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BioOne sees sustainable scholarly publishing as an inherently collaborative enterprise connecting authors, nonprofit publishers, academic institutions, research libraries, and research funders in the common goal of maximizing access to critical research. Changes In the Vascular Flora of the Middlesex Fells Reservation, Middlesex County, Massachusetts, from 1895 To 2011 Author(s): Bryan T. Hamlin, Walter T. Kittredge, Donald P. Lubin, and Elizabeth Barton Wright Source: Rhodora, 114(959):229-308. 2012. Published By: The New England Botanical Club, Inc. DOI: http://dx.doi.org/10.3119/11-02 URL: http://www.bioone.org/doi/full/10.3119/11-02 BioOne (www.bioone.org ) is a nonprofit, online aggregation of core research in the biological, ecological, and environmental sciences. BioOne provides a sustainable online platform for over 170 journals and books published by nonprofit societies, associations, museums, institutions, and presses. Your use of this PDF, the BioOne Web site, and all posted and associated content indicates your acceptance of BioOne’s Terms of Use, available at www.bioone.org/page/terms_of_use . Usage of BioOne content is strictly limited to personal, educational, and non-commercial use. Commercial inquiries or rights and permissions requests should be directed to the individual publisher as copyright holder.

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Page 1: Changes In the Vascular Flora of the Middlesex Fells Reservation, Middlesex County, Massachusetts, from 1895 To 2011

BioOne sees sustainable scholarly publishing as an inherently collaborative enterprise connecting authors,nonprofit publishers, academic institutions, research libraries, and research funders in the common goal ofmaximizing access to critical research.

Changes In the Vascular Flora of the Middlesex FellsReservation, Middlesex County, Massachusetts, from1895 To 2011Author(s): Bryan T. Hamlin, Walter T. Kittredge, Donald P. Lubin,and Elizabeth Barton WrightSource: Rhodora, 114(959):229-308. 2012.Published By: The New England Botanical Club, Inc.DOI: http://dx.doi.org/10.3119/11-02URL: http://www.bioone.org/doi/full/10.3119/11-02

BioOne (www.bioone.org) is a nonprofit, online aggregation of coreresearch in the biological, ecological, and environmental sciences. BioOneprovides a sustainable online platform for over 170 journals and bookspublished by nonprofit societies, associations, museums, institutions, andpresses.

Your use of this PDF, the BioOne Web site, and all posted and associatedcontent indicates your acceptance of BioOne’s Terms of Use, available atwww.bioone.org/page/terms_of_use.

Usage of BioOne content is strictly limited to personal, educational, andnon-commercial use. Commercial inquiries or rights and permissionsrequests should be directed to the individual publisher as copyright holder.

Page 2: Changes In the Vascular Flora of the Middlesex Fells Reservation, Middlesex County, Massachusetts, from 1895 To 2011

CHANGES IN THE VASCULAR FLORA OF THE

MIDDLESEX FELLS RESERVATION,

MIDDLESEX COUNTY, MASSACHUSETTS,FROM 1895 TO 2011

BRYAN T. HAMLIN

56 Wyman Street, Medford, MA 02155e-mail: [email protected]

WALTER T. KITTREDGE

Harvard University Herbaria, 22 Divinity Avenue, Cambridge, MA 02138

DONALD P. LUBIN

8 Ashford Street, Allston, MA 02134

ELIZABETH BARTON WRIGHT

420 Hemenway St. #253, Marlborough, MA 01752

ABSTRACT. A study of the vascular flora of the Middlesex Fells Reservation, asuburban forest reserve in greater Boston, was made over nine years from 2003 to2011. We compiled a checklist, determining the frequency of each taxon anddocumenting all natural community types occurring in the reservation. Wecompared our findings to those made in the 1890s, and attempted to explainchanges in species composition and frequency. Despite many impacts on thereservation in the last century, a high level of species richness was observed, with868 vascular taxa extant, including 563 native taxa. There was little net change inthe number of native taxa found, but a near tripling of non-native taxa from 110to 305. Habitat destruction, fragmentation, forest succession, fire suppression,trends favoring mesophytic vegetation, climate change, and invasive species areall believed to be factors in the changes observed in the flora.

