fox

Naturalist, Interstate State Park
St. Croix Falls, WI 54024

ABSTRACT

The vascular plants and plant communities observed at Interstate State Park, located in Polk
County, on the south side of the city of St. Croix Falls in northwestern Wisconsin are presented. The
most commonly encountered plant communities were mesic and dry-mesic hardwood forest. The
vascular plant list includes 652 taxa from 118 families, including 35 ferns and fem allies, nine gym-
nosperms, and 608 angiosperms. Fifty-nine county records were discovered in the course of the sur¬
vey as well as two species listed as Threatened, and five as Special Concern, by the State of Wis¬
consin. These data are presented to document floral composition and to provide a base for future
research activities within the park and the region.

KEYWORDS: Interstate State Park, flora of Wisconsin, St. Croix River

INTRODUCTION

Interstate State Park (the Park) is located in Polk County, on the south side of
the city of St. Croix Falls in northwestern Wisconsin, approximately 70 kilome¬
ters (45 miles) northeast of St. Paul-Minneapolis, Minnesota. Formed in 1900, it
is Wisconsin’s oldest state park and is approximately 538 hectares (1330 acres)
in area. There are three State Natural Areas within the boundary of the Park. The
St. Croix National Scenic Riverway borders the Park to the west, the western ter¬
minus of the Ice Age National Scenic Trail is in the Park, and the Park is also a
unit of the Ice Age National Scientific Reserve. The main features attracting the
public to the Park are glacial potholes and scenic basalt cliffs that form The
Dalles of the St. Croix River. This project was undertaken to document the flora
of this Park and of the greater region, since these data are generally lacking and
incomplete for northwestern Wisconsin.

The Park is primarily forested, and the most co mm on forest types are mesic
forest, dry-mesic forest, floodplain forest, and oak woodlands, following the
classification of natural communities established by the Wisconsin Department
of Natural Resources (2012b). The site is also situated in the middle of the ten¬
sion zone. This zone roughly follows the border separating the northern hard¬
woods province and the prairie-forest province, and as a result, contains species
from both (Curtis 1959). Compiling information on the range limits of 182

* Author for correspondence (derekanderson06@gmail.com)

2014 THE MICHIGAN BOTANIST 3

TABLE 1: Collection contributions by major collectors at Interstate State Park, showing for each
total collections and collections of the four largest plant families at the Park.

Collector

Plant Families

Total

Asteraceae

Poaceae

Cyperaceae

Rosaceae

Other

Anderson, Derek S.

722

73

83

70

31

465

Cochrane, Theodore S.

63

8

10

4

0

41

Fassett, Norman C.

84

15

12

8

2

47

Patman, Jacqueline

98

8

11

11

7

61

Pohl, Richard W.

187

21

22

7

13

124

plants, Curtis created maps identifying this zone where a greater number of
northern and southern species reach the edges of their range (Curtis 1959; An¬
derson 2005). Several uncommon habitats such as cliffs (dry and moist), forest
seeps, bedrock glades, sedge meadows and emergent aquatic communities add to
an already diverse flora.

The Park has a long history of botanizing, but until this survey, there had not
been any exhaustive documentation of the flora. Richard Pohl conducted one of
the more intensive surveys of the Park in 1936. Other noteworthy collecting trips
were made by Norman Fassett in 1927 and 1934, Jacqueline Patman in 1959,
and Theodore Cochrane in 1972 (Robert W. Freckman Herbarium 2015, Wis¬
consin State Herbarium 2015). A comparison of the contributions by past col¬
lectors with those of the authors is presented in Table 1.

A review of online databases of Wisconsin herbaria revealed a number of
species that had been collected in the past and deposited in the state’s herbaria
but that were not relocated during this study. These species are indicated in Ap¬
pendix I by the citation of collector and collection number, year of collection,
and herbarium acronym.

Early public land survey records (Haight 1847-1848, Jenkins 1847, Whitcher
1847) indicate that the tree composition in the area of the Park in the mid-1800s
is similar to that of today. Surveyors characterized the Park as being “very bro¬
ken,” since it “abounds with traprock [basalt]” (Haight 1847-1848). Other sur¬
veyors described the landscape in and around the Park as hilly and stony. They
noted that the trees typical in the area of this rocky region of the Park included
linden, birch, white oak, elm, hickory, pine, and butternut (presumably Tilia
americana, Betula sp., Quercus alba and/or Q. macrocarpa, Ulmus sp., Cary a
cordiformis, Pinus strobus, and Juglans cinerea, respectively). The understory
in these rocky areas also included hazel and prickly ash (presumably Corylus sp.,
and Zanthoxylum americanum ). Interestingly, the notes that describe section 36
in Township 34N, Range 19W mention prairie, which is consistent with the ex¬
tensive bedrock glade community seen in the vicinity today.

The section line notes for the area near the St. Croix River indicate the pres¬
ence of such species as white maple, black ash, willow, elm and linden (pre¬
sumably Acer saccharinum, Fraxinus nigra, Salix sp., Ulmus sp., and Tilia
americana, respectively), which confirms the wetter nature of this area. The
notes also reveal that in upland areas beyond the basalt exposures, species in¬
dicative of a mesic forest became common, including sugar maple, linden, white

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THE MICHIGAN BOTANIST

Vol. 53

walnut, white ash, pine, elm and oak (presumed to be Acer saccharum, Tilia
americana, Juglans cinerea, Fraxinus americana, Pinus strobus, Ulmus sp., and
Quercus rubra, respectively), and, in the understory, ironwood ( Ostrya virgini-
ana) and prickly ash.

The climate of the region is continental. Based on weather data from St. Croix
Falls, the mean annual precipitation is 77.7 cm, and August is the month with the
greatest mean monthly precipitation (11.9 cm). The mean annual temperature for
the area is 7° C; July is the hottest month (mean 28.9° C), and January is the
coldest (mean -17.6° C). The length of the frost-free growing season ranges
from 117 days to 172 days with an average of 144 days per year (Midwestern
Regional Climate Center 2015).

Development within the Park includes a contact station, a nature interpretive
center, two campgrounds, a swimming beach, a boat landing, approximately
14.4 kilometers of hiking trails, and 20.9 kilometers of winter recreation trails
(Wisconsin Department of Natural Resources 2012a). The most disturbed areas
of the Park are a gas pipeline right-of-way and the Silverbrook Mansion site on
the south end of the Park. Although most of the infrastructure of the mansion is
gone, foundation remnants and trout ponds that were created by the berming of
springs remain. Other portions of the Park continue to recover from logging,
mining, grazing, and agricultural activities that took place around the turn of the
twentieth century.

Geology

The complex landscape of the Park is the result of geological events spanning
a billion years. Evidence of lava flows, faulting, flooding, glaciation, and erosion
can be seen in the Park. The geologic events of the past have directly and indi¬
rectly impacted the landscape and plant co mm unities as they exist in the Park
today.

Late in the Precambrian, about 1.1 billion years ago, tectonic forces began to
pull the North American continent apart. Outpourings of lava, called fissure
flows, continued for millions of years, completely burying the pre-existing land¬
scape in this area beneath thousands of feet of Keweenawan basalt. Eight indi¬
vidual basalt flows have been mapped in the area (Cordua 1989). The weight of
the lava and accumulated sediment at the surface resulted in the Lake Superior
syncline, a down-warping of the earth’s surface. The northeast-southwest axis of
the syncline lies west of the Park; as a result, the lava flows in this area dip to¬
ward the west. Faulting also occurred in many places within the syncline.

Toward the end of Precambrian time, a long period of erosion began that
lasted a few hundred million years and wore deep valleys into the Keweenawan
sediments and the rocks below. Then, starting in the Late Cambrian, this region
was submerged under a shallow sea. Hundreds of feet of sediments reburied the
Keweenawan rocks under layers of sedimentary rock. Where sea cliffs stood at
the edge of the sea, boulders fell into the wave zone where the surf tumbled,
broke, and ground them smooth, resulting in a conglomerate rock layer at the
base of the much thicker overlying sandstone (Mickelson et al. 2011). Ravines

2014

THE MICHIGAN BOTANIST

5

containing examples of basal conglomerate are evidence of the ancient exhumed
landscape of this area.

Eventually the area rose above sea level, followed by another long period of
erosion lasting over 300 million years. Gradual erosion ended two million years
ago when the Ice Age began in Wisconsin. During the Pleistocene, several
episodes of glacial and interglacial periods altered the landscape, each advance
leaving little evidence of earlier glaciations.

The most recent glacial episode was the Wisconsin Glaciation, which began
about 100,000 years ago. The best record of late Wisconsin ice cover is pre¬
served on bedrock surfaces as striations, which indicate the direction of ice flow.
Johnson (2000) interpreted the striations at the tops of Observation Rock and
Eagle Peak as having been formed by the Superior Lobe during the last glacia¬
tion. The Superior Lobe was one of six major lobes of ice that advanced over
Wisconsin during the Wisconsin Glaciation. This lobe of ice deepened the Lake
Superior basin as the ice advanced along the Lake Superior syncline, and is thus
named after Lake Superior (Johnson 2000).

Extensive erosion of the St. Croix River gorge and the formation of the pot¬
holes occurred very late during the retreat of the Superior Lobe. The retreating
ice exposed the Lake Superior basin. Melt water filled the basin to form glacial
Lake Duluth at the ice margin. Glacier ice blocked drainage of the melt water to
other outlets. As a result, glacial Lake Duluth eventually overflowed, flooding
the Bois Brule-St. Croix valley. At the Park, torrents of glacial melt water filled
the valley to an elevation of at least 274 meters (900 feet), as is evident at Eagle
Peak by rock eroded by the river (Mickelson et al. 2011).

The rapidly flowing river repeatedly ripped away pieces of the highly frac¬
tured basalt. Continued cutting of the gorge resulted in the deep, narrow forma¬
tion known as The Dalles of the St. Croix. Lake o’ the Dalles is another feature
of the torrential flow of the river. It is a plunge pool basin formed at the base of
cascading water as the glacial St. Croix River tumbled over the bluffs of Summit
Rock.

The rushing melt water also formed glacial potholes in the bedrock 6-21 m
(20-70 ft) above the present level of the river (Figure 1). In the fast-flowing cur¬
rent of the river, recurring eddies moved debris in a circular motion on the
riverbed. Over time, the spinning debris drilled holes into the bedrock. Rocks
caught in the swirling water were worn to a smooth, spherical shape. These rocks
are called grindstones.

Land History

Little is known about the earliest inhabitants of the area now preserved in the
Park. Settlers tell of finding stone spear points and axes and other prehistoric
tools about the Park (Pond 1937). Archeological surveys have not identified any
definite village or campsite dating to prehistoric times. In 1936, an important
archeological discovery provided evidence of prehistoric mammals inhabiting
this area. A number of large bones were uncovered by Civilian Conservation
Corps workmen while digging in peat not far from Lake o’ the Dalles. The bones
were identified as Bison occidentals, an extinct species of bison. Based on bone

6

THE MICHIGAN BOTANIST

Vol. 53

FIGURE 1. A glacial pothole carved into the basalt bedrock along the St. Croix River. Photograph
by Derek Anderson, October 1, 2010.

dating, B. occidentalis persisted in the region for about 4,000 years, with the
greatest presence between ca. 8,000-7,000 calibrated years B.P. (Hawley et al.
2013). Over 1400 bison, deer, and elk bones were recovered from the excavation
site, as well as one copper tool.

The first European to record travel from Lake Superior to the Mississippi
River via the Bois Brule-St. Croix portage was Daniel Greysolon, Sieur du Lhut,
in 1679 (du Lhut is the namesake of Duluth, Minnesota). It was the French, dur¬
ing this early period of exploration, who named this river the St. Croix, or Holy
Cross river, and described the river gorge as the Dalles —a word to describe
rivers with steep walls and fast-flowing water. Du Lhut claimed the region for
France and offered gifts to the Dakota Indians, the first historic inhabitants of the
St. Croix Valley (McMahon and Karamanski 2009).

The river became a regular route for missionaries, fur traders, and other voy¬
agers. Ojibwa Indians began migrating into the valley in the seventeenth century
and became involved in the fur trade. McMahon and Karamanski (2009) dis¬
cussed the eventual and mounting tension and violence between the Ojibwa and
Dakota that escalated into an epic battle fought here at the Dalles of the St. Croix
around 1770. Combined war parties of Fox and Dakota Indians met Ojibwa war¬
riors in combat on the portage trail around the St. Croix Falls. After a fierce bat¬
tle with heavy losses on all sides, the Ojibwa emerged victorious. An uneasy
boundary between the Ojibwa and Dakota was established, yet skirmishes con¬
tinued for the next several decades.

By the mid-1800s, the fur trade had diminished and the logging industry

2014

THE MICHIGAN BOTANIST

7

began in earnest due to the rising market for lumber in the Mississippi River Val¬
ley and beyond. For more than a half century, the St. Croix River was used to
transport pine logs to mills and markets downstream. The most spectacular event
of the logging days occurred in the Dalles in the spring of 1886. More than 150
million feet of pine were stacked in a logjam that extended three miles upstream
of the 90-degree bend in the river known as the “Elbow of the St. Croix.” By
1914, the last log drive had reached the mills and the logging boom was over.
Other business ventures such as copper mining, traprock excavation and blast¬
ing, and construction of the hydroelectric dam at the “falls” of Taylors Falls and
St. Croix Falls had also begun by the 1890s (St. Croix Falls Area History, com¬
piled by Knudson, unpublished).

Local citizens became concerned about preserving the scenic beauty of the
river gorge. In the mid-1890s, George Hazzard of Taylors Falls, Minnesota,
began encouraging citizens of that community to lobby their legislators to create
a state park on the Minnesota side of the Dalles. As a result, Minnesota Interstate
State Park was established in 1895. Harry Baker of St. Croix Falls, Wisconsin,
was doing the same in his community. After five years of numerous trips to
Madison, and fundraisers by local businessmen to support the lobbying efforts,
the legislators finally agreed to set aside funds to purchase land on the Wiscon¬
sin side of the Dalles of the St. Croix River. On September 20, 1900, the State of
Wisconsin acquired the first parcel of land here, officially establishing Interstate
State Park as Wisconsin’s oldest state park (Baker, unpublished letter 1961).

Approximately 30 hectares (a little more than 72 acres) of land was purchased
in 1900, mostly from within the city limits of St. Croix Falls. In 1907 and 1908,
more than 200 additional hectares (500 acres) were purchased. These purchases
included what are now the Dalles of the St. Croix and the Lake o’ the Dalles
areas. Over the next several years lands were acquired bit by bit from several ad¬
jacent landowners, thereby providing protection for the more sensitive areas of
the Park (Interstate State Park acquisition files, unpublished).

Despite land acquisition, little development took place within the Park until
the 1930s and the creation of the Civilian Conservation Corps (CCC). In the
summer of 1935 the barracks of CCC Camp Interstate were constructed not far
from Lake o’ the Dalles. In November, 1935, CCC Company 633 arrived from
Grafton, Illinois, to begin work in the Park. Development projects included ex¬
tension and improvement of the park road; construction of stone buildings, shel¬
ters, and other structures; and construction of 16 kilometers (10 miles) of hiking
trails (Pond 1937). Late in 1937 Camp Interstate was abandoned until the fol¬
lowing year when CCC Company 4610 arrived to continue the work until they
were disbanded in 1940. Many of the stone structures and nine miles of the hik¬
ing trails built by the CCC are still in use today.

Additional land acquisitions included approximately 100 hectares (240 acres)
purchased in Osceola Township from the Riegel family in 1964 and about 80
hectares (196 acres) purchased from the Mills family in 1970, the latter of which
nowconstitutes the Silverbrook area (Interstate State Park acquisition files, un¬
published).

Significant to further protection of the natural and cultural resources of the
Park was the passage of two acts of Congress. In 1968 the Upper St. Croix River

THE MICHIGAN BOTANIST

Vol. 53

was one of eight original rivers protected by the National Wild and Scenic
Rivers Act. In 1972 that act was amended to include the Lower St. Croix. In
1971 another act of Congress created the Ice Age National Scientific Reserve,
established to preserve Wisconsin’s glacial heritage. Interstate State Park is one
of nine units of the Reserve. As a result, federal funding was received to aid in
new development at the Park. In 1982 a new park entrance and office, a mainte¬
nance building, and the Ice Age Interpretive Center were completed.

METHODS

Study Area

Interstate State Park is a 538 hectare park located along the east edge of the St. Croix River, south
of St. Croix Falls in Polk County, Wisconsin (Figure 2). This area was well kn own throughout his¬
tory and was often referred to as the “Falls of the St. Croix”. The actual falls were a series of rapids
where the St. Croix River dropped about 12 meters (40 feet) in elevation over the course of 9.6 kilo¬
meters (6 miles). Today, most of these rapids are hidden underwater because of a hydroelectric dam
that was completed in 1906.

There are a number of places within the Park referenced by nicknames used to describe the dif¬
ferent features and places. Eagle Peak, a basalt exposure, is 274 meters (900 feet) above sea level. It
is one of the highest points in the Park. Geographically, it is located in the center of the Park. Sum¬
mit Rock, another basalt rock feature, is 250 meters (830 feet) above sea level. It is the highest point
in the Park located along the St. Croix River gorge (the area described as the Dalles). Observation
Rock is a third basalt feature nicknamed for the panoramic view it provides of the St. Croix River
Valley. It is found on the last 1.5 kilometers (ca 1 mile) of the Ice Age Trail.

The lakes observed within the Park are connected to the St. Croix River. Fake o’ the Dalles is a
spring fed lake centrally located within the Park, just south of Summit Rock. A small stream flows
from this lake along a former river channel to the St. Croix River. Folsom Fake is on the southern
boundary of the Park and is backwater slough.

Also found at the southern end of the Park is an area referred to as the Silverbrook Mansion.
When it was in private ownership, the estate developed the area. A downhill ski slope was cleared,
and several trout ponds were created by berming natural springs near the home site. The mansion
was razed in 1974, and now only foundational ruins remain.

Data for this flora project were collected during the growing seasons of 2011 to 2013. Seven hun¬
dred twenty-two voucher specimens representing 612 taxa were collected, identified, and deposited
in the Robert W. Freckmann Herbarium of the University of Wisconsin-Stevens Point in Stevens
Point, Wisconsin (UWSP). These are listed in Appendix I. An additional 40 taxa that were previ¬
ously documented at the Park and represented by collections at UWSP or the Wisconsin State
Herbarium of the University of Wisconsin-Madison (WIS) but not located during the present survey
are also included in this list. The following sources were used for plant identification: Fassett (1951,
1978), Gleason and Cronquist (1998), Holmgren (1998), Hipp (2008), Crow and Hellquist (2000),
Smith (2008), and Voss and Reznicek (2012). Nomenclature follows Reznicek et al. (2011) for the
ferns and fem allies and Voss and Reznicek (2012) for the seed plants. Six taxa not included in Voss
and Reznicek (2012) follow the nomenclature of the Flora of North America (1993+)- These are An-
tennaria plantaginifolia, Delphinium carolinianum, Phemeranthus rugospermus, Rubus alumnus,
Tradescantia occidentalis, and Vernonia fasciculata.

General species composition was noted for plant communities in the Park. We visited each com¬
munity throughout the Park recording the dominant plants in each stratum (canopy, understory and
ground layers) if present. This information was used to describe the basic composition of the plant
communities. The nomenclature for the plant communities follows that used by the Wisconsin DNR
Natural Heritage Inventory (Wisconsin Department of Natural Resources 2012b). One or two visits
were made to the Park each week during the growing season. An extra effort was placed on visiting
forested communities in the spring to capture ephemerals, while later-season survey effort was fo¬
cused on wetlands and other more open communities. Meander surveys to locate rare species (Goff
et al. 1982) were undertaken in the more uncommon communities of the Park.

A list of target species to search for while conducting surveys was compiled from several re-

2014

THE MICHIGAN BOTANIST

9

Plant Communities of Interstate State Park

Legend

Cliff

Sandstone

Plant Communities

Bedrock Glade
~j Developed Area
Emergent Marsh
1:-^^ Floodplain Forest

Forested Seeps - VWet-Mesic Forest
Formerly Disturbed Area
11 Northern Dry-Mesic Forest
: Northern Sedge Meadow
I | Oak Woodland
Open Water

[ T 7'' T | Pine Plan la lion

^ Southern Dry-Mesic Forest
■ Southern Mesic Forest

FIGURE 2. Map of the major plant communities of Interstate State Park.

10

THE MICHIGAN BOTANIST

Vol. 53

sources. These included databases of previous collections from the Park housed at Wisconsin
herbaria (Robert W. Freckmann Herbarium 2015; Wisconsin State Herbarium 2015) and previously
prepared flora checklists on file at the Park (Crawford 1987, unpublished; Delaney 2000, unpub¬
lished). These unpublished checklists were based on observation rather than the collection of speci¬
mens preserved in an herbarium. Effort was placed on surveying plant communities that would likely
provide habitat for the species on these lists.

The map of native plant communities was created using ESRI ArcGIS 9.3 software. Plant com¬
munity boundaries were drawn using US Geologic Survey topographic quadrangle maps (1:24,000),
the 2008 Farm Service Agency’s National Agriculture Imagery Program (NAIP) color and color in¬
frared aerial images of Minnesota, and notes collected during ground surveys.

RESULTS

A total of 652 vascular plant taxa representing 366 genera and 118 families
were documented from the Park (see Appendix I). Ferns and fern allies were rep¬
resented by 35 taxa, gymnosperms by 9 taxa, and angiosperms by 608 taxa. Of
the angiosperms, 184 taxa were monocots and 424 were dicots. The predominant
plant families were Asteraceae with 68 taxa, Poaceae with 67 taxa, Cyperaceae
with 57 taxa, and Rosaceae with 33 taxa. These totals include two rare species
listed as Threatened and five species listed as Special Concern by the Wisconsin
Department of Natural Resources, Bureau of Endangered Resources (Wisconsin
Department of Natural Resources 2011c) (Table 2). In addition, three of the
species were previously tracked by the Bureau of Endangered Resources, but
were removed from the state’s endangered species list during the course of the
survey. These are Carex assiniboinensis, Juglans cinerea, and Taxus canaden¬
sis. Ninety-four of the species are introduced, some of which are ecologically in¬
vasive.

When comparing the species we collected with species previously docu¬
mented by collections in the State’s herbaria (UWSP, WIS), we identified 59
species not previously reported from Polk County (Table 3). These include 25
species that are introduced and naturalized in the state of Wisconsin.

TABLE 2: Documented rare plant species at Interstate State Park (Wisconsin Department of Natural
Resources 2012c). Species indicated as a “New Record” were not previously known from the Park.
“Not Actively Tracked” indicates species that were formerly tracked by the Bureau of Endangered
Resources at the start of this survey but are no longer included in the department’s database.

Species

New Record

State Status

Besseya bullii

Threatened

Carex assiniboinensis

Not Actively Tracked

Carex backii

X

Special Concern

Cystopteris laurentiana

X

Special Concern

Dryopteris fragrans

Special Concern

Juglans cinerea

Not Actively Tracked

Opuntia fragilis

Threatened

Panax quinquefolius

X

Special Concern

Phemeranthus rugospermus

Special Concern

Taxus canadensis

X

Special Concern

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TABLE 3: New Polk County Records (an asterisk indicates introduced species).

Species

Abies balsamea

*Iris pseudacorus

*Acer ginnala

Lactuca biennis

Actaea pachypoda

Liparis liliifolia

*Alliaria petiolata

*Malus pumila

Anemone cylindrica

*Matricaria discoidea

*Berberis thunbergii

*Medicago lupulina

Carex backii

Menispermum canadense

Carex buxbaumii

Mertensia virginica

Carex haydenii

Oxalis dillenii

Carex hirtifolia

Panax quinquefolius

Carex lurida

Poa annua

Carex muhlenbergii

Prunus pumila

Carex radiata

Quercus alba

Carex stricta

Ranunculus hispidus var. nitidus

Carex typhina

*Robinia pseudoacacia

*Convallaria majalis

*Rumex crispus

Cystopteris laurentiana

Salix petiolaris

*Dianthus armeria

*Salix xrubens

Dicentra cucullaria

*Sedum acre

Dichanthelium linearifolium

*Sorbus aucuparia

*Draba vema

*Stellaria media

*Elaeagnus angustifolia

*Tanacetum vulgare

Fragaria vesca

Tsuga canadensis

*Galinsoga quadriradiata

*Ulmus pumila

Gaylussacia baccata

Vaccinium myrtilloides

*Gypsophila muralis

*Veronica officinalis

*Hieracium caespitosum

*Veronica vema

Hierochloe hirta

*Vicia cracca

Hypoxis hirusta

Impatiens pallida

Zannichellia palustris

PLANT COMMUNITIES

The location of the Park within the tension zone described by Curtis (1959)
makes it difficult to distinguish plant communities with southern and with north¬
ern affinities (in particular, the forests) from each other. The following descrip¬
tions of plant communities within the Park indicate, among other things, the
dominant species observed in each community. The map in Figure 2 highlights
the major plant communities found within the Park, generated on the basis of a
field survey and the examination of aerial photographs.

Southern Mesic Forest

Southern mesic forest, together with southern dry-mesic forest, comprises the
majority of the acreage of the Park. These two communities grade into one an¬
other with changes in topography, slope aspect, and soil composition. The south¬
ern mesic forest is most commonly encountered in the eastern half of the Park. It
is found on gentle to steep slopes, and there is little or no influence from the un¬
derlying bedrock on its composition. The southern mesic forests within the Park

FIGURE 3. Typical forest community on south-facing slopes and ridge tops that lack bedrock near
the surface. The forest is dominated by red oak (Quercus rubra) and basswood (Tilia americana).
Sugar maple (Acer saccharum ) is common in the understory, and there is a diverse ground layer.
Photograph by Derek Anderson, May 10, 2012.

are dominated by Acer saccharum. Occasionally Quercus rubra, Tilia ameri¬
cana, Carya cordiformis, Fraxinus americana, and F. pennsylvanica are canopy
co-dominants. The understory is dominated by Acer saccharum and Ostrya vir-
giniana. The ground layer is diverse and consists of Trillium grandiflorum, T.
cernuum, Hepatica americana, H. acutiloba, Hydrophyllum virginianum, Adi-
antum pedatum, Athyrium filix-femina, Caulophyllum thalictroides, and many
other spring ephemerals.

Southern Dry-Mesic Forest

As noted above, southern dry-mesic forests in the Park are often found in
close association with southern mesic forest. Dry-mesic forest often occurs near
ridge tops with moderate to well-drained soils (Figure 3) and on south-facing
slopes. It also occurs near oak woodland communities where bedrock outcrops
are more prevalent. The canopy is dominated by Quercus rubra and Tilia amer¬
icana. The understory is often dominated by Acer saccharum. The ground layer
is usually diverse. The more common ground species observed include Gera¬
nium maculatum, Aralia nudicaulis, Hylodesmum glutinosum, and Arisaema tri-
phyllum.

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13

FIGURE 4. The floodplain forest along the St. Croix River in early spring. The forest is dominated
by silver maple (Acer saccharinum ); hackberry (Celtis occidentalis) and cottonwood (Populus del-
toides ) are also present. Water begins to pool in old channels as the snow and ice begin to melt. The
water in this area of the Park can easily rise an additional 3 to 6 meters (10 to 20 feet) as spring rain
and seasonal snowmelt occurs in the greater watershed. Photograph by Derek Anderson, April 2,
2011 .

Floodplain Forest

A well-developed example of floodplain forest in northern Wisconsin can be
observed on the western edge of the Park. This riparian zone is dominated by
Acer saccharinum, wad. Populus deltoides, Celtis occidentalis, and Acer negundo
are occasional co-dominants (Figure 4). Spring floods inundate the area for sev¬
eral weeks each year, and there are several natural, well-developed channels
throughout the area of this community. Species common in the herbaceous layer
include Viola sororia, Laportea canadensis, Matteuccia struthiopteris, and Rud-
beckia laciniata. Lobelia cardinalis is one of the more striking species observed
in this community in late summer, especially along the seasonal water channels.
As these channels dry out over the course of the summer, they are dominated by
Eleocharis acicularis, E. obtusa, E. ovata, Cyperus erythrorhizos, C. odoratus,
and C. strigosus.

Forested Seeps (with Southern Wet-Mesic Forest Inclusions)

A number of forested seeps can be found along the steep slopes above the St.
Croix River. This is especially true in the southern reaches of the Park where the
more porous sandstone facilitates the movement of groundwater. The seepage

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THE MICHIGAN BOTANIST

Vol. 53

FIGURE 5. A forested seep located along the slopes above the St. Croix River. The canopy is patchy
and dominated by black ash (Fraxinus nigra). The ground layer is dominated by skunk cabbage
(Symplocarpus foetidus). Photograph by Derek Anderson, May 12, 2012.

zones in these areas underlie a patchy canopy dominated by Fraxinus nigra, with
Ulmus americana and Betula allegheniensis as occasional co-dominants. Slight
rises in the topography surrounding the seepage areas tend to support Acer sac-
charum and Tilia americana. In the areas of ground water seepage, Symplocar¬
pus foetidus is dominant (Figure 5), and there is a continuous cover in the ground
layer. Other species present in the seeps include Impatiens capensis, Hydro-
cotyle americana, Viola sororia, Poa palustris, Packera aurea, and Micranthes
pensylvanica. Many of these seeps feed small streams that eventually reach the
St. Croix River.

Dry and Moist Cliffs

There are several cliffs throughout the Park, which are composed of one of
two basic lithologies, basalt or sandstone. The many exposures of these cliffs
provide a full spectrum of moisture conditions. Several of these cliffs are di¬
rectly exposed to the sun, whereas others are completely shaded by well-devel¬
oped forest canopies (for example, some of the largest individuals of Pinus
strobus in the Park can be found growing from the cliffs). Several of the cliffs
are wet from groundwater seepage. The most prominent area occupied by this
plant community is the Dalles of the St. Croix River. Basalt cliffs rise 30 meters
(100 feet) above the river. Smaller basalt cliffs of 5-15 meters (15-50 feet) can
be found a few hundred meters from the St. Croix River and along abandoned

2014

THE MICHIGAN BOTANIST

15

FIGURE 6. The state special concern fragrant fern {Dryopteris
fragrans) has long been documented in the area of the Park. It
is typically found growing from the cracks of basalt near the St.
Croix River. Photograph by Derek Anderson, 2007.

river channels, such as in the area of Meadow Valley. At the bases of some of
these cliffs are large talus slopes and fields where the basalt has weathered from
the cliffs. The sandstone cliffs are most prominent in the southern region of the
Park west of the Silverbrook Mansion and are not readily accessible by trail.

Vegetation is sparse on these cliffs, and both types of cliffs tend to be domi¬
nated by a wide variety of pteridophytes. The most common of these include
Polypodium virginianum, Woodsia ilvensis, and Cystopteris fragilis. Three ferns
that are rare in the state of Wisconsin, Dryopteris fragrans (Figure 6),
Cystopteris laurentiana, and Woodsia oregana, are found in this community.
Woodsia oregana had been previously documented at the Park but was not relo¬
cated during the course of this survey. Herbaceous species that are also found
along these cliffs include Aquilegia canadensis. Campanula rotundifolia, Cory-
dalis aurea, and Capnoides sempervirens.

16

THE MICHIGAN BOTANIST

Vol. 53

Northern Dry-Mesic Forest

This community occupies a fairly small portion of the overall acreage of the
Park. It is associated with draws and ravines of the basalt cliffs along the St.
Croix River. The canopy is dominated by Pinus strobus with occasional P.
resinosa, Quercus rubra and Q. ellipsoidalis. The shrub layer contains Vac-
cinium angustifolium, and the ground layer contains Maianthemum canadense,
Aralia nudicaulis, Cornus canadensis, and Trientalis borealis.

Oak Woodland

The oak woodland plant community is found between the bedrock glades and
the dry-mesic forests throughout the Park. It is also found as an inclusion within
the mesic forests where large portions of bedrock are exposed. Typically, this
community has an interrupted or patchy canopy, dominated by Quercus macro-
carpa, Q. alba, and Q. ellipsoidalis. Juniperus virginiana is an associate in some
stands. Shrubs are sparse, but include Corylus americana, C. cornuta, Rhus ty-
phina, R. glabra, and Zanthoxylum americanum. The ground layer includes Ely-
mus hystrix, Andropogon gerardii, Solidago spp., Toxicodendron rydbergii, and
Galium boreale. This community also provides habitat for the state Threatened
Besseya bullii (Figure 7).

mEa. 7 ' jjjg 1 m

#■ u SM

^ | FIGURE 7. Kitten tails (Besseya bullii ) in

an oak woodland near a transition be-
lUuhL 1 tween woodland and bedrock glade. This

species is listed as threatened in Wiscon-
^ sin. Photograph by Derek Anderson, June

| 2,2013.

2014

THE MICHIGAN BOTANIST

17

FIGURE 8. One of several bedrock glades found throughout the Park. Lichen and moss covered
basalt is in the foreground. Prairie vegetation and oak woodlands with Quercus macrocarpa, Q. el-
lipsoidalis, and Q. alba surround the exposed bedrock. Photograph by Derek Anderson, July 23,
2011 .

Bedrock Glade

The bedrock glade community in the Park is found where basalt bedrock is
exposed or is near the surface. These sites tend to be dry, and they support a
unique assemblage of plants adapted to xeric conditions, such as Selaginella ru-
pestris, lichens, and mosses. Where soil has developed and accumulated, prairie,
savanna, and barrens species can be observed (Figure 8). These species include
Schizachyrium scoparium, Andropogon gerardii, Sorghastrum nutans. Coreop¬
sis palmata, Lespedeza capitata, Liatris aspera, and Quercus ellipsoidalis. Op-
untia fragilis (Figure 9) and Phemeranthus rugospermus are two rare species
that are found within this community. Bedrock glades are found throughout the
Park; the largest areas are west of Eagle Peak and in the southern reaches of the
Park. Smaller inclusions of this community occasionally occur within mesic
forests where basalt bedrock is exposed.

Northern Sedge Meadow

There are a number of sedge meadows throughout the Park, the largest ap¬
proaching five hectares (12 acres) in area. The majority of the sedge meadow
communities in the Park are dominated by Phalaris arundinacea. Closer exami¬
nation reveals a few diverse, intact remnants of native vegetation within the

18

THE MICHIGAN BOTANIST

Vol. 53

FIGURE 9. The brittle prickly pear cactus (Opuntia fragilis) is listed as threatened in Wisconsin.
This species is restricted to areas of bedrock glade habitat within the Park. Photograph by Derek An¬
derson June 28, 2011.

larger context of Phalaris arundinacea. These remnants tend to be dominated by
Car ex lacustris, although a few are dominated by Car ex stricta. Other
graminoids found in this community include Glyceria canadensis, G. striata,
Poa palustris, Calamagrostis canadensis, and several species of Carex and Scir-
pus. Several forbs are also present, including Doellingeria umbellata, Eu-
trochium maculatum, Epilobium ciliatum, E. leptophyllum, Asclepias incarnata,
and Campanula aparinoides.

Emergent Marsh and Submergent Aquatic

Aquatic communities are found in a few locations within the Park. A large
emergent marsh is present in the backwaters of the St. Croix River, in the area of
Folsom Lake. This backwater is inundated most of the year with 15-100 cm
(6^10 inches) of water (Figure 10). In drought years mud flats become exposed.
The emergent marsh is dominated by Bolboschoenus fluviatilis, and lesser
amounts of Sagittaria latifolia and S. rigida are present.

Submerged aquatic communities are present in the areas of Folsom Lake and
of Lake o’ the Dalles. In these bodies of water, Potamogeton nodosus and P.
pusillus are common. Elodea canadensis, E. nuttallii, and Ceratophyllum de-
mersum are present in pockets within these lakes. Zannichellia palustris is found
in water channels running through the emergent marsh in Folsom Lake. This is
a new discovery for the Park and for Polk County. Potamogeton crispus, an in¬
vasive non-native species, occurs in both Lake o’ the Dalles and Folsom Lake.

2014

THE MICHIGAN BOTANIST

19

FIGURE 10. The north end of the river backwater named Folsom Lake. River bulrush ( Bol-
boschoenus fluviatilis) dominates in the right center area of the photograph. The open water contains
long-leaved pondweed (Potamogeton nodosus) and slender pondweed (P. pusillus). Photograph by
Derek Anderson, July 27, 2012.

DISCUSSION

The forest flora at the Park has been affected by about 100 years of secondary
successional regeneration of forest in northwestern Wisconsin. The diverse flora
of the site is also influenced by its location in the tension zone and by such abi¬
otic factors as bedrock, topography, and groundwater seepage. The high diver¬
sity of plant species found in the Park becomes evident when comparing this site
with Amnicon Falls State Park, approximately 150 km northeast, which was the
subject of a previous floristic study. Hlina et al. (2008) identified a total of 400
taxa at Amnicon Falls State Park. The two parks share 297 species in common.
One hundred three of the species at Amnicon Falls State Park have not been doc¬
umented at Interstate State Park. These include several species with cooler cli¬
mate (or northern) affinities, including Carex castanea, C. ormostachya, Hale-
nia deflexa, Petasites frigidus var. sagittatus, Rubus parviflorus, Shepherdia
canadensis, and Sparganium angustifolium. In contrast, Interstate State Park has
355 species that were not documented at Amnicon Falls State Park. Several of
these species, including Boechera canadensis, Carex blanda, Ludwigia poly-
carpa, Ranunculus fascicularis, Staphylea trifolia, and Trillium flexipes, have
not been documented north of the tension zone described by Curtis (1959).

A total of 163 species were documented for the Park for the first time and, of
those, 59 were also newly documented for Polk County. It is evident from these

20

THE MICHIGAN BOTANIST

Vol. 53

numbers that gaps still remain in our knowledge of plant distributions within the
state, particularly in those regions located the farthest from the state’s universi¬
ties. The myth of a well-catalogued flora is not unique to the Park, and, as Ertter
(2000) points out, there are several examples to the contrary throughout North
America. Although many new species records were discovered for the Park and
for Polk County, 40 taxa previously collected from the Park were not relocated
during this three-year study. There is no evident correlation between the indi¬
vidual taxa and the inability to relocate them within the Park (such as the disap¬
pearance of northern species that may have been lost to a changing climate or the
loss of a given community). Instead, they may actually no longer be found
within the Park, or they may have been simply overlooked.

The diverse landscape of the Park supports several rare species (Table 2). One
of these species, Besseya bullii (Figure 7), was rediscovered during the course of
the survey. An older collection of the species from 1959 did not include a spe¬
cific location in the Park. As a result, it is uncertain whether the new record is
from the same population as the older collection. Another species of particular
interest is the fern, Dryopteris fragrans (Figure 6), which was first collected at
the “Falls of St. Croix” by Charles Parry in 1848. According to correspondence
between Parry and Dr. John Torrey, this was the first time the species was doc¬
umented within the limits of the United States. Parry (1852) indicated that the
plant was “quite abundant.” This species now appears to be quite rare in the
Park. Between 2011 and 2013, only a few small populations were discovered on
the extensive cliffs. litis and Judziewicz (1994) did not locate this species during
their rare plant survey of the St. Croix National Scenic Riverway. They noted
that extensive collections were made dating back to 1861 and that over-collect¬
ing at this site had likely contributed to the decline.

In addition to rare species, other surprise discoveries were made during the
course of this survey. One was of a lone eastern hemlock ( Tsuga canadensis )
discovered in a mesic hardwood forest in the north end of the Park in the region
that was first acquired in 1900. The tree was growing a few meters from a long-
abandoned hiking trail. This hemlock tree had a diameter at breast height of 17.5
cm (6.8 in.). By comparing this measurement to the average diameters of trees in
Michigan as documented by Bums and Honkala (1990), we conclude that this
tree is most likely between forty and sixty years old. Its origins remain a bit of a
mystery, since the nearest known site is approximately one hundred kilometers
to the east.

Invasive species are another factor that impacts the diversity and abundance
of native plant species in forests. Many of the non-native species in the Park are
not particularly invasive and are restricted to previously disturbed land, such as
the gas pipeline right-of-way, roadways, trails, and campgrounds. However, one
of the more aggressively invasive species discovered at the Park in the course of
the survey was garlic mustard ( Alliaria petiolata). This species should be moni¬
tored and managed, particularly since Knight et al. (2009) found that the pres¬
ence of white-tailed deer can lead to a significant divergence in plant community
stmcture, and to an increase in the relative cover of garlic mustard. In addition,
the non-native honeysuckles, Lonicera tatarica and L. xbella, and common
buckthorn, Rhamnus cathartica, were observed in populations ranging from

2014

THE MICHIGAN BOTANIST

21

scattered individuals to dense patches throughout the Park. Management activi¬
ties have been directed toward these species when funding allows. Phalaris
arundinacea has become a dominant species in several larger wetlands that were
likely sedge meadows prior to the invasion.

As similarly reported in an earlier floristic study of Amnicon Falls State Park
(Hlina et al. 2008), Interstate State Park has become a refuge for white-tailed
deer ( Odoceoileus virginianus), despite a large portion of the Park being open to
hunting. Evidence of deer browse and winter deer yards can be observed
throughout. The white-tailed deer appear to be impacting the forests within the
Park, as reported in a number of studies examining such impacts (Graham 1954;
Stoeckeler et al. 1957; Beals et al. 1960; Alverson et al. 1988; Balgooyen and
Waller 1995; Mudrak et al. 2009). It appears that some woody species such as
Thuja occidentalis and Taxus canadensis are not regenerating. Impacts are not
restricted to woody species, however; the herbaceous layer in these forests has
also been impacted in plant communities of northern Wisconsin with high deer
populations (Rooney et al. 2004; Holmes et al. 2008).

The documentation of the flora at Interstate State Park sets the stage for fu¬
ture research and management activities at the Park. Follow-up activities could
include: 1) monitoring and removal of invasive plant populations; 2) monitoring
rare plant populations; 3) monitoring impacts of the white-tailed deer population
on plant species and communities; 4) assessing comparisons of phenological
shifts in certain species over time; and 5) using the results of this study as a base¬
line for future floristic work in the Park and the surrounding area.

ACKNOWLEDGMENTS

Thanks are extended to the staff of Interstate State Park. Thanks are also extended to Barb De¬
laney for sharing her knowledge of the Park’s flora and for her assistance in locating several species
in the Park. Special thanks are also extended to Dr. Robert Freckmann and Dr. Emmet Judziewicz of
the University of Wisconsin—Stevens Point for their assistance in plant identification and for sharing
their knowledge of plants and their distribution in the state of Wisconsin. We would also like to
thank Maureen Yunker for compiling the land history information relating to the general acquisition
of Park lands. We would like to tha nk Merel Black for running database queries. Thanks are ex¬
tended to Paul Hlina, Janeen Ruby, Nancy Sather, and Daniel Wovcha for the helpful comments they
provided in reviewing this manuscript. And finally, we would like to thank two anonymous review¬
ers and the editor, Michael Huft, for their helpful comments and suggestions.