Key Words: afforestation, biodiversity, climate change, disturbance ecology,fire, forest succession, fragmentation, frequency, invasive plants,land-use history, mesophication, Middlesex Fells, plant commu-nities, vascular flora

The Middlesex Fells (hereafter referred to as the Fells) is a

suburban forest of 1400 ha within Middlesex County, Massachu-

setts. About a quarter of the area is open water, mostly in the form

of reservoirs. Incorporated as a Metropolitan Parks reservation in

1894, the majority of the Fells is now under the jurisdiction of the

Massachusetts Department of Conservation and Recreation

RHODORA, Vol. 114, No. 959, pp. 229–308, 2012

E Copyright 2012 by the New England Botanical ClubDOI: 10.3119/11-02

229

Page 3: Changes In the Vascular Flora of the Middlesex Fells Reservation, Middlesex County, Massachusetts, from 1895 To 2011

(DCR). We conducted a floristic study of the Fells for several

reasons: 1) the floral survey of the 1890s provided an importantbasis for comparison for change over 116 years (Deane 1896); 2) the

initial survey indicated the existence of a high level of biodiversity;

3) the reservation is a large, isolated fragment in a densely

populated area; 4) the DCR-controlled area has not undergone

logging or other forms of active forestry management since its

creation; 5) a study of part of the reservation in the 1990s indicated

a heavy loss of species (Drayton 1993; Drayton and Primack 1996).

A preliminary survey in 2003 raised some doubts about thesefindings, indicating the need for a more thorough study.

In 1894, the landscaping firm founded by famed landscaper,

Frederick Law Olmsted, was employed to conduct a survey of all

plants in the Fells and the three other forest reservations also

created that year: Blue Hills, Beaver Brook, and Stony Brook. The

Olmsted company gave the job of organizing the survey to its

employee Warren Manning, who assembled a team of over twenty

volunteers to conduct the fieldwork. Where help was needed inidentifying taxonomically difficult groups, Manning engaged the

assistance of local experts, including several at Harvard University,

such as M. L. Fernald, to whom samples were submitted for

determination. Field survey work was conducted during 1894 and

1895 (Manning 1894).

In December of 1895, botanist Walter Deane was hired to

compile and publish the Flora of the Blue Hills, Middlesex Fells,

Stony Brook, and Beaver Brook Reservations (hereafter referred toas Deane’s Flora). Deane’s Flora was published in May 1896, with

the words ‘Preliminary Edition’ on the title page. The 1894–1895

survey vouchers were deposited in the herbarium of the New

England Botanical Club (NEBC), which Deane had helped found in

early 1896, largely as a result of the cooperation developed between

amateur and professional botanists during the survey (Weatherby

1933). In compiling his report, Deane worked from the card index

that was used to collect data from field observations and vouchers.This card index contains many observations on localities and details

of frequency of occurrence not reported by Deane in the Flora, or

on the voucher labels (Anonymous ca. 1896).

After the publication of Deane’s Flora, no further reports were

written about the Fells for 100 years. However, new taxa were

found and vouchered from 1896 onwards, and particularly in the

1920s by Nathaniel Kidder, then President of the NEBC. Kidder

230 Rhodora [Vol. 114

Page 4: Changes In the Vascular Flora of the Middlesex Fells Reservation, Middlesex County, Massachusetts, from 1895 To 2011

vouchered plants that had not been reported in 1895, adding a

further 41 taxa to the Fells flora, while vouchers from otherbotanists added a further 23 taxa in the first half of the 20th

century. As the centenary of the Fells’ formation approached, a

floristic survey was made by Brian Drayton from 1990 through

1992 (Drayton 1993). Drayton confined his area of study to

approximately 40% of the Fells. Unfortunately, the Metropolitan

District Commission did not grant permission for vouchers to be

made to document the findings. Drayton compared what he found

to the results of the 1895 survey and published these findings aftersome minor revisions (Drayton and Primack 1996). As the first

major survey of the flora of the Fells in a century, the 1996 paper

was cited widely for its conclusion that many taxa had been lost

from the area of study.