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787-798.

Smith, W. R. (2008). Trees and shrubs of Minnesota. University of Minnesota Press, Minneapolis.

Stoeckeler, J. H., R. O. Strothman, and L. W. Krefting. (1957). Effect of deer browsing on repro¬
duction in the northern hardwood-hemlock type in northeastern Wisconsin. Journal of Wildlife
Management 21: 75-80.

Voss, E. G. and A. A. Reznicek. (2012). Field Manual of Michigan Flora. The University of Michi¬
gan Press, A nn Arbor, Michigan.

Whitcher, J. E. (1847). Survey and subdivision of township 33 north range 19 west of the 4th merid¬
ian Wisconsin in Wisconsin Public Land Survey Records: Original field notes and plat maps.
(1833—1866). State of Wisconsin Board of Commissioners of public lands. Available at
http://digicoll.library.wisc.edu/SurveyNotes/SurveyNotesHome.html. (Accessed June 22, 2015).

Wisconsin Department of Natural Resources. (2015a). Interstate State Park. Available at
http://dnr.wi.gov/topic/parks/name/interstate/. (Accessed June 22, 2015).

Wisconsin Department of Natural Resources. (2015b). Wisconsin Natural Heritage Inventory (NHI)
Recognized Natural Communities. Available at http://dnr.wi.gov/topic/endangeredresources/
communities.asp. (Accessed June 18, 2015).

Wisconsin Department of Natural Resources. (2015c). Wisconsin Natural Heritage Working List.
Available at http://dnr.wi.gov/topic/NHI/documents/NHIWorkingList.pdf. (Accessed June 22,
2015).

Wisconsin State Herbarium (2015). University of Wisconsin - Madison. Available at
http://herbarium.wisc.edu/. (Accessed June 11, 2015).

APPENDIX I: INTERSTATE STATE PARK FLORA LIST

The following list is organized alphabetically, first by family, then by genus, and finally by
species, in each major group of plants. Non-native species are indicated with an asterisk. All collec¬
tions in the course of this survey were made by Anderson and are deposited in the Robert W. Freck¬
mann Herbarium at the University of Wisconsin-Stevens Point (UWSP). The name of each species
is followed by a co mm on name and, in parentheses, Anderson’s collection numbers. This list also
contains forty species collected in the past but not relocated during the course of our study. Each of
these species is followed by a common name, the collector’s name in italics, their collection number,
year of collection and the herbarium acronym. Nomenclature follows Reznicek et al. (2011) for the
ferns and fem allies and Voss and Reznicek (2012) for the seed plants. Six taxa not included in Voss
and Reznicek (2012) follow the nomenclature of the Flora of North America (1993+)- These ar eAn-
tennaria plantaginifolia. Delphinium carolinianum, Phemeranthus rugospermus, Rubus alumnus,
Tradescantia occidentalis, and Vernonia fasciculata as they are not known to occur in Michigan.

PTERIDOPHYTES

ASPLENIACEAE (Spleenwort Family)

Asplenium rhizophyllum L., walking fern (1576)

ATHYRIACEAE (Lady Fem Family)

Athyrium fdix-femina (L.) Roth, common lady fem (1498)

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CYSTOPTERIDACEAE (Brittle Fern Family)

Cystopteris bulbifera (L.) Bernh., bladder fern (1839, 1842)

Cystopteris fragilis (L.) Bernh., brittle bladder fern (1555)

Cystopteris laurentiana (Weath.) Blasdell, Laurentian bladder fern (1554)

Cystopteris tenuis (Michx.) Desv., MacKay’s brittle fern (1618)

Gymnocarpium dryopteris (L.) Newman, oak fern (1586)

DENNSTAEDTIACEAE (Bracken Fern Family)

Pteridium aquilinum (L.) Kuhn. var. latiusculum (Desv.) A. Heller, bracken fern (1647)

DRY OPTERID ACEAE (Wood Fern Family)

Dryopteris carthusiana (Vill.) H. P. Fuchs, spinulose wood fern (1532)

Dryopteris cristata (L.) A. Gray, crested shield fem (1522)

Dryopteris fragrans (L.) Schott, fragrant fem (1553)

EQUISETACEAE (Horsetail Family)

Equisetum arvense L., field horsetail (1720, 1721)

Equisetum hyemale L., common scouring rush (1531, 1624)

Equisetum pratense Ehrh., meadow horsetail (1961)

Equisetum scirpoides Michx., dwarf scouring rush (1620)

Equisetum sylvaticum L., wood horsetail (2226)

Equisetum xferrissii Clute, Ferris’ horsetail; Fassett 15557, 1927 (WIS)

LYCOPODLACEAE (Club-Moss Family)

Dendrolycopodium obscurum (L.) A. Haines, princess’-pine (1934)

Diphasiastrum digitatum (A. Braun) Holub, southern ground-cedar (1443, 1935)

Huperzia lucidula (Michx.) Trevis., shining club moss (1933)

ONOCLEACEAE (Ostrich Fem Family)

Matteuccia struthiopteris (L.) Todaro, ostrich fem (1958)

Onoclea sensibilis L., sensitive fem (1869)

OPHIOGLOSSACEAE (Adder’s-Tongue Family)

Botrypus virginianus (L.) Michx., rattlesnake fem (1598)

Sceptridium dissectum (Spreng.) Lyon, cut-leaved grape fem (810, 1468, 1936)

OSMUNDACEAE (Royal Fem Family)

Osmunda cinnamomea L., cinnamon fem (1612)

Osmunda claytoniana L., interrupted fem (1611)

Osmunda regalis L., royal fem (1672, 1676)

POLYPODIACEAE (Polypody Fem Family)

Polypodium virginianum L., common polypody fem (1641)

PTERIDACEAE (Maidenhair Fem Family)

Adiantum pedatum L., maidenhair fem (1575)

SELAGINELLACEAE (Spikemoss Family)

Selaginella rupestris (L.) Spring, rock spikemoss (1473)

THELYPTERIDACEAE (Marsh Fem Family)

Phegopteris connectilis (L.) Slosson, northern beech fem; Fassett 15506, 1927, (WIS)
Thelypteris palustris Schott var . pubescens (Lawson) Femald (1870)

WOODSIACEAE (Woodsia Fem Family)

Woodsia ilvensis (L.) R. Br., rusty cliff fem (1971)

Woodsia obtusa (Spreng.) Torn, blunt-lobed cliff fem (1523)

Woodsia oregana D. C. Eaton subsp. cathcartiana (B. L. Rob.) Windham, Oregon cliff fem,
Tyrone 4143, 1938 (WIS)

GYMNOSPERMS

CUPRESSACEAE (Cypress Family)

Juniperus communis L., common juniper (1748)

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Juniperus virginiana L., eastern red-cedar (1743)

Thuja occidentalis L., northern white cedar (1719)

PINACEAE (Pine Family)

Abies balsamea (L.) Mill., balsam fir (2138)

Picea glauca (Moench) Voss, white spruce (1938)

Pinus resinosa Aiton, red pine (1986)

Pinus strobus L., white pine (1977)

Tsuga canadensis (L.) Carriere, eastern hemlock (2139, 2140)

TAXACEAE (Yew Family)

Taxus canadensis Marshall, American yew (1441)

DICOTYLEDONS

ADOXACEAE (Moschatel Family)

Sambucus canadensis L., American elderberry (1804)

Sambucus racemosa L., red-berried elder (1473)

Viburnum lentago L., nannyberry (1565, 1573)

Viburnum rafinesquianum Schult., downy arrow-wood (1584)

Viburnum trilobum Marshall, American cranberry bush (1984)

AMARANTH ACE AE (Amaranth Family)

Amaranthus retroflexus L., rough pigweed (2051)

Amaranthus tuberculatus (Moq.) J. D. Sauer, rough-fruited amaranth (2245)
Chenopodium album L., common lamb’s quarters (1983)

Chenopodium simplex (Torn) Raf., maple-leaved goosefoot (1774)

ANACARDIACEAE (Cashew Family)

Rhus glabra L., smooth sumac (2000)

Rhus typhina L., staghorn sumac (2030)

Rhus xpulvinata Green, hybrid sumac; Patman s.n., 1959 (WIS)
Toxicodendron rydbergii (Rydb.) Greene, western poison ivy (2127)

APIACEAE (Parsley Family)

Angelica atropurpurea L., purple-stem angelica (1764)

Cicuta maculata L., water hemlock (1707)

Cryptotaenia canadensis (L.) DC., honewort (1596, 1648)

Osmorhiza claytonii (Michx.) C. B. Clarke, bland sweet cicely (1621)
Osmorhiza longistylis (Torn) DC., anise-root (1528)

Sanicula marilandica L., black snakeroot (1604, 1619)

Sanicula odorata (Raf.) Pryer & Phillippe, clustered black snakeroot (1599)
Zizia aurea (L.) W. D. J. Koch, golden alexanders (1546)

APOCYNACEAE (Dogbane and Milkweed Family)

Apocynum androsaemifolium L., spreading dogbane (1640)

Apocynum cannabinum L., Indian-hemp (2009)

Asclepias exaltata L., poke milkweed (1791)

Asclepias incamata L., swamp milkweed (1750)

Asclepias syriaca L., common milkweed (2020)

Asclepias tuberosa L., butterfly weed (1776)

Asclepias verticillata L., whorled milkweed (2070)

AQUIFOLIACEAE (Holly Family)

Ilex verticillata (L.) A. Gray, winterberry (1671)

ARALIACEAE (Ginseng Family)

Aralia nudicaulis L., wild sarsaparilla (1510)

Aralia racemosa L., American spikenard (1809)

Hydrocotyle americana L., marsh pennywort (1841)

Panax quinquefolius L., American ginseng (1990)

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ARISTOLOCHIACEAE (Birthwort Family)

As arum canadense L., wild ginger (1457)

ASTERACEAE (Sunflower Family)

Achillea millefolium L., yarrow (1710)

Ageratina altissima (L.) R. M. King & H. Rob., white snakeroot (1901)

Ambrosia artemisiifolia L., co mm on ragweed (1813)

Antennaria neglecta Greene, field pussy-toes (1480, 1488)

Antennaria plantaginifolia (L.) Hook., plantain pussy-toes (1469)

* Arctium minus (Hill) Berhn., common burdock (2028)

Artemisia campestris L. subsp. caudata (Michx.) H. M. Hall & Clem, field wormwood
(1899)

Artemisia ludoviciana Nutt, subsp. ludoviciana, white sage (1854)

Artemisia serrata Nutt., saw-toothed sagebrush (1867)

Bidens cemua L., nodding beggar-ticks (2116)

Bidens comosa (A. Gray) Wiegand, straw-stem beggar-ticks (2113)

Bidens connata Willd., purple-stem beggar-ticks; Koch 7158, 1971 (UWSP)

Bidens frondosa L., common beggar-ticks (2112)

*Centaurea stoebe L., spotted knapweed (1897)

*Cirsium arvense (L.) Scop., Canada thistle (2071)

Cirsium discolor (Willd.) Spreng., field thistle (2075)

Cirsium muticum Michx., swamp thistle (1887)

*Cirsium vulgare (Savi) Ten., bull thistle (2050)

Conyza canadensis (L.) Cronquist, Canadian horseweed (1843)

Coreopsis palmata Nutt., prairie coreopsis (1775)

*Crepis tectorum L., hawk’s beard (1635)

Doellingeria umbellata (Mill.) Nees, flat-topped aster (1871)

Erechtites hieraciifolius (L.) DC., American bumweed (2059)

Erigeron philadelphicus L., common fleabane (1614, 2006)

Erigeron strigosus Willd., daisy fleabane (2189)

Eupatorium perfoliatum L., boneset (1860)

Eurybia macrophylla (L.) Cass., big-leaved aster (1824)

Euthamia graminifolia (L.) Nutt., grass-leaved goldenrod (2076)

Eutrochium maculatum (L.) E. E. Lamont, spotted Joe-Pye weed (1825)

Eutrochium purpureum (L.) E. E. Lamont, purple-node Joe-Pye weed (1429)

*Galinsoga quadriradiata Cav., common quickweed (1928)

Gnaphalium uliginosum L., marsh cudweed (2115)

Helenium autumnale L., common sneezeweed (1922)

Helianthus giganteus L., giant sunflower; Pohl 560, 1936 (WIS)

Helianthus pauciflorus Nutt., stiff sunflower (1434)

Helianthus strumosus L., woodland sunflower (1821, 1915)

Helianthus tuberosus L., Jerusalum-artichoke (2033)

Heliopsis helianthoides (L.) Sweet, ox-eye (1755)

*Hieracium aurantiacum L. orange hawkweed (1605)

*Hieracium caespitosum Durmot., yellow king-devil (1663)

Hieracium umbellatum L., northern hawkweed (1919, 2069)

Lactuca biennis (Moench) Femald, tall blue lettuce (2233)

Lactuca canadensis L., Canada lettuce (2065)

*Leucanthemum vulgare Lam., ox-eye daisy (1715)

Liatris aspera Michx., rough blazing star (1855)

* Matricaria discoidea DC., pineapple-weed (2012)

Packera aurea (L.) A. Love & D. Love, golden ragwort (1627)

Prenanthes alba L., white-lettuce (2141)

Rudbeckia hirta L. var .pulcherrima Farw., black-eyed Susan (1757)

Rudbeckia laciniata L., cut-leaved coneflower (1828)

Solidago canadensis L., Canada goldenrod (2052)

Solidago flexicaulis L., zig-zag goldenrod (1918)

Solidago gigantea Aiton, giant goldenrod (1820)

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Solidago hispida Willd. var. hispida, hairy goldenrod (1435)

Solidago juncea Aiton, early goldenrod (1436)

Solidago nemoralis Aiton, gray goldenrod (1856)

Symphyotrichum laeve (L.) A. Love & D. Love, smooth aster; Sayre s.n., 1937 (WIS)
Symphyotrichum lanceolatum (Willd.) G. L. Nesom, panicled aster (2053, 2077)
Symphyotrichum novae-angliae (L.) G. L. Nesom, New England aster (1879)
Symphyotrichum oolentangiense (Riddell) G. L. Nesom, sky-blue aster (1929)
Symphyotrichum puniceum (L.) A. Love & D. Love, swamp aster (2064)

Symphyotrichum sericeum (Vent.) G. L. Nesom, silky aster (2126)

Symphyotrichum urophyllum (DC.) G. L. Nesom, arrow-leaved aster (1917, 1932)
*Tanacetum vulgare L., common tansy (2049)

* Taraxacum officinale F. H. Wigg., common dandelion (1954)

*Tragopogon dubius Scop., goat’s beard (1592, 1609)

Vernonia fasciculata Michx., ironweed (1851, 1852)

Xanthium strumarium L., common cocklebur (2107)

BALSAMINACEAE (Touch-Me-Not Family)

Impatiens capensis Meerb., orange jewelweed (1859)

Impatiens pallida Nutt., yellow jewelweed (1878)

BERBERIDACEAE (Barberry Family)

*Berberis thunbergii DC., Japanese barberry (1654)

Caulophyllum thalictroides (L.) Michx., blue cohosh (1693)

BETULACEAE (Birch Family)

Alnus incana (L.) Moench subsp. rugosa (Du Roi) R. T. Clausen, speckled alder (1682)
Betula alleghaniensis Britton, yellow birch (1477)

Betula papyrifera Marshall, paper birch (1795)

Carpinus caroliniana Walter, American hornbeam (1941)

Corylus americana Walter, American hazelnut (1937)

Corylus cornuta Marshall subsp. cornuta, beaked hazelnut (1940)

Ostrya virginiana (Mill.) K. Koch, ironwood (1763)

BORAGINACEAE (Borage Family)

Hackelia deflexa (Wahlenb.) Opiz var. americana (A. Gray) Femald and I. M. Johnst., cliff
stickseed (1792)

Hackelia virginiana (L.) I. M. Johnst., beggar’s-lice (1752)

Hydrophyllum virginianum L., Virginia waterleaf (1533)

Mertensia virginica (L.) Pers. ex Link, Virginia bluebells (1967)

BRAS SIC ACEAE (Mustard Family)

*Alliaria petiolata (M. Bieb.) Cavara & Grande, garlic mustard (1481)

Arabidopsis lyrata (L.) O’Kane & Al-Shehbaz, sand cress (1474, 1497)

*Barbarea vulgaris Aiton, yellow rocket (1517)

*Berteroa incana (L.) DC., hoary-alyssum (2001)

Boechera canadensis (L.) Al-Shehbaz (1652, 1653)

Boechera grahamii (Lehmann) Windham & Al-Shebaz (1513, 1534)

*Capsella bursa-pastoris (L.) Medik., shepherd’s-purse (1560)

Cardamine bulbosa (Muhl.) Britton, Stems & Poggenb., spring cress (1559)

Cardamine parviflora L. var. arenicola (Britton) O. E. Schulz, dry-land bitter-cress (1494)
Cardaminepensylvanica Willd., Pennsylvania bitter-cress (1527)

*Draba verna L., spring whitlow grass (1496)

*Erysimum cheiranthoides L., worm-seed mustard (1798)

*Hesperis matronalis L., dame’s rocket (1995)

*Lepidium densiflorum Schrad., prairie pepper-weed (2247)

* Nasturtium officinale R. Br., watercress (1610)

Rorippa palustris (L.) Besser subsp. palustris, marsh cress (2015)

Sisymbrium altissimum L., tall tumble mustard; Patman s.n., 1959 (WIS)

Turritis glabra L., tower mustard (1651)

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CACTACEAE (Cactus Family)

Opuntia fragilis (Nutt.) Haw., brittle prickly pear (1738)

CAMPANULACEAE (Bellflower Family)

Campanula aparinoides Pursh, marsh bell flower (1758)

* Campanula rapunculoides L., European bell flower (1891)

Campanula rotundifolia L., harebell (1630)

Lobelia cardinalis L., cardinal flower (1836)

Lobelia inflata L., Indian tobacco (1872, 1895)

Lobelia siphilitica L., great blue lobelia (1428)

CANNABACEAE (Hemp Family)

Humulus lupulus L., common hops (1868)

Celtis occidentalis L., hackberry (1681)

CAPRIFOLLACEAE (Honeysuckle Family)

Lonicera dioica L., red honeysuckle (1970)

*Lonicera tatarica L., Tatarian honeysuckle (1536, 1563)

* Lonicera xbella Zabel, showy honeysuckle (1978)

Symphoricarpos albus (L.) S. F. Blake, snowbeny (1639)

Triosteum aurantiacum E. P. Bicknell, early horse gentian (1507)

CARYOPHYLLACEAE (Pink Family)

*Cerastium fontanum Baumg., common chickweed (1666)

*Dianthus armeria L., Deptford pink (1823)

*Gypsophila muralis L., cushion baby’s breath (2013)

Moehringia lateriflora (L.) Fenzl, sandwort (1495)

*Myosoton aquaticum (L.) Moench, giant chickweed (1543, 2183)

*Saponaria officinalis L., bouncing-bet (2043)

Silene antirrhina L., sleepy catchfly (2186)

*Silene latifolia Poir, bladder campion (1646)

Stellaria longifolia Willd., long-leaved stichwort (1594)

*Stellaria media (L.) Vill., co mm on chickweed (2184)

CELASTRACEAE (Bittersweet Family)

Celastrus scandens L., American bittersweet (1862)

CERATOPHYLLACEAE (Homwort Family)

Ceratophyllum demersum L., coon’s tail (1803,1908)

CISTACEAE (Rock-Rose Family)

Crocanthemum bicknellii (Femald) Janch., hoary frostweed (2230)

Lechea intermedia Britton, intermediate pinweed (1814)

CONVOLVULACEAE (Morning Gloiy Family)

Calystegia sepium (L.) R. Br., hedge bindweed (1754)

Cuscuta gronovii Roem. & Schult. var. gronovii, common dodder (1877)

CORNACEAE (Dogwood Family)

Cornus altemifolia L.f., pagoda dogwood (1617)

Cornus amomum Mill, (blue-fruited dogwood); Pohl 550, 1936 (WIS)

Cornus canadensis L., bunchberry (1582)

Cornus foemina Mill, subsp. racemosa (Lam.) J. S. Wilson, gray dogwood (1667, 1670)
Cornus rugosa Lam., round-leaved dogwood (1648)

Cornus sericea L., red osier dogwood (1762)

CRASSULACEAE (Sedum Family)

*Sedum acre L., gold-moss stonecrop (1686)

CUCURBITACEAE (Gourd Family)

Echinocystis lobata (Michx.) Torr. & A. Gray, wild cucumber (2072)

DIERVILLACEAE (Bush-honeysuckle Family)

Diervilla lonicera Mill., northern bush-honeysuckle (1628)

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ELAEAGNACEAE (Oleaster Family)

*Elaeagnus angustifolia L., Russan olive (1987)

ERICACEAE (Heath Family)

Arctostaphylos uva-ursi (L.) Spreng., bearberry (1746, 1955)

Chimaphila umbellata (L.) W. P. C. Barton, pipsissewa (1962)

Gaylussacia baccata (Wangenh.) K. Koch, black huckleberry (1972)

Monotropa uniflora L., Indian-pipe (1893)

Pyrola elliptica Nutt., large-leaved shin-leaf (1973)

Vaccinium angustifolium Aiton, early low blueberry (1472)

Vaccinium myrtilloides Michx., velvet-leaved blueberry (1585)

EUPHORBIACEAE (Spurge Family)

Acalypha rhomboidea Raf., three-seeded mercury (1838, 1914)

Euphorbia corollata L., flowering spurge (1784)

Euphorbia cyparissias L., cypress spurge (1601)

Euphorbia glyptosperma Engelm., ridge-seeded spurge; Pohl 583, 1936 (WIS)
Euphorbia maculata (L.) Small, milk purslane (1927, 2074)

Euphorbia nutans Lag., nodding spurge; Fassett 5483, 1927 (WIS)

FABACEAE (Bean or Pea Family)

Amorpha canescens Pursh, leadplant (1745)

Amorpha fruticosa L., false indigo (1649)

Amphicarpaea bracteata (L.) Femald, hog peanut (1921)

Apios americana Medik., common groundnut (1866)

Dalea purpurea Vent., purple prairie clover (1911)

Desmodium canadense (L.) DC., showy tick-trefoil (2191)

Hylodesmum glutinosum (Willd.) H. Ohashi & R. R. Mill, pointed tick-trefoil (1787)
Lathyrus ochroleucus Hook., white pea (1539)

Lathyrus venosus Willd., forest pea (1564)

Lespedeza capitata Michx., round-headed bush clover (1873)

* Lotus comiculatus L., bird’s-foot trefoil (1714)

*Medicago lupidina L., black medick (1669)

*Melilotus albus Medik., white sweet clover (1744)

*Melilotus officinalis (L.) Pall., yellow sweet clover (1633)

*Robinia pseudoacacia L., black locust (2192)

*Securigera varia L., crown-vetch (1713)

*Trifolium arvense L., rabbit’s-foot clover (1778)

* Trifolium campestre Schreb., field clover (1779)

* Trifolium hybridum L., alsike clover (2193)

* Trifolium pratense L., red clover (1615)

*Trifolium repens L., white clover (1623)

Vicia americana Willd., American vetch (1580)

*Vicia cracca L., cow vetch (1997)

FAGACEAE (Beech Family)

Quercus alba L., white oak (1709)

Quercus ellipsoidalis E. J. Hill, northern pin oak (1718)

Quercus macrocarpa Michx., bur oak (1874)

Quercus rubra L., northern red oak (1991)

Quercus xbebbiana C. K. Schneid., Bebb’s oak; Fassett 15552, 1927 (WIS)

GENTIANACEAE (Gentian Family)

Gentiana andrewsii Griseb., bottle gentian (1923, 1924)

GERANIACEAE (Geranium Family)

Geranium maculatum L., wild geranium (1524)

GROSSULARIACEAE (Gooseberry Family)

Ribes cynosbati L., prickly wild gooseberry (1504)

Ribes hirtellum Michx., swamp gooseberry (1518)

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HAMAMELIDACEAE (Witch Hazel Family)

Hamamelis virginiana L., American witch hazel (2134)

HYPERICACEAE (St. John’s Wort Family)

Hypericum ascyron L., giant St. John’s wort (1766)

Hypericum majus (A. Gray) Britton, larger St. John’s wort; Pohl 571, 1936 (WIS)
*Hypericum perforatum L., common St. John’s wort (1655)

Triadenum fraseri (Spach) Gleason, marsh St. John’s wort (2061)

JUGLANDACEAE (Walnut Family)

Carya cordifonnis (Wangenh.) K. Koch, bittemut hickory (1679)

Juglans cinerea L., butternut (1688)

Juglans nigra L., black walnut (1595)

LAMIACEAE (Mint Family)

Agastache scrophulariaefolia (Willd.) Kuntze, purple giant hyssop (1827, 1853)

*Galeopsis tetrahit L., common hemp-nettle (2029)

*Glechoma hederacea L., creeping Charlie (1471)

Hedeoma hispida Pursh., rough false pennyroyal (1625)

*Leonurus cardiaca L., motherwort (1756)

Lycopus americanus W. P. C. Barton, American water-horehound (1840)

Lycopus uniflorus Michx., northern bugleweed (2036)

Mentha canadensis L., wild mint (2040)

Monarda fistulosa L., bee balm (1770)

Physostegia virginiana (L.) Benth. subsp. virginiana, false-dragonhead (1903)

Prunella vulgaris L., heal-all (1694)

Pycnanthemum virginianum (L.) B. L. Rob & Femald, Virginia mountain mint (1704, 1822)
Scutellaria lateriflora L., skullcap (1425, 2035)

Scutellaria parvula Michx. var. missouriensis (Torr.) Goodman & C. A. Lawson, smooth
small skullcap (1637)

Stachys arenicola Britton., marsh hedge-nettle (2045)

Teucrium canadense L., American germander (1767, 1896)

Trichostema brachiatum L., false-pennyroyal (1817, 2232)

LIMNANTHACEAE (Meadow-Foam Family)

Floerkea proserpinacoides Willd., false mermaid (1530)

LINDERNIACEAE (False Pimpernel Family)

Lindernia dubia (L.) Pennell, false pimpernel (2114)

MALVACEAE (Mallow Family)

*Malva neglecta Wallr., cheeses (1785)

Tilia americana L., basswood (1780)

MENISPERMACEAE (Moonseed Family)

Menispermum canadense L., Canada moonseed (2045)

MOLLUGINACEAE (Carpetweed Family)

*Mollugo verticillata L., carpetweed (1910)

MONTIACEAE (Blinks Family)

Claytonia virginica L., spring beauty (1460)

MYRSINACEAE (Myrsine Family)

Lysimachia ciliata L., fringed loosestrife (1742, 2027)

Lysimachia hybrida Michx., river loosestrife (2042)

Lysimachia terrestris (L.) Britton, Stems & Poggenb., swamp loosestrife (2235)

Trientalis borealis Raf., American starflower (1538)

NYCTAGINACEAE (Four-O’clock Family)

Mirabilis nyctaginea (Michx.) MacMill., wild four-o’clock (1797)

NYMPHAEACEAE (Water-Lily Family)

Nymphaea odorata Aiton, fragrant water-lily (2047)

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OLEACEAE (Olive Family)

Fraxinus americana L., white ash (2137)

Fraxinus nigra Marshall, black ash (1574)

Fraxinus pennsylvanica Marshall, green ash (1683)

ONAGRACEAE (Evening-Primrose Family)

Chamerion angustifolium (L.) Holub, fireweed (1760)

Circaea canadensis (L.) Hill subsp. canadensis, enchanter’s nightshade (1741)
Epilobium ciliatum Raf., American willow-herb (1427, 1876, 2067, 2228)
Epilobium leptophyllum Raf., American marsh willow-herb (2066)

Ludwigia palustris (L.) Elliot, marsh purslane (1837)

Ludwigia polycarpa Short & R. Peter, false-loosestrife (2237)

Oenothera biennis L., evening-primrose (1831)

Oenotheraperennis L., small sun-drops (1650, 1703, 2190)

OROBANCHACEAE (Broom-rape Family)

Agalinis tenuifolia (Vahl) Raf., common false foxglove (1920)

Aureolaria pedicularia (L.) Raf., false foxglove; Pohl 588, 1936 (WIS)
Pedicularis canadensis L., wood-betony (1501)

Pedicularis lanceolata Michx., swamp lousewort (1898)

OXALIDACEAE (Wood-Sorrel Family)

Oxalis dillenii Jacq., southern yellow wood-sorrel (1998)

Oxalis stricta L., common wood-sorrel (1662)

Oxalis violacea L., violet wood-sorrel (1515)

PAPAVERACEAE (Poppy Family)

Capnoides sempervirens (L.) Borkh., pale corydalis (1540)

Corydalis aurea Willd. subsp. aurea, golden corydalis (1556)

Dicentra cucullaria (L.) Bemh., Dutchman’s breeches (1458)

Sanguinaria canadensis L., bloodroot (1447)

PENTHORACEAE (Stonecrop Family)

Penthorum sedoides L., ditch stonecrop (2025, 2236)

PHRYMACEAE (Lopseed Family)

Mimulus ringens L. var. ringens, monkey-flower (2110)

Phryma leptostachya L., American lop-seed (1753)

PLANTAGINACEAE (Plantain Family)

Besseya bullii (Eaton) Rydb., kittentails (2142)

Chelone glabra L., turtlehead (1861)

Gratiola neglecta Torn, clammy hedge-hyssop (2111,2118)

*Linaria vulgaris Mill., butter and eggs (1992)

*Plantago major L., common plantain (2129)

Plantago rugelii Decne., American plantain (1890)

Veronica beccabunga L. var. americana Raf., American brooklime (1607)

* Veronica officinalis L., common speedwell (1613)

* Veronica serpyllifolia L., thyme-leaved speedwell (1590)

*Veronica verna L., spring speedwell (1508)

Veronicastrum virginicum (L.) Farw., culver’s root (1830)

POLEMONIACEAE (Phlox Family)

Phlox divaricata L., forest phlox (1492)

Phlox pilosa L., prairie phlox (1591)

POLYGALACEAE (Milkwort Family)

Polygala sanguinea L., field milkwort (1886, 2231)

POLYGONACEAE (Buckwheat Family)

Fallopia cilinodis (Michx.) Holub, fringed bindweed (1658, 1690)

*Fallopia convolvulus (L.) A. Love, black-bindweed (2185)

Persicaria amphibia (L.) A. Gray, water smartweed (1912)

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*Persicaria hydropiper (L.) Delarbre (marsh-pepper knotweed); Fassett 12756, 1927 (WIS)
Persicaria pensylvanica (L.) Gomez, Pennsylvania smartweed (1863, 2194)

Persicaria punctata (Elliot) Small, dotted smartweed; Pohl 572, 1936 (WIS)

Persicaria sagittata (L.) H. Gross, arrow-leaved tear-thumb (1888)

Persicaria virginiana (L.) Gaert., jumpseed (1844)

Polygonum tenue Michx., pleat-leaf knotweed (1818)

*Rumex acetosella L., sheep sorrel (1514)

Rumex altissimus A. W. Wood, pale dock (2026)

*Rumex crispus L., curly dock (2018)

*Rumex obtusifolius L. (bitter dock); Pohl 594, 1936 (WIS)

Rumex orbiculatus A. Gray, great water dock (2062)

PORTULACACEAE (Purslane Family)

Phemeranthus rugospermus (Holz.) Kiger, prairie fame-flower (1812)

*Portulaca oleracea L., common purselane (2032)

PRIMULACEAE (Primrose Family)

Androsace occidentalis Pursh, rock-jasmine (1948)

RANUNCULACEAE (Buttercup Family)

Actaea pachypoda L., white baneberry (1439)

Actaea rubra (Aiton) Willd., red baneberry (1519)

Anemone canadensis L., Canada anemone (1578)

Anemone cylindrica A. Gray, thimbleweed (2182)

Anemone quinquefolia L., wood anemone (1451)

Anemone virginiana L., tall anemone (1593)

Aquilegia canadensis L., wild columbine (1509)

Caltha palustris L., marsh marigold (1452)

Clematis virginiana L., virgin’s bower (1829)

Delphinium carolinianum Walter subsp. virescens (Nutt.) R. E. Brooks, Carolina larkspur
(2181)

Enemion biternatum Raf., false rue-anemone (1953)

Hepatica acutiloba (DC.) G. Lawson, sharp-lobed hepatica (1449)

Hepatica americana (DC.) H. Hara, round-lobed hepatica (1446)

Ranunculus abortivus L., little-leaf buttercup (1535)

*Ranunculus acris L., tall buttercup (1685)

Ranunculus fascicularis Muhl., thick-root buttercup (1448)

Ranunculus hispidus Michx. var nitidus (Champ.) T. Duncan, bristly buttercup (1476, 1969)
Ranunculus pensylvanicus L. f., bristly buttercup; Fassett 8265, 1936 (WIS)

Ranunculus recurvatus Poir. var. recurvatus, hooked buttercup (1519)

Thalictrum dasycarpum Fisch. & Ave-Lall., tall meadow-rue (1687)

Thalictrum dioicum L., early meadow-rue (1500)

Thalictrum thalictroides (L.) A. J. Eames & B. Boivin, rue-anemone (1445)

RHAMNACEAE (Buckthorn Family)

Ceanothus americanus L., New Jersey tea (2019)

*Rhamnus cathartica L., common buckthorn (1989)

ROSACEAE (Rose Family)

Agrimonia gryposepala Walk., common agrimony (1826)

Amelanchier laevis Wiegand, Allegheny serviceberry; Pohl 489, 1936 (WIS)

Amelanchier sanguinea (Pursh) DC., New England serviceberry (1463, 1487, 1944, 1951)
Comarum palustre L., marsh cinquefoil; Pohl 504, 1936 (WIS)

Drymocallis arguta (Pursh) Rydb., prairie cinquefoil (1699)

Fragaria vesca L. subsp. americana (Porter) Staudt, thin-leaved wild strawberry (1493)
Fragaria virginiana Mill., wild strawberry (1525)

Geum aleppicum Jacq., yellow avens (1749)

Geum canadense Jacq., white avens (1659)

Geum triflorum Pursh, prairie smoke (1700)

*Malus pumila Mill., apple (1516)

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Physocarpus opulifolius (L.) Maxim., ninebark (1629, 2145)

Potentilla argentea L., silvery cinquefoil (1661)

Potentilla norvegica L., rough cinquefoil; Patman s.n., 1959 (WIS)

*Potentilla recta L., sulpher cinquefoil (1772)

Potentilla simplex Michx., common cinquefoil (1673)

Prunus americana Marshall, wild plum (1947, 1950)

Prunus nigra Aiton., Canada plum (1464)

Prunus pumila L., sand cherry (1483)

Prunus serotina Ehrh., black cherry (1572)

Prunus virginiana L. var. virginiana, chokecherry (1512)

Rosa acicularis Lindl., bristly rose (1632)

Rosa arkansana Porter, prairie rose; Patman s.n., 1959 (WIS)

Rosa blanda Aiton, smooth rose (1602)

Rubus allegheniensis Porter, common blackberry (2187)

Rubus alumnus L. H. Bailey, old field blackberry (1603)

Rubus occidentalis L., black raspberry (1668)

Rubus pubescens Raf., dwarf red raspbeny (1936)

Rubus strigosus Michx., red raspberry (1660)

Rubus superioris L. H. Bailey, Superior blackberry; Pohl 529, 1936 (WIS)

*Sorbus aucuparia L., Eurasian mountain-ash (1552)

Sorbus decora (Sarg.) C. K. Schneid., northern mountain-ash; Pohl 506, 1936 (WIS)

Spiraea alba Du Roi var. alba, white meadowsweet (1805)

RUBIACEAE (Madder Family)

Cephalanthus occidentalis L., buttonbush (1833)

Galium aparine L., cleavers (1505)

Galium asprellum Michx., rough bedstraw (1562)

Galium boreale L., northern bedstraw (1583)

Galium concinnum Torr. & A. Gray, pretty bedstraw (1581)

Galium obtusum Bigelow subsp. obtusum, blunt-leaf bedstraw (1678)

Galium triflorum Michx., sweet-scented bedstraw (1544)

Houstonia longifolia Gaertn., long-leaved bluets (1549, 1664)

Mitchella repens L., partridgeberry (1747, 1974)

RUTACEAE (Rue Family)

Zanthoxylum americanum Mill., prickly ash (1680)

SALICACEAE (Willow Family)

Populus deltoides Bartram. ex Marshall subsp. monilifera (Aiton) Eckenw., cottonwood
(1751)

Populus grandidentata Michx., big-toothed aspen (2128)

Populus tremuloides Michx., quacking aspen (1988)

Salix discolor Muhl., pussy willow (1485, 1486, 1939)

Salix eriocephala Michx., diamond willow (2037)

Salix interior Rowlee, sandbar willow (2068)

Salix petiolaris Sm., slender willow (1484)

* Salix xrubens Schrank, hybrid crack willow (1952)

SANTALACEAE (Sandalwood Family)

Comandra umbellata (L.) Nutt., bastard toadflax (1542)

SAPINDACEAE (Soapberry Family)

*Acer ginnala Maxim., amur maple (2008)

Acer negundo L., boxelder (1949)

Acer rubrum L. var. rubrum, red maple (1959)

Acer saccharinum L., silver maple (2022)

Acer saccharum Marshall var. saccharum, sugar maple (1964)

SAXIFRAGACEAE (Saxifrage Family)

Chrysosplenium americanum Hook., American golden saxifrage (1980)

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Heuchera richardsonii R. Br., prairie alum root (1511)

Micranthes pensylvanica (L.) Haw., swamp saxifrage (1616)

Mitella diphylla L., two-leaved miterwort (1470)

SCROPHULARIACEAE (Figwort Family)

Scrophularia lanceolata Pursh, American figwort (1631)

*Verbascum thapsus L., common mullein (2031)

SOLANACEAE (Nightshade Family)

Solarium dulcamara L., bittersweet nightshade (1716)

Solarium ptycanthum Dunal, black nightshade (1796)

STAPHYLEACEAE (Bladdemut Family)

Staphylea trifolia L., American bladdemut (1945)

THYMELAEACEAE (Mezereum Family)

Dirca palustris L., leatherwood (1942)

ULMACEAE (Elm Family)

Ulmus americana L., American elm (1943)

*Ulmuspumila L., Siberian elm (1946)

Ulmus rubra Muhl., red elm (2023)

Ulmus thomasii Sarg., rock elm (1993)

URTICACEAE (Nettle Family)

Boehmeria cylindrica (L.) Sw., small spike false nettle (1777, 1833)

Laportea canadensis (L.) Wedd., wood nettle (1875)

Parietaria pensylvanica Muhl. ex Willd., Pennsylvania pellitory (1608)

Pilea pumila (L.) A. Gray, clearweed (1864)

Urtica dioica L. subsp. gracilis (Aiton) Selander, stinging nettle (1865)

VERBENACEAE (Vervain Family)

Verbena hastata L., blue vervain (1806)

Verbena urticifolia L., white vervain (1849)

VIOLACEAE (Violet Family)

Viola labradorica Schrank, American dog violet (1461)

Viola macloskeyi F. E. Lloyd subsp. pallens (DC.) M. S. Baker, small white violet (1454,
1956)

Viola pedatifida G. Don, prairie violet (2143)

Viola pubescens Aiton, yellow forest violet (1491)

Viola sagittata Aiton, arrow-leaved violet (1541)

Viola sororia Willd., door-yard violet (1475, 1478)

VITACEAE (Grape Family)

Parthenocissus inserta (A. Kem.) Fritsch, grape woodbine (2234)

Vitis riparia Michx., riverbank grape, frost grape (2055)

MONOCOTYLEDONS

ACORACEAE (Sweet Flag Family)

Acorus americanus (Raf.) Raf., sweet flag; Pohl 490, 1936 (WIS)

ALISMATACEAE (Water-Plantain Family)

Alisma triviale Pursh, northern water-plantain (2039)

Sagittaria latifolia Willd., broad-leaved arrow-head (1902)

Sagittaria rigida Pursh, stiff arrow-head (1808, 2241)

ALLIACEAE (Onion Family)

Allium canadense L., wild onion (1579)

Allium stellatum Ker Gawl., prairie onion (1811)

Allium tricoccum Aiton, wild leek (1521)

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ARACEAE (Arum Family)

Arisaema triphyllum (L.) Schott subsp. triphyllum, Jack-in-the-pulpit (1561)

Calla palustris L., wild calla (1727)

Lemna trisulca L., forked duckweed (1957)

Lemna turionifera Landolt, Turion duckweed (1905)

Spirodela polyrrhiza (L.) Shleid., greater duckweed (1906)

Symplocarpus foetidus (L.) W. P. C. Barton, skunk cabbage (1444)

ASPARAGACEAE (Asparagus Family)

*Asparagns officinalis L., asparagus (1577)

COMMELINACEAE (Spiderwort Family)

Tradescantia occidentalis (Britton) Smyth var. occidentalis, prairie spiderwort (1773)

CONVALLARIACEAE (Lily-of-the-valley Family)

Clintonia borealis (Aiton) Raf., blue-bead lily (1588)

*Convallaria majalis L. var. majalis, European lily-of-the-valley (1979)

Maianthemum canadense Desf., wild lily-of-the-valley (1537)

Maianthemum racemosum (L.) Link, false Solomon’s-seal (1587)

Polygonatum biflorum (Walter) Elliott, giant Solomon’s-seal (1638)

Polygonatum pubescens (Willd.) Pursh, hairy Solomon’s-seal (1790)

Uvularia grandiflora Sm., bellwort (1490)

Uvularia sessilifolia L., sessile bellwort (1467)

CYPERACEAE (Sedge Family)

Bolboschoenus fluviatilis (Torr.) Sojak, river bulrush (1900)

Bulbostylis capillaris (L.) C. B. Clarke, hair sedge; Fassett 17705, 1934 (WIS)

Carex assiniboinensis W. Boott, Assiniboine sedge (1551)

Car ex backii W. Boott, Rocky Mountain sedge (1976)

Carex bicknellii Britton, Bicknell’s oval sedge (1636)

Carex blanda Dewey, common wood sedge (1499, 1545)

Carex brevior (Dewey) Mack. Ex Lunell, fescue sedge (1994)

Carex bromoides Willd., brome-like sedge (1453)

Carex buxbanmii Wahlenb., Buxbaum’s sedge (1548)

Carex cephalophora Willd., oval-headed sedge; Fassett 7382, 1927 (WIS)

Carex crinita Lam. var. crinita, fringed sedge (1622)

Carex cristatella Britton, crested oval-sedge (1737)

Carex deweyana Schwein., Dewey’s sedge (1571)

Carex emoryi Dewey, Emory’s sedge; Pohl 484, 1936 (WIS)

Carex gracillima Schwein., graceful sedge (1506)

Carex haydenii Dewey, Hayden’s sedge (1568, 2146)

Carex hirtifolia Mack., hairy sedge (1982)

Carex intumescens Rudge, greater bladder sedge (1574)