METHODS

Study site. The Fells reservation comprises parts of the citiesand towns of Malden, Medford, Melrose, Stoneham, and Win-

chester in Middlesex County, Massachusetts (ca. 42u269 to 42u289N

and ca. 71u4.59 to 71u7.59W). The southern boundary is 9 km north

of the city center of Boston, and its northern boundary is 4 km

south of and inside the metropolitan area’s inner ring road,

Interstate Route 95. The Fells is an isolated area of native

vegetation in the highly developed suburban landscape of the

Boston Basin Ecoregion. Almost 80 percent of this ecoregion hasbeen developed, making the remaining conservation areas, totalling

over 2000 ha, critical for maintaining local biodiversity (Friends of

Alewife Reservation 2011; Shafer 1995).

The area of this study included: 1) the forest and pond of

Wright’s Park, now under the jurisdiction of the city of Medford; 2)

the land surrounding the Winchester reservoirs in the west; and 3)

Spot Pond and the Fells Reservoir in the east, which are

administered by the Massachusetts Water Resources Authority(MWRA; Figure 1). Each of these areas had been surveyed as part

of the Fells in the 1890s. The Stone Zoo and the DCR maintenance

yard to the zoo’s north, both areas within the 1895 survey area,

were excluded because they are now separate and greatly disturbed.

The Langwood Commons inholding, which originally encompassed

the Langwood Hotel, Stoneham’s adjoining Whip Hill Park, and

Medford’s Victory Park, none of which were in the original

2012] Hamlin et al.—Middlesex Fells Flora 231

Page 5: Changes In the Vascular Flora of the Middlesex Fells Reservation, Middlesex County, Massachusetts, from 1895 To 2011

1895 survey area, were not included. The area of study was

approximately 1400 ha, larger than the official DCR Fells

Reservation area of 1042 ha because of the inclusion of other

jurisdictional areas which are mostly, but not entirely, open water.

Open water occupies approximately 334 ha and access to the deeper

water of the reservoirs was restricted and therefore could not be

fully explored.

Lawrence Woods (Sector 1) was added to the Fells in 1925,

adding approximately 132 ha that had not been part of the 1895

survey. An approximately equal area of land was later removed

Figure 1. A map of the Middlesex Fells showing the eight sectors with theirboundaries, the three Winchester reservoirs—North, Middle and South—SpotPond, the Fells Reservoir, Lawrence Woods (Sector 1) and Wright’s Park. Thebase map is reproduced with permission from the DCR. For more detailedfeatures in the Fells see the map at www.foundinthefells.com.

232 Rhodora [Vol. 114

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from other parts of the Fells with the construction of Interstate

Route 93 and the taking of other parcels of land around theperimeter.

Physiography, geology, and hydrology. The Fells is an upland

dissected by north-south ridges and valleys. The highest point is

Bear Hill at 96.6 m elevation, which is at the northern end of thecentral ridge. The lowest elevation is 19.8 m, at Gate 1, where

Whitmore Brook exits the reservation. The steep southern scarp in

the eastern part of the Fells is part of the rim of an ancient caldera in

what is now the Boston basin, part of the North Boundary Fault

(Rast and Skehan 1990). Bedrock formations are part of the Avalon

Terrane, which was derived from part of a Paleozoic micro-continent

that accreted to coastal New England and Canada due to continental

drift. This complex tectonic geology consists of Lynn volcanic felsicrhyolites in the south and east, plutonic Dedham granite and

granodiorite in the western and northern parts, and a broad band of

Cambrian metasedimentary rock on top of this layer, running from

the northeast, across the reservation in a west-southwest direction

(Cardoza et al. 1990). The landscape also includes the Medford Dike,

a 100–200 m wide strip of diabase forming a valley between Pine and

Little Pine hills, which was extensively quarried in the 19th century.

The Fells is also cross-hatched with numerous swarms of narrowintrusive mafic dikes (Ross 1990).

Repeated glacial advances and retreats have left a thin layer of

glacial till over the bedrock. Twenty-six soil types occur, with Hollis

Rock Outcrop being the most common (Bitterman et al. 2006). The

last receding glacier also created small kettle hole ponds, such as

Shiner Pool (in Sector 6) and the larger Spot Pond.

The Fells lies within the Mystic River watershed with six small,

and sometimes intermittent, streams from the uplands formingminor tributaries to the river. In the western valley of the

reservation, the three man-made Winchester reservoirs predate

1894 (Figure 1).

Climate. Coastal eastern Massachusetts is in USDA Plant

Hardiness Zone 6a, where the lowest temperatures range from

220.6uC to 223.3uC. Based on data between 1986 and 2002,

average monthly temperatures range from 21.9uC in January to

23.1uC in July. Average annual precipitation in recent years has

been 105 cm. The heaviest precipitation occurs in the months of

November, December, and March, and the driest month is July.

2012] Hamlin et al.—Middlesex Fells Flora 233

Page 7: Changes In the Vascular Flora of the Middlesex Fells Reservation, Middlesex County, Massachusetts, from 1895 To 2011

Precipitation occurs in every month, and varies on average between

a low of 7.1 cm in July, and a high of 10.7 cm in November

(Anonymous 2011; National Oceanic and Atmospheric Adminis-

tration 2011).

Land use history. When the Metropolitan Park Commission

(MPC) created the Middlesex Fells Reservation, much of the land

acquired was composed of woodlots (areas that had been regularly

logged) along with several small farms.

Major construction projects began in the Fells immediately

following the publication of Deane’s Flora. Between 1896 and 1900

the level of Spot Pond was raised three meters, expanding its area

from 105 ha to 119 ha, entailing dam construction, stream

diversion, and the destruction of the pond’s natural littoral, leaving

a shoreline of rip-rap in many places (Olmsted 1900). At the same

time, the smaller Melrose Pond was greatly expanded to create what

is now Basin 1 of the Fells Reservoir (Howard 1900). In the early

years of the reservation, the MPC cleared the forest of ‘‘masses of

dead wood’’ for protection against fire (de las Casas 1898; Wellman

1914). A trolley line was constructed through the Fells. It operated

from 1910 to 1946 and had a stop at the Sheepfold. The rail bed and

two viaducts are still evident (Levin and Mahlstedt 1990).

Currently, the Sheepfold is maintained as an open, grassy area of

approximately four hectares, which is mowed twice a year.

In 1918, the town of Winchester began removing all hardwood

trees for up to 100 m from the shore around the entire perimeter of its

three reservoirs, replacing them in subsequent years with plantings of

conifers, many non-native (Medford Historical Society 1935).

During the 1930s, Works Progress Administration (WPA) and

Civilian Conservation Corps (CCC) crews carried out many projects

in the Fells, including much tree-spraying following outbreaks of

gypsy moth infestations and the Chestnut blight. Projects were

carried out at Bellevue Pond and Spot Pond Brook, involving the

construction of dams and redirection of waterways (Levin and

Mahlstedt 1990). From 1945 to 1950, an army unit manning an anti-

aircraft battery was based on a hill in the northwest corner of

Lawrence Woods. The foundations of a few army huts remain and

the area is currently maintained as meadow habitat for birds and

butterflies. Between 1956 and 1961, the eight-lane Interstate 93

highway was built through the middle of the Fells, splitting the

reservation in two and creating several rock dumps in the forest.