Carex lacustris Willd., lake sedge (2010)

Carex lupulina Willd., common hop sedge (804, 1732)

Carex lurida Wahlenb., shallow sedge (1769)

Carex muhlenbergii Willd., Muhlenberg’s sedge (2005)

Carex normalis Mack., greater straw sedge (1567)

Carex pedunculata Willd., long-stalk sedge (1455, 1965)

Carexpellita Willd., broad-leaved woolly sedge (1600, 1708)

Carex pensylvanica Lam., Pennsylvania sedge (1450)

Carex projecta Mack., loose-headed oval sedge (1569, 1701)

Carex radiata (Wahlenb.) Small, eastern star sedge (1507)

Carex retrorsa Schwein., deflexed bottlebrush sedge (524)

Carex sprengelii Spreng., Sprengel’s sedge (1503)

Carex stipata Willd. var. stipata, common fox sedge (1597)

Carex stricta Lam., tussock sedge (2147)

Carex tribuloides Wahlenb., awl-fruited oval sedge (1782)

Carex tuckermanii Dewey, Tuckerman’s sedge (1705, 1706)

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Carex typhina Michx., cattail sedge (1733, 1734, 1783, 2046)

Carex umbellata Schkuhr. ex Willd., early oak sedge (1960)

Carex vesicaria L., blister sedge (1736)

Carex vulpinoidea Michx., brown fox sedge (1735)

Cyperus bipartitus Torn, slender flat sedge (1847, 1926)

Cyperus diandrus Torn, umbrella flat sedge; Pohl 598, 1936 (WIS)

Cyperus erythrorhizos Muhl., red-rooted sedge (2124)

Cyperus esculentus L., yellow nut sedge (1846)

Cyperus odoratus L., flat sedge (802)

Cyperus schweinitzii Tom, Great Plains sand sedge (1794, 1857, 1894)

Cyperus squarrosus L., bearded flat sedge (2246)

Cyperus strigosus L., straw colored cyperus (1925)

Eleocharis acicularis (L.) Roem. & Schult., needle spike-rush (2123)

Eleocharis erythropoda Steud., bald spike-rush (2122)

Eleocharis intermedia Schult., matted spike-rush (2095)

Eleocharis obtusa (Willd.) Schult., blunt spike-rush (2016, 2120, 2243)

Eleocharis ovata (Roth) Roem. & Schult., oval spike-rush (2119)

Schoenoplectus tabernaemontani (C. C. Gmel.) Palla, soft stem bulrush (1807)

Scirpus atrovirens Willd., black bulrush (1765, 1845)

Scirpus cyperinus (L.) Kunth, wool-grass (2041)

Scirpus microcarpus J. Presl. & C. Presl, panicled bulrush (1644, 1645)

Scirpus pedicellatus Femald, stalked wool-grass (1786)

DIOSCOREACEAE (Yam Family)

Dioscorea villosa L., wild-yam (1691, 1789)

HEMEROCALLIDACEAE (Day-lily Family)

*Hemerocallis fulva (L.) L., orange day-lily (2014)

HYDROCHARITACEAE (Frog’s-Bit Family)

Elodea canadensis Michx., common waterweed (1802)

Elodea nuttallii (Planch.) St. John, slender waterweed (1909)

Najas flexilis (Willd.) Rostk. Schmidt, northern water-nymph (2240)

Najas guadalupensis (Spreng.) Magnus, southern water-nymph (1800)

Vallisneria americana Michx., American eel-grass (2058)

HYPOXIDACEAE (Star-grass Family)

Hypoxis hirsuta (L.) Coville, yellow star-grass (1698, 2144)

IRIDACEAE (Iris Family)

*Iris pseudacorus L., yellow flag (1676)

Iris virginica L., var. shrevei (Small) E. S. Anderson, southern blue flag (1606, 2011)
Sisyrinchium campestre E. P. Bicknell, prairie blue-eyed grass (1526)

JUNCACEAE (Rush Family)

Juncus effusus L., common rush; Pohl 497, 1936 (WIS)

Juncus tenuis Willd., path rush (1665, 2024)

Luzula acuminata Raf. var. acuminata, hairy wood rush (1462)

Luzida multiflora (Ehrh.) Lej. subsp. multiflora, common wood rush (1502)

LILIACEAE (Lily Family)

Erythronium albidum Nutt., white trout lily (1459)

Lilium michiganense Farw., Michigan lily (1761)

ORCHIDACEAE (Orchid Family)

Cypripedium acaule Aiton, moccasin flower (1692)

Cypripedium parviflorum Salib. var. pubescens (Willd.) O. W. Knight, large yellow lady’s
slipper (1971)

Galearis spectabilis (L.) Raf., showy orchis (1557)

Goody era pubescens (Willd.) R. Br., downy rattlesnake-plantain (1430)

Liparis liliifolia (L.) Rich, ex Lindl., lily-leaved twayblade (1771, 2188)

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Platanthera psycodes (L.) Lindl., purple fringed orchid (520)

POACEAE (Grass Family)

Agrostis perennans (Walter) Tuck. var. perennans, autumn bent grass (2034)

Agrostis scabra Willd., tickle grass (1730, 1731)

Andropogon gerardii Vitman, big bluestem grass (1816)

Brachyelytrum aristosum (Michx.) P. Beauv. Ex Branner & Coville, northern shorthusk
(1674)

Brachyelytrum erectum (Schreb.) P. Beauv., long-awned wood grass (1432)

Bromus ciliatus L., fringed brome (534)

*Bromus inermis Leyss., smooth brome (1697, 2002)

Bromus latiglumis (Shear) Hitchc., ear-leaved brome; Skinners 2858, 1940 (WIS)

Bromus pubescens L., hairy woodland brome (1696, 1810)

Calamagrostis canadensis (Michx.) P. Beauv., blue-joint grass (1832)

Cenchrus longispinus (Hack.) Femald, sandbur (1848)

Cinna arundinacea L., common wood reed (2135, 2136, 2227)

Cinna latifolia (Gopp.) Griseb., drooping wood reed; Fassett 5392, 1929 (WIS)

*Dactylis glomerata L., orchard grass (1626)

Danthonia spicata (L.) Roem. & Schult., poverty oatgrass (527, 1643, 1858)

Dichanthelium acuminatum (Sw.) Gould & C. A. Clark subsp. fasciculatum (Torr.) Freck-
mann & LeLong, western panic grass (1711)

Dichanthelium boreale (Nash) Freckmann, northern panic grass; Frecbnann 4324, 1996
(UWSP)

Dichanthelium latifolium (F.) Harvill., broad-leaved panic grass (1656)

Dichanthelium leibergii (Vasey) Scribn., Feiberg’s panic grass; Cochrane 5332, 1927 (WIS)
Dichanthelium linearifolium (Scribn.) Gould, linear-leaved panic grass (1712)
Dichanthelium oligosanthes (Schult.) Gould subsp. scribnerianum (Nash) Freckman & Fe-
Long (1634)

Dichanthelium perlongum (Nash) Freckmann, long-stalked panic grass (1657)
Dichanthelium xanthophysum (A. Gray) Freckmann, pale panic grass; Freckmann 4322,
1996 (UWSP)

*Digitaria ischaemum (Schreb.) Muhl., smooth crabgrass (2073)

Echinochloa muricata (P. Beauv.) Femald, barnyard grass (537, 2057)

Echinochloa walteri (Pursh) A. Heller, coast barnyard grass (2108, 2117)

Elymus canadensis F., Canada wild-rye (1889)

Elymus hystrix F., bottlebrush-grass (521)

* Elymus repens (F.) Gould, quackgrass (1717)

Elymus villosus Muhl., downy wild-rye (1819, 1852)

Elymus virginicus F. var. virginicus, Virginia wild-rye (1426)

Eragrostis capillaris (F.) Nees, lace grass; Fassett 5472, 1927 (WIS)

Eragrostis cilianensis (All.) Vignolo ex Janch., sti nk grass (2096)

Eragrostis hypnoides (Earn.) Britton, Stems & Poggenb., creeping love grass (2129)
Eragrostis pectinacea (Michx.) Nees var. pectinacea, low love grass (2056)

Festuca subverticillata (Pers.) E. B. Alexeev, nodding fescue (526, 528, 1729)

Glyceria canadensis (Michx.) Trin., rattlesnake mannagrass (2060)

Glyceria grandis S. Watson, American mana grass; Patman s.n., 1959 (WIS)

Glyceria striata (Earn.) Hitchc., fowl mannagrass (529, 1675, 1724, 1725)

Hesperostipa spartea (Trin.) Barkworth, porcupine grass (1999)

Hierochloe hirta (Schrank) Borbas, hairy sweetgrass (1981)

Koeleria macrantha (Ledeb.) Schult., Junegrass (1642)

Leersia oryzoides (F.) Sw., rice cut grass (2054)

Leersia virginica Willd., white grass (1838, 1892)

Milium effusum F., American millet grass (1566, 1695)

Muhlenbergia frondosa (Poir.) Fern., common satin grass; Shinners 2856, 1940 (WIS)
Muhlenbergia mexicana (F.) Trin., leaft satin grass; Shinners 2854, 1940 (WIS)

Oryzopsis asperifolia Michx., rough-leaved rice grass (1456, 1489, 1966)

Panicum capillare F., common witch grass (2079, 2121)

Panicum philadelphicum Trin., Philadelphia panic grass (2109)

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THE MICHIGAN BOTANIST

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Panicum virgatum L., switchgrass (532, 535, 536)

Phalaris arundinacea L., reed canary grass (1996)

*Phleum pratense L., subsp. pratense, timothy (1788)

Phragmites australis (Cav.) Trin. Ex Steud., common reed grass (2063)
Piptatherum racemosum (Sm.) Eaton, black-seeded rice grass (2094)

*Poa annua L., annual bluegrass (2007)

*Poa compressa L., Canada bluegrass (2003)

Poapalustris L., marsh bluegrass (1529, 1722, 1723)

*Poa pratensis L., Kentucky bluegrass (1550)

Schizachne purpurascens (Torr.) Swallen, false melic (1482)

Schizachyrium scoparium (Michx.) Nash, little bluestem (1930)

*Setaria faberi R. A. W. Herrm., giant foxtail (2078)

*Setaria pumila (Poir.) Roem. & Schult. subsp. pumila, yellow foxtail (2038)
*Setaria viridis (L.) P. Beauv., green foxtail (1793)

Sorghastrum nutans (L.), Nash., Indian grass (1433)

Spartina pectinata Link, prairie cord-grass (530)

Sporobolus heterolepis (A. Gray) A. Gray, prairie drop seed (2017, 2126)

POTAMOGETONACEAE (Pondweed Family)

Potamogeton crispus L., curly pondweed (1801)

Potamogeton nodosus Poir., long-leaf pondweed (1904)

Potamogeton pusillus L., slender pondweed (1799, 1907)

Zannichelliapalustris L., homed-pondweed (2238, 2239)

SMILACACEAE (Catbrier Family)

Smilax ecirrhata S. Watson, upright carrion flower (1558, 1968, 1975)
Smilax hispida Raf., bristly greenbriar (1689)

Smilax lasioneura Hook., common carrion flower (1985)

TRILLIACEAE (Trillium Family)

Trillium cernuum L., nodding trillium (1589)

Trillium flexipes Raf., declined trillium (1466)

Trillium grandiflorum (Michx.) Salisb., big white trillium (1465)

TYPHACEAE (Cat-Tail Family)

Sparganium eurycarpum Englm., common bur-reed; Pohl 527, 1936 (WIS)
Typha angustifolia L., narrow-leaved cat-tail (1739)

Typha latifolia L., broad-leaved cat-tail (1759)

Typha xglauca Godr., hybrid cat-tail (1740)

2014

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39

COMPARATIVE ANALYSIS OF URBAN AND RURAL FOREST
FRAGMENT STRUCTURE AND DIVERSITY IN
NORTHEASTERN INDIANA

Kristine D. Arvola, Parker F. Booth, Charles C. Ellinwood, Angela J. Fry,
Jason L. Furge, Kaitlyn A. Haehnle, Lauren E. Hall, Melanie A. Jablonski,
Daniel K. Jones, Justin T. Martin, Kevin M. McLane, Kevan C. Mensch,
Victoria A. Mumaw, Rebeca N. Quirindongo, Michael J. Ravesi,

Jesse J. Rinard, Patrick R. Selig, Andrew P. Sellan, Maja B. Sljivar,
Emily A. Stulik, Tasha R. Sunday, Alison N. Turley, Mark T. Voors,
Adam R. Warrix, Tyler C. Wood, and Jordan M. Marshall 1
Department of Biology, Indiana University-Purdue University
Fort Wayne, Fort Wayne, IN 46805

ABSTRACT

Although it was once continuously forested, the land cover in northeastern Indiana is now dom¬
inated by agriculture, and sparsely occurring forest fragments now constitute only approximately 8%
of the land cover. A majority of these forest fragments are privately owned and have a history of
some form of active forest management. We conducted a systematic ecological survey of understory,
midstory, and overstory plant species in three forests that have differing protection and management
histories to compare the effects of these different histories. Historical aerial images of each forest
were compared to gauge the canopy structure and to clarify the management history for the forests.
The percentage of canopy cover and the floristic quality indices (FQI) each followed expected
trends, whereby the highest FQI value and percentage of canopy cover occurred in the forest with the
longest history of preservation. Lower values of species richness for the understory, midstory, and
overstory strata, respectively, were found in the forest that has a history of overstory management
and for which there is no defined protection status. The understory species were each generally lim¬
ited to one of the forests, whereas the species composition of the midstory and overstory strata were
much more similar among the three forests. Measurements of forest basal area and percentage of
canopy cover provide some explanation of the distribution of understory and midstory species in
nonmetric multidimensional scaling ordination plots. The amount of forest protection, measured by
the time since disturbance and the percentage of canopy closure, influenced the ric hn ess of the un¬
derstory and the FQI of a given plot. Furthermore, the location of a forest was an important factor in
the relative occurrence of non-native species, the most rural forest having no non-native species.

KEYWORDS: Fogwell Forest Nature Preserve, Mengerson Nature Preserve, fragmentation,
floristic quality index, forest management

INTRODUCTION

According to the National Land Cover Database (Fry et al. 2011), the Mid¬
west region of the United States is dominated by agriculture with over 60% of
land cover in cultivated crops, pasture land, or other open-field agricultural prac¬
tices; forests account for only 20% of the land cover. Urban and suburban de¬
velopment has increased fairly continuously in the region for well over 60 years

1 Author for correspondence (marshalj@ipfw.edu)

40

THE MICHIGAN BOTANIST

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(Radeloff et al. 2005). Because there are large areas of contiguous forested land
in northern Michigan, Minnesota, and Wisconsin, the percentage of land cover
devoted to agricultural uses in the Midwest as a whole understates the situation
in the southern part of the region, where cultivated and pastoral agricultural
lands account for 80-90% of the land cover (Radeloff et al. 2005) and where
forests are fragmented into relatively small woodlots. These woodlots generally
have high edge-to-interior ratios, thereby increasing the area of forest subject to
influence by the surrounding land matrix, which is typically agricultural, but also
includes developed urban and suburban land (Brothers and Spingam 1992; Gon¬
zalez et al. 2010). Urbanization results in changes to the understory of forest
fragments, shifting community composition away from native species toward
non-native species, as well as potentially homogenizing previously distinct com¬
munities (Kuhn and Klotz 2006; Dolan et al. 2011).

Much of the Central Till Plain region of Indiana was forested prior to European
settlement and was dominated by flatwoods (Hedge 1997). This physiographic re¬
gion covers much of the central and northeastern portions of Indiana with the
southern boundary delineated by the southern reaches of the Wisconsinan ice sheet
(Hedge 1997). As a result of extensive agricultural and urban development, forests
now account for only 20.3% of the land cover in the entire state, the majority of
which is privately owned (Woodall et al. 2009). Forest conservation efforts in In¬
diana have increased steadily with the inclusion of private forests in classified for¬
est and cooperative forest management programs (IDNR 2010).

Understory and midstory plant communities are directly influenced by the
structure and composition of the overstory community (Jameson 1967; Roberts
1992). Changes in the overstory by anthropogenic manipulation will alter those
lower strata, which could be positive or negative depending on the community
and the manipulation (Meier et al. 1995; Albrecht and McCarthy 2006). Addi¬
tionally, isolation of forest fragments from other forests alters understory com¬
position, negatively affecting a large proportion of species (McKinney and
Lockwood 1999). Fragmentation impacts are long term and may persist on local
or regional scales (Vellend et al. 2006). Land managers are therefore interested
in quantifying the floristic integrity of a given plant community in light of forest
fragmentation and anthropogenically induced disturbances (Rothrock 2004).
Floristic quality assessments, as defined by Swink and Wilhelm (1994), have
been applied to several different ecosystem types and, within a single ecosystem
type, to those under different management strategies (e.g., Francis et al. 2000;
Lopez and Fennessy 2002, Bacone et al. 2007; Rothrock et al. 2011). The need
to understand the influence of forest protection and preservation on the floristic
structure and composition of forest fragments is the principal reason we have un¬
dertaken this study.

The objectives of this study were 1) to quantify the understory, midstory, and
overstory plant communities in three forests in northeastern Indiana with respect
to species richness, diversity, and evenness, 2) to relate the composition and di¬
versity of plant co mm unities to characteristics of forest structure, and 3) to test
the hypothesis that the management history and the characteristics of the sur¬
rounding land matrix will influence the species composition of the understory
and midstory of these three forest fragments.

2014

THE MICHIGAN BOTANIST

41

Legend

Allen County

I I Fort Wayne City Boundary
Forest Location
O Plot Location

FIGURE l.A. Study forest locations within Allen County,
Indiana. B-D. Plot locations for Fogwell Forest Nature Pre¬
serve (B), Indiana University-Purdue University Fort Wayne
Forest (C), and Mengerson Forest Nature Preserve (D).
White lines indicate property boundaries.

MATERIALS AND METHODS

Study Sites

Three forests in Allen County, Indiana, were selected for comparison (Figure 1A). Two of them,
Fogwell Forest Nature Preserve (Fogwell) (40°59'50" N, 85°14'37" W; Figure IB) and Mengerson
Nature Preserve (Mengerson) (41°7'35" N, 85°4'4" W; Figure ID), are owned and managed by
ACRES Land Trust and are designated nature preserves by the Indiana Department of Natural Re¬
sources. The third forest (IPFW) (41°7'22" N, 85°7'18" W; Figure 1C) is owned and managed by In¬
diana University-Purdue University Fort Wayne. IPFW and Mengerson are in the Auburn Morainal

42

THE MICHIGAN BOTANIST

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Complex physiographic division, and Fogwell is in the Bluffton Till Plain physiographic division
(Franzmeier et al. 2004). All three forests are located adjacent to the boundary between these two
physiographic divisions, which are contained in the broader Central Till Plain Natural Region, and
all three are dominated by Blount-Morley silt loam soils (NRCS 2013).

Fogwell Forest Nature Preserve is a 24.8 ha property, approximately 12.3 ha of which has a con¬
tinuous forest canopy (ACRES 2008). A small housing subdivision lies to the north of Fogwell, and
the remaining adjacent land consists of cultivated agricultural land and privately-owned hardwood
forest. Rothrock (1997) described the forested portion of Fogwell as being donated to ACRES Land
Trust in 1976, prior to which the land had been designated a classified forest in the 1930s, which
placed limits on the removal of trees from the forest. The Classified Forest program in Indiana is
managed by the Department of Natural Resources Division of Forestry and provides protection of
privately-owned forests against large anthropogenic disturbances, while providing the land owner
with a reduced tax assessment (IDNR 2015). Because of this designation, Fogwell has remained a
mature closed canopy forest since the 1930s (Figure 2A).

IPFW, a forested tract of land approximately 13.8 ha in area that is adjacent to the university cam¬
pus, was acquired by the university in 2004. Little documentation is available regarding its manage¬
ment history, but historical aerial photos indicate that the property has been continuously forested since
the late 1930s (Figure 2B). However, the canopy does not appear to have been as dense or as com¬
pletely closed as the canopy at Fogwell. IPFW is bounded by residential neighborhoods to the north and
west, commercial properties to the west and south, and intensely managed athletic fields to the east.

Mengerson Nature Preserve is an approximately 14.4 ha forest that was donated to ACRES Land
Trust in 1973 (ACRES 2008). Unlike Fogwell and IPFW, Mengerson was forested only in the north¬
ern 1/3 during the 1930s (Figure 2C). Over the subsequent decades, several species of trees have col¬
onized the southern 2/3 of Mengerson, resulting in an early successional forest. Even now, however,
the southern 1/3 of Mengerson is only sparsely forested (Figure ID).

Methods

Within each forest, an initial grid of points with 25.25 m spacing was overlaid on aerial images,
and 20 grid points were randomly selected to serve as plot center locations, using ArcMap (version
9.3.1, ESRI, Redlands, California). The spacing of the initial grid points was selected in a manner to
ensure that adjacent overstory plots (defined below) did not overlap. At each plot center, understory,
midstory, and overstory survey plots were established. Each understory plot consisted of two 1 m 2
quadrats diagonal from each other with sides parallel to the cardinal directions (the southwest comer
of one quadrat and the northeast comer of the other were each at the plot center). Within the under¬
story survey plots, all individual plants less than 2 m in height were identified to the finest taxonomic
level possible (typically species) and counted. Each midstory survey plot consisted of a 5 * 5 m
square plot with the sides parallel to the cardinal directions. All individual stems greater than 2 m in
height and less than 8 cm diameter at breast height (dbh) were identified to species and counted.
Overstory survey plots consisted of 500 m 2 circular plots (12.62 m radius). All overstory stems (i.e.,
those greater than 8 cm dbh) were measured for dbh, identified to species, and counted. Voucher
specimens for each taxon identified in the understory plots were deposited in the Indiana University-
Purdue University Fort Wayne Department of Biology herbarium. All surveys and data collection
were conducted during September and October 2013.