234 Rhodora [Vol. 114

Page 8: Changes In the Vascular Flora of the Middlesex Fells Reservation, Middlesex County, Massachusetts, from 1895 To 2011

In the last two decades of the 20th century, the Fells Reservoir

was greatly expanded with the addition of Basins 2 and 3, involving

major dam construction. Covered reservoirs were built on the top

of Bear Hill and south of the Fells Reservoir. The latter site was

seeded with non-native grasses and forbs. Large increases in human

population density surrounding the Fells have led to increased

human usage and impact, including fragmentation. There are now

110 miles of trails and fire roads fragmenting the woodland—far

more than in the 1890s (Lawrence 1886; Massachusetts Department

of Conservation and Recreation 2010).

In addition to these human impacts, certain natural factors

should be mentioned. In 1938, a major hurricane hit the area and

brought down many trees. With the exception of the tree-cutting by

the town of Winchester surrounding its three reservoirs, most of the

rest of the Fells has experienced ongoing forest maturation since

1894 and, in the last 30 years, the reservation has seen an influx of

deer, coyote, fisher, and beaver (Brown 2009).

Survey methodology. We divided the Fells into eight sectors

(Figure 1). Although we were conscious of the need for approxi-

mate equal size, decisions on sector boundaries were influenced by

the placement of roads and waterways, as well as the need for a

multiple of four, for the four levels of frequency measurement.

What then followed, in effect, were eight mini-surveys (Table 1).

Sector 1 represents Lawrence Woods, which was not included in the

original 1895 survey. Sectors 2, 3, and 4 are together equivalent to

the area surveyed by Drayton (1993).

Searches were mostly conducted individually, with occasional

joint trips which sometimes included specialists. Targeted surveys

for particular taxa, such as Amelanchier spp. and Carex spp., were

also conducted. Areas were criss-crossed between trails, and a

particular effort was made to survey unusual or difficult-to-access

areas such as large, deep swamps. Observations of aquatic taxa

were mostly made from shore, and the MWRA shared with us their

data for Spot Pond and the Fells Reservoir, adding four deep-water

aquatic species to our checklist (MWRA 2006). Although we

became aware of the abundance, or the lack thereof, of many

species, a precise quantitative measurement of abundance for the

868 extant taxa was beyond the scope of this survey.

The survey was conducted over nine years from April 2003 to

September 2011. A log was kept, with the lead surveyor making

2012] Hamlin et al.—Middlesex Fells Flora 235

Page 9: Changes In the Vascular Flora of the Middlesex Fells Reservation, Middlesex County, Massachusetts, from 1895 To 2011

,530 visits to the Fells, each averaging 1.7 hours for a total of

900 hours. The total team hours spent in the Fells is estimated to be

2000 hours. Visits were made on a regular basis from March

through November during the most intense study period from 2005

to 2010.

Nomenclature mostly follows Flora Novae Angliae (Haines 2011),

as well as Flora of North America (Flora of North America Editorial

Committee, eds. 1993+) and Gleason and Cronquist (1991).

Data analysis. At the start of the 1890s survey, Manning

suggested using four frequency categories: common, frequent,

occasional, and rare, and gave definitions to his surveyors

(Manning 1894). However, these frequencies were largely based

on subjective individual assessments, as one person’s ‘occasional’

might be another’s ‘frequent,’ and the levels of surveying and

reporting were inconsistent. In many cases, particularly for

graminoids, no frequency was reported for a taxon, leaving Deane

to later make an assessment from the field reports and vouchers.

When Deane only listed one or two localities from which a plant

was reported, with no specific frequency, we assigned a frequency

value of rare for the taxon. The Deane card index and Flora also

revealed an inconsistency of vocabulary with the use of words and

phrases such as: ‘occurs,’ ‘not uncommon,’ and ‘not generally

distributed but found in many places,’ for which judgements of the

intended frequency had to be made. We attempted to apply a

Table 1. Numbers of plant taxa found in each of 8 sectors in the MiddlesexFells, MA, 2011 survey. Data are arranged in ascending order of size ofapproximate hectares (ha) of land (Land Area), with the total sector area (ha),including open water, in parentheses.