At each plot center, basal area per species was assessed with a basal area 10-factor prism.
Canopy cover was measured using a concave spherical densiometer following standard protocols
(i.e. taking measurements 1 m above ground, averaging measurements taken facing the four cardinal
directions). Litter depth was also measured in each plot at the plot center and at a point 6.3 m from
the center toward each of the four midstory plot comers. Measurements of canopy cover and litter
depth were compared between forests using analysis of variance (ANOVA) with a Tukey-Kramer
post-hoc test. Relationships between litter depth, total overstory density (stems / 500 m 2 plot), and
total midstory density (stems / 25 m 2 plot) were tested with a Spearman-Rank correlation (due to vi¬
olations of normality assumptions).

Species richness (S = number of species), Shannon’s diversity index (H’ = -2 p\ In p x , where p t
is the proportion of the rth species), and Pielou’s evenness index (J' = H' / In S), were calculated for
each strata at the plot level. These three measures were then compared among the forests using
ANOVA with a Tukey-Kramer post-hoc test. Using the coefficient of conservatism for the under¬
story species listed by Rothrock (2004), we calculated an unweighted mean coefficient of conser-

2014

THE MICHIGAN BOTANIST

43

FIGURE 2. Aerial images from 1938 for (A) Fogwell (MAPI 2013a), (B) IPFW (MAPI 2013b), and
(C) Mengerson (IHAPI 2013c) forests. White forest boundaries are from the 2013 field study. Geo-
rectification complications resulted in boundary errors on the eastern edges of Fogwell and IPFW.

44

THE MICHIGAN BOTANIST

Vol. 53

vatism (C mean ) for each forest and the floristic quality index (FQI = C mean * Vn umber of native plant
species) for each forest. Coefficient of conservatism (C-value) is a numerical value used to describe
the “nativeness” of a plant species in relation to anthropogenic disturbance (Swink and Wilhelm
1994, Rothrock 2004); the greater the C-value assigned to a species, the more likely it is associated
with remnant habitats similar to those existing prior to European settlement (range 1-10, 0 for non¬
native species). FQI provides a calculated value for the quality and natural importance of a plant
community based on the C-values assigned to the species within that community (Rothrock 2004).
Similarities among the forests in the understory and midstoiy survey plots were visualized with non¬
metric multidimensional scaling (NMDS) ordination using methods described by Kruskal (1964a,b).
Dissimilarities were characterized using Bray-Curtis distances, which allow for a visual representa¬
tion of dissimilarity in species abundances. Within the NMDS figure, species plotted closer together
are less dissimilar than those farther apart. For both understory and midstoiy NMDS ordinations, re¬
lationships between species and forest basal area, percent canopy cover, and litter depth were visu¬
alized with vector plots associated with the ordination (cutoff a = 0.1, iterations = 1000). We se¬
lected our cutoff for the vectors in order to increase the likelihood of displaying basal area, canopy
cover, and litter depth on the NMDS ordination plot. Importance values for each overstory species in
each forest were calculated as the sum of the relative frequency, the relative dominance, and the rel¬
ative density of that species; where relative frequency = number of plots in which a species occurred
/ total number of plots * 100, relative dominance = basal area of a species / total forest basal area *
100, and relative density = number of stems of a species / total number of stems * 100. All statisti¬
cal analyses were conducted using R (version 3.0.2, The R Foundation for Statistical Computing)
base and vegan packages.

RESULTS

In both IPFW and Mengerson, all 20 plot locations were surveyed. However,
due to time constraints, only 17 plots at Fogwell were surveyed. In Mengerson,
six plots were outside of the north property boundary at the time of study, but
were in a section of the forest in the process of being acquired by ACRES Land
Trust (Figure ID). Canopy cover was significantly greater in Fogwell than in the
other two forests (F = 5.77, df = 2,54, P = 0.005; Figure 3A). Litter depth at

Forest Forest

FIGURE 3. Percentage of canopy cover (A) and litter depth (B) in Fogwell, IPFW, and Mengerson
forests. Bars that do not share the same letter are significantly different. Error bars represent one
standard deviation about the mean.

2014

THE MICHIGAN BOTANIST

45

Mengerson was significantly deeper than at the other two forests (F = 9.76, df =
2,54, P< 0.001; Figure 3B).

Understory

A total of 75 species was encountered within the understory strata of the sur¬
veyed forests (Appendix I). Fogwell and IPFW were significantly different in
species ric hn ess (F = 3.82, df = 2,54, P = 0.028) (Figure 4A). However, the three
forests were not significantly different in understory diversity (F = 2.67, df = 2,53,
P = 0.079) (Figure 4B), which is likely related to a lack of difference in eve nn ess
between forests (F = 1.10, df = 2,46, P = 0.341). Similarly, C mean was not signifi¬
cantly different between the forests (F = 0.94, df = 2,84, P = 0.396) (Table 1).
However, Fogwell had both the greatest percentage of species with a coefficient of
conservatism > 5 and the highest FQI value (Table 1). The forest understories had
limited overlap in species composition, sharing only 14.6% of species between
Fogwell and IPFW, 14.3% between IPFW and Mengerson, and 12.5% between
Fogwell and Mengerson. The limited overlap in species was visually evident in the
NMDS ordination analysis (Figure 5A). The basal area, the canopy cover, and the
litter depth each met the cutoff for inclusion as vectors (Figure 5B).

Midstory

A total of 16 species was encountered within the midstory strata of the sur¬
veyed forests (Appendix II). IPFW and Mengerson differed significantly in both
species richness (F = 10.23, df = 2,54, P < 0.001) (Figure 4C) and in diversity (F
= 5.03, df = 2,49, P = 0.010) (Figure 4D). Species evenness was not significantly
different between the forests (F = 3.07, df = 2,48, P = 0.056). Midstory density
(stems / 25 m 2 plot) was significantly greater in Mengerson, with 4 and 3 times
more stems per plot than in Fogwell and IPFW, respectively (F = 20.24, df =
2,54, P < 0.001). Furthermore, midstory density and litter depth were positively
correlated (r = 57.00, P = 0.003). However, litter depth was not correlated with
either midstory species richness (r = 0.07, P = 0.611) or diversity (r = 0.06, P =
0.635). Unlike the understory strata, the midstory strata in the three forests were
fairly similar in species composition, sharing 40.0% of species between Fogwell
and IPFW, 31.3% between IPFW and Mengerson, and 35.7% between Fogwell
and Mengerson. The similarity was visually evident in the NMDS ordination
analysis (Figure 6A). Vector angle indicates relative direction of influence and
vector length indicates relative strength of the influence. For example, Acer sac-
charum (ACSA3) and Ostrya virginiana (OSVI), both shade-tolerant species, in¬
creased in abundance as the percentage of canopy cover increased. Similarly, the
shade-intolerant Crataegus mollis (CRM02) decreased in abundance as the per¬
centage of canopy cover increased (Figure 6B).

Overstory

A total of 34 species was encountered within the overstory strata of the sur¬
veyed forests (Appendix III). The relationship of the species richness of the

46

THE MICHIGAN BOTANIST

Vol. 53

Forest

Forest

FIGURE 4. Species richness and species diversity for understory (A, B), midstory (C, D), and over¬
story (E, F) strata plants in Fogwell, IPFW, and Mengerson forests. Bars that do not share the same
letter are significantly different. Error bars represent one standard deviation about the mean.

2014

THE MICHIGAN BOTANIST

47

TABLE 1. For the understory stratum in each forest studied, the mean coefficient of conservatism
(Cmean) (standard error in parentheses), the count of species for which the coefficient of conservatism
(C) is > 5 (the percentage of those species relative to all species in the midstory stratum of the forest
is given in parentheses), and the floristic quality index (FQI).

Forest

Cmean (standard error)

Count for C > 5 (percentage)

FQI

Fogwell

4.1 (0.4)

13 (41.9%)

23.4

IPFW

3.4 (0.5)

8 (28.6%)

18.6

Mengerson

3.6 (0.4)

7 (25.0%)

18.9

overstory strata of the three forests followed a pattern similar to that observed in
the midstory strata. IPFW and Mengerson differed significantly from each other
(F = 3.20, df = 2,54, P = 0.049) (Figure 4E). However, the overstory diversity of
IPFW was significantly lower than that of Fogwell and Mengerson (F = 6.09, df
= 2,54, P = 0.004) (Figure 4F). As a result, IPFW had the lowest evenness value
of the three forests (F = 7.18, df = 2,54, P = 0.002). At IPFW, A. saccharum ac¬
counted for 62.1% of the overstory individuals. Overstory densities were not sig¬
nificantly different between the three forests, with an overall mean of 19.6 stems
/ 500 m 2 (± 8.3 stems) (F = 0.68, df = 2,54, P = 0.509). Litter depth was not cor¬
related with overstory density (r = -0.07, P = 0.588), species richness (r = 0.07,
P = 0.644), or diversity (r = 0.04, P = 0.784). As with the midstory, the overstory
strata of the three forests were more similar in species composition than were the
understory strata, sharing 35.7% of species between Fogwell and IPFW, 37.5%
between IPFW and Mengerson, and 58.6% between Fogwell and Mengerson. A
strong visual overlap can be seen in the NMDS ordination analysis between
Mengerson and the other two forests, but less so between Fogwell and IPFW
(Figure 7A). Basal area and percentage of canopy cover vectors were not in¬
cluded in the NMDS ordination analysis of pooled species due to collinearity
(Figure 7B).

Acer saccharum was the most important overstory species in all three forests
(Table 2). It had the greatest values for density of stems, for frequency, and for
basal area. There was some overlap in the top five overstory species. Tilia amer-
icana occurred in all three forests, Quercus rubra in Fogwell and Mengerson,
and Ulmus americana in IPFW and Mengerson (Table 2). Some species occu¬
pied high ranking positions in only one of the three forests. For example A. sac-
charinum was among the top five in Mengerson (#3), but did not occur at all in
the other two forests; Carya ovata (#2) and Fagus grandifolia (#3) were both in
the top five in Fogwell, but were less important in the other two forests; and
Juglans nigra (#5) and Ulmus rubra (#4) were much more important in IPFW
than in the other two forests (Table 2).

DISCUSSION

The three selected forests are located within close proximity of each other
(each is approximately 4-16 km from the other two) and, prior to the large-scale

48

THE MICHIGAN BOTANIST

Vol. 53

AXIS 1

CN

(/>

X.

<

B

BIFR

PLMA2+
QXST LEVU

AMAR3

+ LEVI2 FRPE +

GLST +

LIBE3

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po g§AL3 + ROfijjGEVE QURU

ClAR2*CACR9

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AXIS 1

FIGURE 5. Nonmetric multidimensional scaling (NMDS) ordination of under¬
story (final stress = 0.12) for (A) survey plots for Fogwell (squares), IPFW (tri¬
angles), and Mengerson (circles) with 95% confidence ellipses for each forest;
and (B) pooled species. Direction and length of vectors (gray) for basal area
(BA), percentage canopy cover (Canopy), and litter depth (Litter) indicate in¬
fluence on species occurrence. Species letter codes follow USDA (2014).

conversion of forest to cultivated agricultural land and urban development, were
likely connected by contiguous forested land. There have been distinct differ¬
ences in the protection and management regimes of these forests. Fogwell has
received protection for over 80 years, while IPFW and Mengerson have received
protection for only 10 years and 40 years, respectively. IPFW has undergone

2014

THE MICHIGAN BOTANIST

49

AXIS 1

AXIS 1

FIGURE 6. Nonmetric multidimensional scaling (NMDS) ordination of mid¬
story (final stress = 0.08) for (A) survey plots for Fogwell (squares), IPFW (tri¬
angles), and Mengerson (circles) with 95% confidence ellipses for each forest;
and (B) pooled species. Direction and length of vectors (gray) for basal area
(BA) and percentage canopy cover (Canopy) indicate influence on species oc¬
currence. Species letter codes follow USDA (2014).

decades of passive protection (i.e. without management), and the closure of its
canopy likely occurred decades after that at Fogwell, but before that at Menger¬
son. Although Mengerson has been actively protected since the 1970s, its
canopy has only recently closed, as is apparent from historical aerial imagery.

50

THE MICHIGAN BOTANIST

Vol. 53

IT)

o

CM

U>

10

o

B

FRNI

CRM 02

JUNI ACSA3

+ +

MORU2

QUVEp6$££2 +

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TIAM

LITU
+ CEOC

cac a'»I a ? L8

„i 1DI 0ACA1B FAGR

jM 1 + GOPA2

CAOV2
QUAL

QUMA2

ACRU^

C0FL20SV1

- 1.0

- 0.5

0.0

0.5

AXIS 1

FIGURE 7. Nonmetric multidimensional scaling (NMDS) ordination of over¬
story (final stress = 0.22) for (A) survey plots for Fogwell (squares), IPFW (tri¬
angles), and Mengerson (circles) with 95% confidence ellipses for each forest;
and (B) pooled species. Species letter codes follow USDA (2014).

The difference observed in canopy cover was expected, given the differences
in the history of management at the three forests. Both IPFW and Mengerson
have histories of overstory tree removal, which is evidenced by the patchy
canopy visible in aerial images from the late 1930s (Figure 2). A simple visual
comparison of aerial images indicates that IPFW and Mengerson are likely now
at a canopy closure state similar to that of Fogwell from the 1930s to the 1950s.

2014 THE MICHIGAN BOTANIST 51

TABLE 2. Importance values of the top 10 overstory species in each of the forests studied. The rank
of each species within a forest is given in parentheses.

Species

Fogwell

IPFW

Mengerson

Acer saccharinum

—

—

22.7 (3)

Acer saccharum

87.6(1)

129.0(1)

40.1 (1)

Carya cordiformis

9.7 (8)

—

—

Carya ovata

39.6 (2)

—

20.4 (6)

Celtis occidentalis

—

9.6 (9)

—

Fagus grandifolia

37.1 (3)

—

—

Juglans nigra

6.4 (10)

21.7 (5)

12.1 (10)

Ostrya virginiana

—

—

15.9 (7)

Platanus occidentalis

—

6.6 (10)

—

Prunus serotina

—

13.2 (6)

—

Quercus alba

18.7 (6)

—

12.6 (9)

Quercus bicolor

—

—

12.7 (8)

Quercus rubra

25.0 (5)

11.0(7)

35.5 (2)

Quercus velutina

—

10.9 (8)

—

Tilia americana

29.2 (4)

28.3 (2)

21.2 (5)

Ulmus americana

7.6 (9)

24.1 (3)

22.0 (4)

Ulmus rubra

15.7 (7)

23.5 (4)

—

Although litter depth was significantly different statistically between the three
forests, of which Mengerson had the deepest litter, the numerical difference
(ranging from 1.1 to 2.8 cm) may not have been biologically significant. Litter
depth is a complex and dynamic relationship between addition and removal of
litter (Facelli and Pickett 1991). Because of the close proximity of the three
forests to each other, climate differences are likely to be very minor, resulting in
little difference in rainfall or length of the growing season; decomposition rates
are typically regulated by moisture and temperature (Facelli and Pickett 1991).
The cause of the differences in litter depth would likely be related to differences
in midstory densities, Mengerson had both the deepest litter, as well as the rich¬
est and densest midstory.

Rothrock (1997) originally calculated a C mean of 5.6 for Fogwell. However,
after Rothrock (2004) modified the coefficient of conservation values for Indi¬
ana specifically, C mean for Fogwell was recalculated as 4.1 (Rothrock and Ho-
moya 2005). This new value aligns exactly with our calculated C mean for Fog¬
well (4.1). However, our calculated FQI (23.4) was less than half the value
(59.3) calculated for previous surveys (Rothrock and Homoya 2005). This
demonstrates the difficulty of comparing floristic survey and ecological survey
studies in relation to various metrics of diversity or ecology. Rothrock (1997)
conducted a floristic survey of Fogwell with the explicit intent of producing a
full inventory of the preserve, which formed the basis of the calculation by
Rothrock and Homoya (2005). In contrast, we conducted an ecological survey
with the intent of providing a comparison between forests with different man¬
agement histories and different types of surrounding development. Thus, C mean
for each of the three forests is based on species with moderately high to high tol¬
erances for disturbance, using the C-values assigned by Rothrock (2004). All
three forests have experienced anthropogenic disturbances related to forest man-

52

THE MICHIGAN BOTANIST

Vol. 53

agement, although at different times for each forest. While using C-values to
compare communities should done cautiously, comparing counts of species
above or below a threshold may provide insight into forest disturbance or per¬
sistence. A C-value > 5 for a given species indicates it is likely to be found in
remnant areas similar to pre-settlement habitats (Rothrock 2004; Rothrock and
Homoya 2005). The relatively high percentage of species in Fogwell with a C-
value > 5 suggests that the understory plant community has recovered or simply
persisted from limited anthropogenic disturbances in the 1930s (i.e., more
species are adapted to less disturbance). Inversely, the lower percentage of
species at IPFW and Mengerson with a C-value > 5 suggests that the recent dis¬
turbances in forest management are still evident in the understory communities,
which is likely related to the later canopy closure in these forests. While C mean
was not significantly different between the three forests, the FQI for Fogwell
was substantially greater than it was for IPFW and Mengerson. Again, the dif¬
ference is related to the time that has elapsed since the most recent disturbance
and to the percentage of canopy closure within the three forests, Fogwell having
experienced little or no disturbance since the 1930s.

The density of the midstory is likely related to the time of canopy closure
(e.g., Mengerson had the most recent canopy closure and the highest density in
midstory individuals), and likely was the driver in accumulation of forest litter.
Although IPFW has had some anthropogenic disturbance related to forest man¬
agement, the canopy appears to have been much denser in the 1930s than the
canopy at Mengerson. The greater canopy density may have been an important
factor in the greater similarity in both the species observed at Fogwell and IPFW
and the midstory and overstory densities in those two forests. Because active for¬
est management has essentially stopped at IPFW and Mengerson, the overstory
density in both forests has reached similar values as that of Fogwell. While the
similarity in overstory species between forests is relatively high, several of the
top five and top ten most important species are unique to a single forest. Those
most important species are providing the physical structure of the forest. In ad¬
dition to being the most important species in the overstory, Acer saccharum was
the most frequent midstory species pooled across the three forests. Due to the
ability to maintain small stature individuals for decades in the shade (Marks and
Gardescu 1998), A. saccharum is a common midstory species in the region.
Once canopy gaps form, A. saccharum responds and can quickly be recruited
into larger size classes (Marks and Gardescu 1998), leading to inclusion in the
canopy.

The location of each forest has likely been an additional factor that has influ¬
enced the development of understory communities in these forests. In other
urban forests, there has been clear increase in non-native plant species (Dolan et
al. 2011). Fogwell, which has the richest understory and the greatest FQI value
among the forests, is a rural forest. Although there is a small subdivision to the
north of Fogwell, it is not surrounded by urban and suburban development, un¬
like IPFW and Mengerson, each of which were lower in species richness and
FQI values. We did not encounter any non-native understory species at Fogwell.
In contrast, non-native species accounted for 16% of the understory individuals
counted at IPFW and 26% at Mengerson. Again, this is likely related to the prox-

2014

THE MICHIGAN BOTANIST

53

imity of a forest to urban and suburban development and to the time that has
elapsed since canopy closure.

While these three forests had similarities in midstory and overstory species,
their importance to the region may exist more in their lack of similarity in un¬
derstory species. Most of the forest fragments in northeastern Indiana and the
surrounding region are small privately owned properties that have undergone a
broad range of protection and use over the past century. Allowing forests to un¬
dergo canopy closure and long-term minimized anthropogenic disturbance may
increase understory plant species richness and FQI, as seen in Fogwell. How¬
ever, as is apparent in the case of IPFW and Mengerson, proximity to urban de¬
velopment may be just as important a factor as time in promoting colonization of
understory communities by non-native species.

ACKNOWLEDGMENTS

The authors would like to tha nk ACRES Land Trust and the Indiana Department of Natural Re¬
sources Division of Nature Preserves for granting permission for access to Fogwell Forest and
Mengerson Nature Preserves. Specimen vouchers were collected under Indiana Department of Nat¬
ural Resources Division of Nature Preserves Research & Collecting Permit #NP 13-46. This research
was conducted as part of a field botany course at Indiana University-Purdue University Fort Wayne
with enrolled students participating equally as authors. Authors are listed alphabetically with the in¬
structor listed last, as the author of correspondence. The authors would like to thank Gary Helmke
for herbarium and identification assistance, as well as the two anonymous reviewers for their con¬
structive comments that improved this manuscript.

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town Square, Pennsylvania.

APPENDIX I. List of understory species collected at each of the forests studied. Frequency is the number of 2 m 2 plots (out of 17 at Fogwell and 20 at each of
IPFW and Mengerson) in which they were encountered. Count is the mean number of individuals (standard deviation in parentheses). Nomenclature follows ITIS
(2013). An asterisk (*) indicates non-native species (USDA NRCS 2013). Collection numbers are in parentheses after each species name..

Fogwell IPFW Mengerson

2014

THE MICHIGAN BOTANIST

55

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*Euonymus fortunei (Turcsz.) Hand.-Maz. (1257) 1 8.0

Fragaria virginiana Duchesne (1302) 4 7.5 (8.2)

(Continued on next page)

56

THE MICHIGAN BOTANIST

Vol. 53

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Penstemon digitalis Nutt, ex Sims (1317) 1 1.0

(Continued on next page)

APPENDIX I. (Continued).