SectorLand Area

(Sector Area) ha

Number of Taxa

Total NativeIntroduced

(%)Total/haof Land

Sector 8 50 (167) 422 266 156 (37) 8.4Sector 5 70 (100) 480 326 154 (32) 6.8Sector 7 105 (110) 488 330 158 (32) 4.6Sector 1 130 (132) 462 319 143 (31) 3.6Sector 4 170 (195) 546 387 159 (29) 3.2Sector 2 180 (245) 480 328 152 (32) 2.7Sector 3 180 (260) 539 362 177 (33) 3.0Sector 6 190 (200) 564 388 176 (31) 3.0All sectors 1075 (1409) 868 563 305 (35) 0.8

236 Rhodora [Vol. 114

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consistent frequency for each Deane taxon, derived from studying

Deane’s Flora in conjunction with his card index. However, Deane’sfrequency categories should be considered more qualitative than

our survey’s more quantitative method, and so the comparisons

have a significant margin of error. Even so, the comparison still

provides a valuable framework for assessing the overall patterns in

changes of frequency. In our survey, frequency was based on the

number of sectors in which a taxon was found: common taxa were

detected in seven or eight sectors; frequent in five or six; occasional

in three or four; and rare in one or two.Native versus introduced status for each taxon was determined

from several sources (Haines 2011; Sorrie and Somers 1999; USDA,

NRCS 2011a) plus discussions with the State Botanist (Bryan

Connolly, Natural Heritage and Endangered Species Program,

Massachusetts Division of Fisheries and Wildlife, pers. comm.

2011). Eleven taxa were considered to be in an intermediate

category for one of three reasons: 1) it was determined that three

species, Betula nigra, Symphyotrichum novae-angliae, and Pyc-

nanthemum incanum, exist in the reservation partly naturally and

partly as plantings; 2) three others—Calystegia sepium, Festuca

rubra, and Prunella vulgaris—are found in the area as both a native

and introduced subspecies, which this survey failed to differentiate

in these taxa; 3) at the time of writing, it was undetermined whether

Acer negundo, Ilex opaca, Lonicera sempervirens, Persicaria hydro-

piper, and Phalaris arundinacea are native or introduced for

Middlesex County, MA.Introduced status was assigned to certain other taxa because we

had reason to believe that they had been planted or seeded into the

study area: Abies balsamea, Asclepias tuberosa, Lobelia siphilitica,

Pinus resinosa, Pycnanthemum torrei, Symphyotrichum pre-

nanthoides, Thuja occidentalis, and Veronicastrum virginicum. For

the purposes of these analyses, the above-mentioned 11 intermedi-

ate-status species were assigned one or the other status, following

discussion with the State Botanist. These allocations (six native andfive introduced) were therefore not made arbitrarily, and at most

would contribute 1% to the error rate of determining native versus

introduced status.

The 1038 taxa of the checklist were compiled, and data were

tracked in a variety of ways, from totals for each sector to the

numbers of taxa that have changed in frequency over the past

century.

2012] Hamlin et al.—Middlesex Fells Flora 237

Page 11: Changes In the Vascular Flora of the Middlesex Fells Reservation, Middlesex County, Massachusetts, from 1895 To 2011

Research of the Deane Flora vouchers and documents. The

Harvard University Herbaria (HUH) were searched for additionalvouchers made since the 1895 survey. This led to the discovery of

Kidder’s work in the Fells in the 1920s, and of other collections in

the early 20th century. The 1895 survey vouchers in the NEBC

herbarium were also studied. The 6th edition of Gray’s Manual

(Watson and Coulter 1890), which was used by Deane et al., was

studied in order to understand some of the differences in

identification and taxonomy compared with today. Several of the

1895 vouchers have been subsequently re-identified as differenttaxa, leading to additions and deletions from those reported by

Deane.