Fogwell IPFW Mengerson

Species Frequency Count Frequency Count Frequency Count

Persicaria virginiana (L.) Gaertn. (1295) 1 2.0 4 6.0 (1.4)

2014

THE MICHIGAN BOTANIST

57

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THE MICHIGAN BOTANIST

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60

THE MICHIGAN BOTANIST

Vol. 53

REVIEWS OF STEVE W. CHADDE’S MISCELLANEOUS FIELD
GUIDES TO THE PLANTS IN THE GREAT LAKES REGION

Neil A. Harriman

Biology Department
University of Wisconsin-Oshkosh
Oshkosh, Wisconsin 54901
harriman@uwosh. edu

Steve W. Chadde 2013. Wisconsin Flora: An Illustrated Guide to the Vascu¬
lar Plants of Wisconsin, vi + 818 pp. ISBN 978-1490550022. Paperback.
$40.99.

Mr. Chadde is a Badger, having grown up in Kenosha, Wisconsin, he tells us
in the Introduction. He also co mm ents on the fact that, despite the long history
of botanical work in the state, there’s never been a comprehensive flora pub¬
lished for the state. The nearest thing to that is the series “Preliminary Reports
on the Flora of Wisconsin,” which have appeared over many decades in the
Transactions of the Wisconsin Academy of Sciences, Arts and Letters. The
Academy has ceased publication of its Transactions, as of 2001. These prelimi¬
nary reports were done under the aegis of Norman Fassett first, and then Hugh
litis. They are all listed on pp. 777-778; Mr. Chadde has done us all a service by
compiling the data, and giving the URL for finding them all online.

The keys work, the descriptions are excellent, and the illustrations are help¬
ful. I’ve used them for over a year. There are some typographic errors, inevitable
in a work this size, but all that I have reported to Mr. Chadde have been cor¬
rected. The core of the work, acknowledged on p. 777, is the online Wisconsin
State Herbarium, which contains names, photographs, and distribution maps, but
no descriptions and no keys. This is based on specimens in the herbarium of the
University of Wisconsin-Madison. The website somet im es deviates from the
classification adopted in Flora of North America; for example, Symphyotrichum
is not recognized, and traditional Aster continues to be used, the molecular evi¬
dence to the contrary notwithstanding. This is also true for the online Freckmann
Herbarium at UW-Stevens Point, which Chadde also cites as a source.

Neither of these online sources follows faithfully the Angiosperm Phylogeny
Group with respect to, for example, dismemberment of the Liliaceae. Chadde
adopts Liliaceae in the broad sense, and provides a listing of the genera of the
segregate families in a table on p. 653; he also puts the variant family name in
parentheses after each genus in the running treatment. This is very helpful to the
reader. The point of the book is to allow one to put a name to a thing; it is not the
place to ponder questions of phylogeny.

Sometimes, the odd introductions are mentioned in passing and are not in¬
cluded in the keys. In the Poaceae alone there are eleven such species—waifs
that have not become established as part of the naturalized flora. Similarly

2014

THE MICHIGAN BOTANIST

61

treated is Thymelaea passerina, an introduced annual; Thymelaeaceae are
treated as comprising only Dirca palustris, a native shrub. If the little annual
shows up again, one will have to go to Gleason & Cronquist to key it out.

But elsewhere, chance introductions or escapes from cultivation are treated
fully, though sometimes without a distribution map—for example, Marrubium
vulgare, in the Lamiaceae. By contrast, Satureja hortensis and Thymus pule-
gioides get no further mention whatever. In the same family, Collinsonia
canadensis is included in the keys (four times) but is neither described nor illus¬
trated; it is said to have been extirpated in Wisconsin, and perhaps so. It’s very
difficult to be consistent in handling these kinds of problems; the easiest (and
best, I think) solution is to include everything, with appropriate warnings to the
reader.

Yet another weedy mint is treated by Chadde as Leonurus marrubiastrum,
with the combination in Chaiturus given in synonymy. (It is known from only
two counties in Wisconsin, having been first discovered in 1970.) Most modem
manuals adopt Chaiturus marrubiastrum ; this is not a problem, because
Chadde’s index is very thorough and gives every entry of a scientific name,
whether adopted or not. The reader needs to be aware, Chadde is not much given
to taxonomic or nomenclatural wrangles. But he has created a landmark volume
in Wisconsin’s botanical history.

Steve W. Chadde. 2013. Minnesota Flora: An Illustrated Guide to the Vascu¬
lar Plants of Minnesota, vi + 781 pp. ISBN 978-1491224243. Paperback.
$39.99.

A flora of Minnesota, with keys, distribution maps, and illustrations, has not
existed until now. There is a rich history of botanical exploration in Minnesota,
briefly summarized in the Introduction, page 1. The dichotomous keys seem to
be entirely workable; they appear to be the same as those in Wisconsin Flora,
which I know to be most useful

The manual includes something over 1900 species. A figure of 2010 species
(1618 native, 392 introduced) for the state is given in Ownbey & Morley, 1991,
Vascular Plants of Minnesota: A Checklist and Atlas. The most recent survey
(available online) is Cholewa, 2011, Comprehensively Annotated Checklist of
the Flora of Minnesota, version 2011.2, where no totals are given.

Cholewa made an effort in her checklist to follow the available treatments in
Flora of North America and the families as recognized by the Angiosperm Phy-
logeny Group. Chadde has done likewise, for the most part. Hence, Aceraceae
disappear into Sapindaceae, Chenopodiaceae into Amaranthaceae, and the redis¬
tribution of genera out of classical Scrophulariaceae is adhered to. He retains
Liliaceae in the Cronquistian sense, but the segregate fa mi lies to which the gen¬
era are assigned in APG III are clearly indicated. The reader will have no diffi¬
culty in squaring these treatments with other recent works.

The problem of treating the non-native species remains. I note the author’s re¬
mark in the treatment of Acer that the widely cultivated Acer ginnala is occa¬
sionally found as an escape throughout the eastern half of Minnesota. The distri¬
bution map (all the maps are downloaded from bonap.net) shows it in a dozen or

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more counties; nonetheless, it is neither keyed, pictured, nor described. The
same fate befell it in Smith, 2008, Trees and Shrubs of Minnesota. In Ownbey &
Morley (see above) it is mentioned but is not granted a distribution map.

Out of idle curiosity, I looked to see how Chadde treated red mulberry, Morus
rubra. It is shown as very scattered, in only five counties, but extending from the
Canadian border south to the Iowa border. In Smith (see above) it is credited to
only Houston County in extreme southeast Minnesota. Ownbey & Morley have
it in two counties. Lastly, we have Nepal & Wichem, 2013, Taxonomic status of
Red Mulberry {Morus rubra, Moraceae) at its northwestern boundary, Proceed¬
ings of the South Dakota Academy of Science 92: 19- 29, where it is concluded
that the species does not occur in Minnesota - the few available specimens are
said to be hybrids with Morus alba, including the very early records that were
thought to vouch for the occurrence of red mulberry in Minnesota. The hybrid
arises naturally, and is also offered in the horticultural trade. None of this is in
Chadde - there’s simply not room.

What is in Chadde is all there in one place, accessible to the reader in plain
language, because he takes pains to avoid technical jargon as much as possible.
One can wish the author and his book every success.

Chadde, Steve W. 2014. Michigan Flora: Upper Peninsula, iv + 824 pp.
ISBN 978-1500566197. Paperback. $33.95.

The Upper Peninsula comprises just 15 counties, and 29% of the land area of
Michigan. Even so, it harbors nearly 1900 species of vascular plants.

The book covers pteridophytes and lycophytes, gymnosperms, and an-
giosperms (dicots and monocots), in that order. Aside from those dividing lines,
it is entirely alphabetical. The keys and descriptions are very well done; they are
adopted from Voss & Reznicek, Field Manual of Michigan Flora, as is properly
acknowledged on page 1. The treatments of pteridophytes and lycophytes rely
strongly on the treatments at michiganflora.net; however, the keys and descrip¬
tions are much easier to use when one has the printed word at hand - working
from a computer screen is relatively more awkward. Because the book is com¬
piled from such exemplary sources, it follows that readers will find it very use¬
ful. The addition of line drawings for most of the species is very helpful. There
are no distribution maps in Chadde’s book. A county map for the UP and for the
entire state is inserted on page 790.

Voss & Reznicek put monocots first, because that was standard when the
original Voss work was done; Chadde puts them last. Chadde does mostly rec¬
ognize the most recent Angiosperm Phylogeny Group families; hence, he leaves
Diervilla in the Caprifoliaceae, not in its own family, but with a note to the
reader. Voss & Reznicek recognize Dipsacaceae, as does Chadde, even though
APG III favors submerging that family into Caprifoliaceae. V&R also accept the
segregate families out of Liliaceae, such as Melanthiaceae, Trilliaceae, and As-
paragaceae, whereas Chadde sticks with traditional inclusive Liliaceae, but with
the other family names given in brackets.

Chadde takes some introduced species out of the keys and reduces their treat¬
ment to a note. One I noticed is Juncus inflexus L., introduced from Europe, not

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63

rare in wet roadside ditches in Houghton County. Omitting it from the key leaves
the reader flummoxed. A much better strategy, I think, is to put it in the book
along with everything else. (The running header for Juncaceae is misspelled, but
easily corrected for future printings.) To the contrary, in the Hydrocharitaceae,
Hydrocharis morsus-ranae L., introduced from Europe and known only from
Chippewa County in the UP, is included in the key, with an illustration, but does
not merit a detailed description. That seems to be a much better way of handling
introductions that may (or may not) turn out to be mere waifs.

In Judziewicz et al., Field Guide to Wisconsin Grasses, the authors made an
effort to track down the waifs reported for the state, to see if they are extant.
Many of them are not, but if they were here once, they could very well be here
again. Hence, they get the full treatment. This strikes me as a sensible way to go.
(The situation changes in California. There, the place is overrun with botanists
and with short-term plant introductions from both the temperate and the tropical
regions of the earth. A goodly number of these are not treated in The Jepson
Manual.)

Chadde’s intended audience is “students of the region’s flora.” He is serving
that cohort well, and I can only hope that his book will be stocked by merchants
in the UP for sale to the hordes of summer tourists; it’s a little tricky to take de¬
livery from amazon.com when you’re on vacation.

Chadde, Steve W. 2013. Midwest Ferns: A Field Guide to the Ferns and Fern
Relatives of the North Central United States, vi + 450 pp. ISBN 978-
1484161388. Paperback. $24.95.

North Central US in this book is taken to mean Minnesota, Wisconsin, Michi¬
gan, Iowa, Illinois, Indiana, and Ohio. The US Bureau of the Census includes in
the definition these states plus the Dakotas, Nebraska, and Kansas.

The work is lavishly illustrated, with both photographs and drawings, includ¬
ing an illustration of the fern life cycle, perhaps for those whose grounding in
Botany was more based on DNA than morphology. The illustrations of sori, leaf
forms, and spores is handy to have in one place.

The author says his book is a field guide, which generally means it’s meant to
aid in the identification of living plants, not dried herbarium specimens. To that
end, it opens not with conventional keys, but keys under nine different habitat
groups: aquatic, calcareous rock, non-calcareous rock, wet forest, etc. More-or-
less conventional keys begin on p. 31. The habitat keys I found difficult to use;
once I decided my fern was on a near-vertical sandstone cliff face (therefore,
non-calcareous), the key led me to ten parallel choices with no simple way to
distinguish among them. As it happened, the fern I was trying to key out is nor¬
mally a forest-floor species, and its occurrence on the cliff face was an oddity.

The descriptions of the species are very thorough. Where appropriate, each
species description is accompanied by county-level range maps for the upper
Midwest as well as maps showing total distribution in North America. The sci¬
entific names are fully explained or translated. The rare hybrids, some known
from only one or two counties, are included.

Traditional Lycopodium is here treated as six genera. The key to distinguish

64

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them is quite straightforward. Because the book is a field guide, not a taxonomic
treatise, there is no argument offered as to why all these genera should be recog¬
nized. The same taxonomic decisions are made in Michigan Flora Online, at
michiganflora.net. (The ferny parts are by Robert E. Preston, for whom an obit¬
uary appeared in Michigan Botanist 52(1-2): 53-54.) Chadde’s keys owe much
to those online, but he improves on them by intercalating explanations of tech¬
nical terms. I searched the book in vain for any acknowledgment of the online
flora. In addition, it might be well to point out an online work for Wisconsin:
Pteridophytes of Wisconsin: Ferns and Fern Allies at uwgb.edu. I was unable to
find similar sites for Illinois, Indiana, Minnesota, or Ohio.

There are references, pp. 429-431, helpfully separated into “Identification
Guidebooks,” “Fern Culture,” and “Selected Technical Reports.” This is a cir¬
cumstance where an annotated bibliography is wanted—the reader needs some
guidance. I looked and looked for Ferns and Fern Allies of Wisconsin, edition 2,
1953. (It is quite dated, but still useful.) Chadde has it, but with Fassett as first
author. No, the first author is Rolla Milton Try on, Jr. (1916-2001), whose name
somehow came to be listed last (of four).

The regularly updated websites for pteridophytes and lycophytes are very
helpful and useful, but they just aren’t the same as a book in hand. Chadde’s bar-
gain-priced book deserves a wide readership.

2014

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65

BOOK REVIEWS

Flora of North America Editorial Committee, editors. 2014 [2015]. Flora of
North America North of Mexico. Volume 9. Magnoliophyta: Picramniaceae to
Rosaceae. Oxford University Press, New York, N.Y. xxiv + 713 pp. ISBN
978-0-19-534029-7. Hardback. $95.00.

The Flora of North America (FNA), which is the first comprehensive flora pro¬
ject for all of North America north of Mexico and which is expected to fill 30
large volumes, is now somewhat more than half complete (this is the 18th pub¬
lished volume). Volume 9, for which 54 individual contributors are listed, con¬
tains treatments of four families, three of which—the Picramniaceae, the
Staphyleaceae, and the Crossosomataceae—treat a mere 11 species between
them and occupy only 15 pages. The remainder of the volume (645 pages) is de¬
voted to an in-depth treatment of the Rosaceae, a large family that, on a world¬
wide basis, is most prominent in the north temperate region. This volume has a
particularly special significance far beyond the borders of North America for
those seeking to understand taxonomic relationships within the Rosaceae. This is
because, although FNA contains families that are larger on a world-wide basis
and that include a larger number of species in the Flora area, such as Asteraceae
(2413 species), Poaceae (1373 species), and Cyperaceae (843 species), the FNA
treatment of Rosaceae contains a larger percentage of its species within the Flora
area (680 species of 3000, or 23%), as compared to 10% for the Asteraceae, 17%
for Cyperaceae, 12% for Poaceae, nearly 20% for the Brassicaceae. More sig¬
nificantly, however, the Flora contains representatives of all three subfamilies
and all 16 tribes of the Rosaceae, as well as 68 of the 88 genera (77%), a breadth
of taxonomic diversity that cannot be said of any other large family in North
America (some medium-sized families, such as Onagraceae or Boraginaceae,
may approach this level of coverage).

For students of the North American flora, including that of the Great Lakes
region, however, the numerous wholly new generic treatments are most wel¬
come. Among these is a completely new treatment of the large genus Potentilla
and its segregates by Barbara Ertter assisted by Reidar Elven, the late James Re¬
veal, and David Murray. This has always been considered a taxonomically com¬
plex group. Ninety-eight species of Potentilla itself are recognized within the
borders of the Flora, including Potentilla anserina L., which has often in recent
years been segregated as the genus Argentina (and which here includes within P.
anserina arctic and west coast populations often recognized as P. egedei ). Sev¬
eral genera traditionally included within Potentilla are recognized, however.
Among them are the following that include species well-known from the Great
Lakes region: Drymocallis (including the prairie cinquefoil, D. arguta = Poten¬
tilla arguta), Dasiphora, which includes the widespread shrubby cinquefoil, D.
fruticosa = Potentilla fruticosa, Sibbaldiopsis (including S. tridentata = Poten¬
tilla tridentata ), and Comarum (including C. palustre = Potentilla palustris).

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The Rosaceae is well known to harbor a number of genera of great taxonomic
complexity resulting from such processes as hybridization, apomixis, and poly¬
ploidy, prominent among them Rubus, Rosa, Alchemilla, and Crataegus. Al-
chemilla is primarily an Old World genus represented in the Flora by just 12
species, mostly arctic. The other three receive conservative treatments in the
Flora, properly so, in my opinion, for a work intended to provide a floristic
overview, albeit detailed and comprehensive, for the broad botanical commu¬
nity, rather than monographic treatments for the specialist. The large and com¬
plex genus Rubus is reduced to 37 species continent-wide, only two of which
were described later than 1899 (1934 and 1943), but a large synonymy is pro¬
vided so that the innumerable microspecies can be placed by the interested
reader. The species better known as Dalibarda repens is here included in Rubus
on molecular grounds, which is necessary, according to the authors, in order to
preserve the monophyletic status of Rubus. The treatment of Crataegus by the
long-time student of that genus, James Phipps, is the first for the entire Flora area
since the work of Torrey and Gray in 1838-1843. It recognizes 169 species; and,
although conservative, an important objective of the treatment is “to draw atten¬
tion to the often great, but little discussed, inffaspecific variation.” The author’s
long and extensive field acquaintance with the genus has made this possible.

I have mentioned only a few of the many significant accomplishments of this
magisterial volume. Readers will enjoy up-to-date and comprehensive treat¬
ments of other favorite genera, including Geum, Prunus, Cotoneaster, Spiraea,
Horkelia, Filipendula, and Sorbus, among many others. The genus Photinia is
treated as including only 4 introduced species, restoring the native chokeberries,
which some recent authors have included here, to their more traditional place¬
ment in Aronia.

As is true of all the volumes of FNA, a particularly welcome feature of this
volume is the often extensive commentary following the descriptions of genera,
and less frequently of subgeneric taxa and species. There is no specific format
for these commentaries, but authors are given free rein to discuss topics such as
circumscription, taxonomic history, and broad issues of distribution, as well as
explanations of and justifications for the particular taxonomic approach taken,
and, in many cases, discussions of special terminology or unusual features in the
keys. This is just one of many factors that will make Flora of North America a
significant contribution lasting long into the future and well beyond the borders
of North America.

-Michael Huft

James W. Byng. 2014. The Flowering Plants Handbook. A practical guide to
families and genera of the world. Plant Gateway Ltd. (plantgateway.com),
619 pp. ISBN 978-0-9929993-0-8. Paperback. $56.90.

If you attempt to order the print copy of this book from the usual online out¬
lets, you may be disappointed, temporarily. But keep trying. It is not available
from the publisher’s website, strangely. There is an eBook edition, which is re¬
viewed by Neil Snow in Systematic Botany 40(1): 366. 2015. Snow called atten-

2014

THE MICHIGAN BOTANIST

67

tion to some “printing” problems; I checked to see if these had found their way
into the printed book, and found that they had not.

The book is intended to get you to family in all cases, to genus in most cases.
On Cover 4, it states plainly “This plant book aims to help identify flowering
plants to genus and family level anywhere in the world.”

Everything follows Angiosperm Phylogeny Group III. One has first to get
one’s plant into one of the major categories: Asterids, Rosids, basal an-
giosperms, etc. Then you choose which Order you think your plant might belong
to. For each Order, there is a conventional key to families. Families are given a
floral formula, explained on p. 1; for larger families, this is necessarily so broad
as to be of little use. For most readers, the whole point is to get to genus. And
here is where problems arise, because there are no longer keys, just synopses of
the included genera, accompanied by beautiful color photographs. But the de¬
scriptions are not parallel; they jump all over. I tried an unknown (from cultiva¬
tion, native range unknown) in the Rosaceae, with three subfamilies. It was all I
could do to decide on a subfamily, among Dryadoideae, Spiraeoideae, and
Rosoideae. Within the Spiraeoideae alone, there are eight tribes and 58 genera -
all this without a key. It is a daunting task, but one is assured all the genera are
there, so one must persevere.

A sterner test might be Fabaceae. Forget it. The treatment is but 1.3 pages
long, ending with an “advertisement” to consult Lewis et al., 2005, Legumes of
the World. This is 577 pages of keen scholarship, elegant language, and magnif¬
icent illustrations, nearly all in full color—$140. But there’s nary a key to be
found in this book, either.

The treatment of Asteraceae is also scanty, 2.3 pages long. There are said to
be 1600 genera and maybe 24,000 species. If the family were treated fully, one
would need a wheelbarrow to carry the book(s) around. Instead, one is referred
to various websites. There is no “advertisement” urging one to consult, for ex¬
ample, Funk et al. 2009, Systematics, Evolution, and Biogeography of Composi-
tae, again beautifully written and illustrated, but without keys of any sort—$110.

The situation for Poaceae (740 genera, 11,500 species) is quite parallel to the
two examples above, 2.5 pages long. You will still need Clayton & Renvoize,
Genera Graminum, with very thorough keys—$90. And these remarks are not
intended as a criticism of the author. He did the best he could, but the amazing
diversity is simply too great to handle in one book. The work may be most use¬
ful for the very small and very obscure families, especially of the tropics, such
as Petenaeaceae, Tapisciaceae, or Dipentodontaceae.

There are 67 pages in 7-point type of “Literature [Cited].” It’s all the latest
findings, mostly molecular, and therefore with no mention of the classical au¬
thors (Linnaeus, the Hookers, Asa Gray, et alii). The index is very thorough, to
scientific names only. This is not a work for the neophyte.