Deane’s vouchers total ,310, and the database with label data,

including synonyms, is available at the Harvard Herbaria website

(Harvard University Herbaria 2011). The vouchers are to be

scanned and their images made available on the website. The

present survey collected ,365 vouchers, including almost all new

native species found. When it was not possible to voucher a plantbecause of insufficient material (e.g., Nabalus serpentarius and

Ranunculus fascicularis), diagnostic photographs were made.

Photographs were also taken of other unvouchered species, with

the date and location recorded. The full photographic record

contains approximately 1700 images documenting over 460 taxa.

Photo images of unvouchered, locally rare, native species have been

placed in the NEBC library.

At the start of the 1895 survey, Manning had stated that the aimwas ‘‘to include in the Flora all plants now growing, or known to

have been growing,’’ (Manning 1894, p. 1). Likewise, George

Davenport, who compiled the ‘Pteridophyta’ section of the report

wrote, ‘‘As the present report is based wholly upon specimens

actually collected, or known to exist in reliable herbaria, a

necessarily incomplete report must be expected’’ (Deane 1896,

p. 107). From this it is clear that the fieldwork was boosted by the

inclusion of earlier private herbaria records and pre-1894 lists fromvarious noted amateur botanists, including William Boott, who had

collected aquatic plants extensively in Spot Pond in the 1850s. For

these reasons, those taxa in the Deane report that were clearly based

on historic specimens and reports prior to 1894 have been excluded.

This is because the Deane report is understood as a flora of the

reservation since its creation in February, 1894. Furthermore, the

term ‘Middlesex Fells’ was used for several years before then, to

238 Rhodora [Vol. 114

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designate a larger area than was eventually incorporated as the

Middlesex Fells Reservation. Lawrence reported the area known as

the Fells as being ‘‘about 4,000 acres’’ or 1600 ha (Lawrence 1886,

p. 200). Other examples of exclusion are planted taxa that persisted

for a short while after the incorporation in 1894 of the several small

farms into the reservation, and various trees and shrubs planted as

ornamentals by the Metropolitan District Commission (MDC) and

the DCR in more recent years. Exclusions also include discovered

or probable errors of identification (Appendix 1).

Plant communities survey. Plant community types in the Fells

were documented using the Natural Heritage and Endangered

Species Program (NHESP) Classification of the Natural Commu-

nities of Massachusetts, version 1.3 (Swain and Kearsley 2001),

hereafter referred to as the NHESP classification. Historical

comparisons were made using a map titled ‘‘Forest Plan for

Middlesex Fells Reservation, 1896’’ (Olmsted and Olmsted 1905),

detailing where woody plant associations occurred at the time.

Habitats were assessed by delimiting 100 m2 plots (10 3 10 m), with

some variation to accommodate odd-shaped habitats such as linear

roadsides and shorelines. Larger plots were used for habitats with

unusually high or low diversity, to capture their full diversity. For

most habitats, additional plots were done to document their

existence. However, since the survey was not able to include

multiple plots for every habitat, none of the plots were chosen

randomly, but were specifically chosen to highlight rare plants and

high-quality habitats. Thus, these plots must be considered to be

more illustrative than statistically definitive. We did not use

NHESP forms, although the data gathered were comparable to

information required by those forms. Observations were made on

ecological aspects including elevation, slope, aspect, hydrology,

bedrock, and soils, along with the presence and abundance of the

plants, to determine the habitat type.

RESULTS

Checklist data. The checklist reports all taxa that have been

found growing, without cultivation, in the Fells since its creation as

an entity in February 1894 (Appendix 2). Exclusions are listed in

Appendix 1, with explanations. It contains eight taxa from Deane’s

Flora, sixteen from Drayton (1993), and eleven from our own list.

2012] Hamlin et al.—Middlesex Fells Flora 239

Page 13: Changes In the Vascular Flora of the Middlesex Fells Reservation, Middlesex County, Massachusetts, from 1895 To 2011

The resultant checklist lists 1038 taxa, of which 868 (84%) were

found to be extant, with 563 native taxa forming 65% of the extant

total (Table 2).