-Neil A. Harriman

Biology Department
University of Wisconsin-Oshkosh
Oshkosh, Wisconsin 54901
harriman@uwosh.edu

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CATALOGUE OF FLORISTIC ARTICLES IN
THE MICHIGAN BOTANIST

Michael Huft

The Michigan Botanist has a long history of publishing floras and floristic
studies of natural areas in the Great Lakes region, including such major units as
counties, national lakeshores, and state parks, but also islands in the Great Lakes
and even such smaller units as individual woodlots, bogs, or defined wetlands, to
name a few. These studies have covered most major groups of plants, including
lichens, algae, bryophytes, and vascular plants (sometimes limited to subdivi¬
sions of the latter, such as pteridophytes or aquatic plants).

Because these articles provide a rich resource for present-day and future ex¬
plorations and studies, it is useful to make available an accessible catalogue so
that readers can readily find what is available and where an item of interest can
be found. It is also hoped that this list will suggest further areas that need floris¬
tic work or those areas where updates would be useful.

Items are arranged by major plant group and, under Vascular Plants, by state
or province. Within each category, the listing is chronological, and the volume
and page citation is given at the end of each item. A very few items that deal
with more than one plant group or more than one state are listed under each ap¬
propriate category.

LICHENS

Henry, Mary Lou, and Raymond E. Hampton. (1974). Macrolichens of Beaver Island, Charlevoix
County, Michigan. 13: 65-67.

Wetmore, Clifford M. (1988). Lichens of Sleeping Bear National Lakeshore. 27: 111-118.

Wetmore, Clifford M. (1990). Lichens of Apostle Islands National Lakeshore, Wisconsin. 29: 65-73.

ALGAE

Andreson, Norman A. (1970). Algae in Park Lake, Clinton County, Michigan. 9: 95-107.

Andresen, N. E., and E. F. Stoermer. (1978). The diatoms of Douglas Lake—A preliminary list. 17:
49-54.

Hoagland Kyle D., and Forrest M. Begres. (1978). The diatom flora of Tiplady Bog, Michigan. 17:
55-64.

Lipsey, Louis L., Jr., and Mason G. Fenwick. (1978). Investigations on the freshwater Algae of Wis¬
consin. I. Plankton diatoms from High Lake (Vilas County). 17: 113-117.

Glime, Janice M., Janet F. Gereau, and Roy E. Gereau. (1980). Bryophytes of Mount Lookout, Ke¬
weenaw County, Michigan. 19: 247-250.

Anderson, Kevin, Mark Fate, Ching-Han (Eric) Hsieh, Lorenz Kim, Katy Lazarus, J. Patrick Koci-
olek, and Rex L. Lowe. (2013). Algal diversity in, and the description of three new diatom
(Bacillariophyta) species from, Lake of the Clouds, Porcupine Mountains Wilderness State Park,
Michigan USA. 52: 3-24.

BRYOPHYTES

Crum, Howard. (1964). Mosses of the Douglas Lake region of Michigan. 3: 3-12, 48-63.

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69

Miller, Norton G. (1972). A bryophyte florula of an area near Grand Ledge, Eaton County, Michi¬
gan. llr 3-12.

Koch, Rudy G., and Duane J. Lahti. (1973). The bryophytes of Amnicon Falls State Park, Douglas
County, Wisconsin. 12: 212-216.

Hollenson, Raymond H. (1984). A checklist of the liverworts of Michigan. 23: 135-139.

Glime, Janice M., and Allison D. Slavik. (1985). A checklist of bryophytes and their critical locali¬
ties in the Keweenaw Peninsula, Michigan. 24: 154-163.

Stevens, Kevin, and Elwood B. Ehrle. (1990). The Hampton Creek wetland complex in southwest¬
ern Michigan. VI. Transect survey of bryophyte populations. 29: 55-64.

Wheeler, Gerald A., Robert P. Dana, and Carmen Converse. (1991). Contribution to the vascular
(and moss) flora of the Great Plains: A floristic survey of six counties in western Mi nn esota. 30:
75-129.

Glime, Janice M. (2002). Bryophytes of the Pictured Rocks National Lakeshore, Alger County,
Michigan. 41: 31—45.

Ehrle, Elwood B. (2005). Mosses and liverworts of the Kalamazoo Nature Center, a preliminary sur¬
vey. 44: 119-126.

VASCULAR PLANTS

Illinois

Moran, Robbin C. (1978). Vascular flora of the ravines along Lake Michigan in Lake County, Illi¬
nois. 17: 123-140.

Indiana

Angstmann, Julia L., Paul E. Rothrock, and Thomas W. Post. (2005). The vascular flora and com¬
munity structure of little Calumet Headwaters Nature Preserve, LaPorte County, Indiana. 45:
153-192.

Namestnik, Scott, and Keith Board. (2010). The vascular flora of Sebert Property, LaPorte County,
Indiana. 49: 1—40.

Michigan

Hagenah, Dale J. (1963). Pteridophytes of the Huron Mountains, Marquette County, Michigan. 2:
78-93. (See additions below in Wahla, 1984)

Crow, Garrett E. (1969). Species of vascular plants of Pennfield Bog, Calhoun County, Michigan. 8:
131-136.

Beaman, John H. (1970). A botanical inventory of Sanford Natural Area. I. The Environment. 9:
116-139. II. Checklist of vascular plants. 9:147-164.

Veldman, Lorraine Calhoun, and Daniel E. Wujek. (1971). Pteridophytes of Beaver Island,
Charlevoix County, Michigan. 10: 194-196.

Wells, James R., and Paul W. Thompson. (1974). Vegetation and flora of Keweenaw County, Michi¬
gan. 13: 107-151.

Read, Robert H. (1975). Vascular plants of Pictured Rocks National Lakeshore, Alger County,
Michigan. 14: 3^43.

Frye, Darlene M. (1976-1977). A botanical inventory of Sandhill Woodlot, Ingham County, Michi¬
gan. I. The vegetation. 15: 131-140. II. Checklist of vascular plants. 15: 195-204. III. Phenology
study. 16: 34-38.

Haynes, Robert R., and C. Barre Hellquist. (1978). The distribution of the aquatic vascular flora of
Douglas Lake, Cheboygan County, Michigan. 17: 183-191.

Beach, James H., and W. D. Stevens. (1979-1980). A study of Baker Woodlot. I. Physical and his¬
torical description. 18: 123-137. II. Description of vegetation. 19: 3-13. III. Checklist of Vascu¬
lar Plants. 19: 51-69 (Stevens and Beach).

Sytsma, Kenneth J., and Richard W. Pippen. (1981-1982). The Hampton Creek wetland complex in
southwestern Michigan. I. History and Physical features. 20: 137-142. II. Community classifica¬
tion. 20: 147-156. III. Structure and succession of tamarack forests. 21: 67-74. IV. Fen succes¬
sion. 21: 105-115. V. Species of vascular plants. 21: 195-204. (See Part VI under Bryophytes,
above).

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Mustard, Timothy S. (1983). The vegetation of the Manistee National Forest, Oceana and Mason
Counties, Michigan. I. Physical, historical, and ecological aspects. 22: 111-122. II. Checklist of
vascular plants. 22: 1151-161.

Weitzman, Anna L. (1984). Sommerby Swamp, an unusual plant community in Mackinac County,
Michigan. 23: 11-18.

Wahla, James. (1984). Additions to the pteridophyte flora of Marquette County, Michigan. 23:
178-179. (Updating Hagenah, 1963, above)

Kron, Kathleen A., and Beverly Scoviak Walters. (1986). A study of Red Cedar Natural Area. I. His¬
torical background and description of the vegetation. 25: 27-34. II. Checklist of vascular plants.
25: 35^44.

Hazlett, Brian T. (1986). The vegetation and flora of the Nordhaus Dunes, Manistee national Forest,
Mason County, Michigan. I. History and present vegetation. 25: 74-92. II. Catalogue of vascular
plants. 25: 125-139.

Slavick, Allison D., and Robert A. Janke. (1987). The vascular flora of Isle Royale National Park.
26: 91-134.

Kron, Kathleen A. (1989). The vegetation of Indian Bowl Wet Prairie and its adjacent plant com¬
munities. I. Description of the vegetation. 28: 179-200. II. Checklist of vascular plants. 28:
201-215.

Hazlett, Brian T. (1991). The flora of Sleeping Bear Dunes National Lakeshore, Benzie and Leelanau
Counties, Michigan. 30: 139-202.

Meagher, Walter L., and Stephen J. Tonsor. (1992). Checklist of the vascular flora of the Augusta
Floodplain Preserve. 31: 83-98.

Forzley, Kathleen C., Thaddeus A. Grudzien, and James R. Wells. (1993). Comparative floristics of
seven islands in northwestern Lake Michigan. 32: 3-21.

Hazlett, Brian T. (1993). The vascular flora of North and South Fox Islands, northern Lake Michi¬
gan. 32: 239-264.

Judziewicz, Emmet J. (1997). Vegetation and Flora of Passage Island, Isle Royale National Park,
Michigan. 36: 35-62.

Judziewicz, Emmet J. (1997). Michigan’s farthest north: A botanical visit to the Gull Islands, Isle
Royale National Park. 36: 78-87.

Laureto, Pamela J. (1998). Vascular flora and plant community types of Seidman Park, Kent County,
Michigan. 37: 67-89.

Voss, Edward G. (2001). Flora of St. Helena Island (Straits of Mackinac), Michigan. 40: 27-47.

Judziewicz, Emmet J. (2001). Flora and vegetation of the Grand Traverse Islands (Lake Michigan),
Wisconsin and Michigan. 40: 81-208.

Ross, Stephen, and Robert J. Krueger. (2002). A botanical survey of the White Pine Trail State Park
(Rails-to-Trails) in Mecosta County, Michigan. 41: 51-84.

Slaughter, Bradford S., and J. Dan Skean, Jr. (2003). Annotated checklist of vascular plants in the
vicinity of Cedar Creek and Brewster Lake, Pierce Cedar Creek Institute, Barry County, Michi¬
gan. 42: 127-148.

McKenna, Duane D. (2004). Flora and vegetation of Kalamazoo County, Michigan. 43: 137-359.

Whately, Cathryn Elizabeth, Daniel E. Wujek, and Edwin E. Leuck II. (2005). The vascular flora of
Hog Island, Charlevoix County, Michigan. 44: 29-48.

Williams, Gretchen M., Gilbert D. Starks, and Daniel E. Wujek. (2005). Annotated checklist of vas¬
cular plants of Neithercut Woodland, Clare County, Michigan. 44: 57-71.

Smith, Pamela F., and Dennis W. Woodland. (2006). Vascular plant study of Warren Dunes State
Park, Berrien County, Michigan. 45: 1-58.

Springer, Joshua C., and Bruce D. Parfitt. (2010). A checklist of vascular plants of the Lapeer State
Game Area, Lapeer County, Michigan, USA. 49: 41-72.

Baker, Kathleen M., and C. Elizabeth Whately. (2010). Vascular plant survey of privately owned
wetlands, Pavilion Township, Kalamazoo County, Michigan. 49: 85-98.

Bassett, Tyler. (2011). Annotated checklist of the flora of the Kalamazoo Nature Center with natural
community descriptions. 50: 41-104.

2014

THE MICHIGAN BOTANIST

71

Minnesota

Joyal, Robert. (1971). The Marshall Bog near Itasca Park, Minnesota. 10: 78-88.

Wheeler, Gerald A., and Paul H. Glaser. (1982). Vascular plants of the Red Lake Peatland, northern
Minnesota. 21: 89-93.

Wheeler, Gerald A., Robert P. Dana, and Carmen Converse. (1991). Contribution to the vascular
(and moss) flora of the Great Plains: A floristic survey of six counties in western Mi nn esota. 30:
75-129.

Anderson, Derek, Jessica Brandt, Lynn Wright, and Donald Davidson. (2005). Ecology and floristics
of Knife Island, a gull and cormorant rookery on Lake Superior, near Two Harbors, Lake
County, Minnesota. 44: 95-104.

Ohio

Stuckey, Ronald L. (1975). A floristic analysis of the vascular plants of a marsh at Perry’s Victory
Monument, Lake Erie. 14: 144-166.

Tyrrell, Lucy E. (1987). A floristic survey of buttonbush swamps in Gahanna Woods State Nature
Preserve, Franklin County, Ohio. 26: 29-38.

Whyte, Robert S., David A. Francko, and David M. Klarer. (2003). The aquatic vegetation of the old
Woman Creek National Estuarine Research Reserve (Huron, Ohio): A Lake Erie coastal wetland.
42: 63-84.

Ontario

Reznicek, A. A., and P. M. Catling. (1989). Flora of Long Point, Regional Municipality of
Haldimand-Norfolk, Ontario. 28: 99-175.

Morton, J. K., and Joan M. Ve nn . (1996). The flora of Caribou Island, Lake Superior. 35: 3-25.

Weatherbee, Ellen Elliott (In Press). The flora of Cockbum Island, Ontario, Canada.

Wisconsin

Musselman, Lytton J., Theodore S. Cochran, William E. Rice, and Marion M. Rice. (1971). The
flora of Rock County, Wisconsin. 10: 147-193.

Dabydeen, Simon, and Rudy G. Koch. (1977). Vegetation and floristics of a sand spoil deposit—
Barker’s Island in Superior Harbor, Douglas County, Wisconsin. 16: 39-46.

Peck, James H., and W. Carl Taylor. (1980). Check list and distributions of Wisconsin ferns and fem
allies. 19: 251-268.

Koch, Rudy G., Patti Younger, and Leo Bruederle. (1983). Vegetation and floristics of Wisconsin
Point, Douglas County, Wisconsin. 22: 35^46.

Cochrane, Theodore S., Marion M. Rice, and William E. Rice. (1984). The flora of Rock County,
Wisconsin: Supplement I. 23: 121-133.

Forzley, Kathleen C., Thaddeus A. Grudzien, and James R. Wells. (1993). Comparative floristics of
seven islands in northwestern Lake Michigan. 32: 3-21.

Judziewicz, Emmet J., and Rudy G. Koch. (1993). Flora and vegetation of the Apostle Islands Na¬
tional Lakeshore and Madeline Island, Ashland and Bayfield Counties, Wisconsin. 32: 43-189.

Eddy, Thomas L. (2001). A vascular flora of the Norwegian Bay Wetlands on Green Lake, Green
Lake County, Wisconsin. 40: 51-69.

Judziewicz, Emmet J. (2001). Flora and vegetation of the Grand Traverse Islands (Lake Michigan),
Wisconsin and Michigan. 40: 81-208.

Fields, Douglas M. (2003). The vascular plants of Taylor County, Wisconsin. 42: 171-282.

Judziewicz, Emmet J. (2004). A thirty-year study of the vascular plants of “Butternut Pines,” a 40-
acre tract in Oconto County, Wisconsin. 43: 81-115.

Hart, Charles R. (2004). A checklist of the vascular flora of Woodland Dunes Nature Center, Mani¬
towoc—Two Rivers, Wisconsin. 43: 361-375.

Herrick, Bradley M. (2005). The vegetation of Point Au Sable Nature Preserve, Brown County, Wis¬
consin, USA. 44: 49-55.

Hlina, Paul, Derek S. Anderson, and Donald W. Davidson. (2008). Flora of Amnicon Falls State
Park, Douglas County, Wisconsin. 47: 121-146.

Hlina, Paul S., and Derek S. Anderson. (2011). Wetland assessment and inventory of the Pokegama
Carnegie Wetland State Natural Area, Douglas County, Wisconsin. 50: 2-25.

Anderson, Derek, and Julie Fox. (2014). Flora of Interstate State Park, Polk County, Wisconsin.

72

THE MICHIGAN BOTANIST

Vol. 53

CALL FOR PAPERS

We currently have articles that will fill the three remaining double issues
planned during 2015. However, we will need more articles to start filling the
2016 issues, and the sooner they can be submitted the better. Please consider
what you have available that can be turned into a paper for The Michigan
Botanist - taxonomy, forest ecology, floristic surveys, conservation, use of
plants by humans, historical studies, and many other topics. If you have any
questions about the suitability of a proposed article, please feel free to contact
the editor. Student-authored papers are always welcome. Short research notes
and book reviews are also welcome.

We’d also like to consider some occasional special topic issues, in which sev¬
eral papers by different authors would explore different aspects of some area of
concern. Some ideas might be Invasive Plants, or The Decline in Field Botany
and Collections-Based Research, or The Role and Management of Nature Pre¬
serves or other conservation topics. Others could be a series of articles (which
need not appear in a single issue, but could appear singly over time) on topics
such as the history of floristics and flora writing in each of the several states and
provinces that make up the Great Lakes region, or articles on the development,
management, and importance of nature preserves, parks, land trusts, and similar
areas in each of these states. These are just a few suggestions that come to mind.
I would welcome any ideas, as well as any volunteers to take on the task (or as¬
sist in the task) of assembling authors and articles for these types of ventures.

INSTRUCTIONS TO AUTHORS

Refer to http://quod.lib.umich.edU/m/mbot/submit for more detailed instructions, especially

for formatting, style conventions, literature cited, and voucher specimen requirements. Please

contact the editor with any questions.

1. Create text in 12-point Times New Roman font and double space paragraphs throughout.
Research articles should be organized as follows: Title, Author(s) and address(es), Abstract with
up to 5 keywords, Introduction, Materials and Methods, Results, Discussion,
Acknowledgements, Literature Cited, Tables, Figure Legends, and Figures. Sections may be
omitted if not relevant. All pages should be numbered.

2. For noteworthy collections, manuscripts should be formatted as follows. The title, “Noteworthy
Collections,” should begin each submitted manuscript, followed on the next line by the State or
Province for the species reported. The next line should list the taxon of interest using the fol¬
lowing format: Species Author(s) (Family). Common name. The rest of the manuscript should
include the following named sections: (i) Significance of the Report, (ii) Previous Knowledge,
(iii) Discussion, (iv) Diagnostic Characters (if desired), (v) Specimen Citations, (vi)
Acknowledgements (if desired), and (vii) Literature Cited. Each of these sections is largely self-
explanatory; however, the “Significance of the Report” section should be limited to a brief sen¬
tence or phrase indicating the significance of the collection(s), and this may be expanded upon
in the “Discussion” section; the “Specimen Citations” section should include the relevant label
data from the voucher specimen(s) including location data, collector(s), collection number, etc.,
as well as the Index Herbariorum acronym(s) of the herbarium or herbaria where the specimen(s)
are deposited. The manuscript should end with the name and address of the author(s).

3. Non-research articles, such as book reviews, letters to the editor, notices, biographies and other
general interest articles can be formatted as general text without the specific sections listed
above. However, literature cited and any tables or figures should be formatted as described
below.

4. Create tables either as an MS Word table or using a tab-delimited format. Each table is to be sub¬
mitted as a separate file. Table captions should be placed at the top of the table. Any footnotes
should appear at the bottom of the table. Please do not insert tables within the body of the text.

5. Send each figure as a separate file in a high-resolution format—eps, jpg, or tif. Figures like bar
graphs that gain their meaning with color won’t work—use coarse-grained cross-hatching, etc.
Create figure legends as a separate text file, and the typesetter will insert them as appropriate.
Please do not insert the figure in the body of the text file.

6 . Citations: Please verify that all references cited in the text are present in the literature cited sec¬
tion and vice versa. Citations within the text should list the author’s last name and publication
year (e. g. Smith 1990). For works with more than 2 authors, use “et al.”, and separate multiple
citations with a semicolon.

7. Literature Cited: List citations alphabetically by author’s last name. The first author’s name is to
be listed with surname first, followed by initials (e.g. Smith, E. B.), and subsequent authors are
to be listed with initials first.Separate author’s initials with a single space. The year of publica¬
tion should appear in parentheses immediately before the title of the citation. The entire journal
name or book title should be spelled out. Please put a space after the colon when citing volume
number and page numbers.

8 . Italicize all scientific names. Voucher specimens must be cited in floristic works and in any other
study whose results depend on the identity of the plant(s). Papers citing plant records without
documenting vouchers are generally not acceptable.

9. Manuscripts must be submitted electronically to the email address of the editor. All manuscripts
will be reviewed by at least two referees.

CONTENTS

In This Issue

Michael Huft

1

Flora of Interstate State Park, Polk County, Wisconsin

Derek S. Anderson and Julie Fox 2

Comparative Analysis of Urban and Rural Forest Fragment
Structure and Diversity in Northeastern Indiana

Kristine D. Arvola, Parker F. Booth, Charles C. Ellinwood,

Angela J. Fry, Jason L. Furge, Kaitlyn A. Haehnle,

Lauren E. Hall, Melanie A. Jablonski, Daniel K. Jones,

Justin T. Martin, Kevin M. McLane, Kevan C. Mensch,

Victoria A. Mumaw, Rebeca N. Quirindongo, Michael J.

Ravesi, Jesse J. Rinard, Patrick R. Selig, Andrew P. Sellan,
Maja B. Sljivar, Emily A. Stulik, Tasha R. Sunday, Alison N.
Turley, MarkT. Voors, Adam R. Warrix, Tyler C. Wood, and
Jordan M. Marshall 39

Reviews of Steve W. Chadde’s Miscellaneous Field Guides to the Plants
in the Great Lakes Region

Neil A. Harriman 60

Book Reviews 65

Catalogue of Floristic Articles in The Michigan Botanist

Michael Huft 68

Call for Papers 72

On the cover: The Dalles of the St. Croix River. The state of Wisconsin is on the left side of the
river, and the State of Minnesota is on the right. Here, the tallest cliffs rise approximately
30 meters (100 ft) above the river. Photograph by Derek Anderson, November 8, 2013.