For strict comparison with Deane’s Flora, one should exclude

taxa only found in Sector 1, as this area was added to the

reservation since the 1890s (Table 2). Seventeen taxa were found

solely in Sector 1, 11 being native. Seven of these taxa, including

one non-native, had been reported by Deane in the original area of

the Fells: Bidens cernua, Carex albicans var. emmonsii, Coptis

trifolia, Cornus canadensis, Doellingeria umbellata, Lysimachia

nummularia, and Thalictrum thalictroides. Numbers from all eight

sectors were used for frequency comparisons. The total number of

extant taxa in the Fells was 28% larger than Deane’s checklist,

largely as a result of a big increase in introduced taxa over the

116 years. The number of taxa found in each of the eight sectors

surveyed is shown in Table 1. Sector 2 had the lowest number of

taxa per unit of land area (2.7 taxa/ha), probably because a higher

proportion of its area had been planted to conifers.

Our survey’s total of vascular plants for Sectors 2, 3, and 4

combined, the same area surveyed by Drayton, was 718 (478

native). Drayton and Primack (1996) had reported 331 taxa (244

native). After adjustments for their exclusion of ferns and allies,

graminoids, and aquatics, our comparative figure is 564 taxa (355

native).

As well as being the smallest in land area, Sector 8 is largely

degraded—leading to the highest percentage (37%) of introduced

species. Seventy percent of Sector 8 consists of the open water of

Spot Pond. Because only 13 species are reported from all the open

water of the Fells, the area excluding open water should be noted

when assessing the richness of plant diversity for a sector (Table 1).

Table 2. Comparison of numbers of total, native, and introduced plant taxafound for 1895 and 2011 surveys of Middlesex Fells, MA. % 5 percent oftotal taxa.

Survey

Number of Taxa (% of Total Taxa)

Total Native Introduced

1895 680 570 (84%) 110 (16%)2011 Sectors 2–8 851 552 (65%) 299 (35%)2011 All sectors (1–8) 868 563 (65%) 305 (35%)

240 Rhodora [Vol. 114

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The frequency totals between the 1895 and 2011 surveys,

expressed as percentages, are compared in Table 3. The proportions

of total species that were rare, occasional, frequent, or common

were similar in both surveys. Similarities diverged when the

percentages were separated into native and non-native (Table 3).

In 1895, a lower proportion of the rare taxa were introduced

(14%) compared to the overall proportion of 16%, whereas in 2011

a larger proportion of rare taxa (40%; Table 3) were introduced

compared to the overall proportion of 35% (Table 2). Conversely,

the proportion of common taxa in 1895 that were introduced was

slightly above the overall proportion at 17%, whereas in 2011 the

proportion of common introduced taxa was 29% (Table 3), well

below the overall proportion of 35%. The rare native taxa, asexpressed as a percentage of total native taxa, increased from 27%

in 1895 to 30% in 2011, whereas the percentage of introduced taxa

that were rare went from 23% in 1895 to 37% in 2011 (Table 3).

Of the native taxa that were rare in 1895, 40% could not be

refound in the 2011 survey, 35% stayed rare, and 25% increased in

frequency (Table 4); whereas a large majority (68%) of introduced

species that were rare in 1895 could not be refound (Table 5). Of the

Table 3. A comparison of percent frequency totals of plant taxa found inMiddlesex Fells, MA in 1895 and 2011 surveys. No. 5 number of taxa. TotalNo. taxa in 1895 5 680; in 2011 5 868.

FrequencyCategory andSurvey Year

For Each Frequency Category

Portion ofTotal Taxa

% (No.)

% of Frequency Total % of Total Taxa

Native Introduced Native Introduced

Rare

1895 27% (181) 86% (156) 14% (25) 27% 23%

2011 32% (278) 60% (166) 40% (112) 30% 37%

Occasional

1895 19% (128) 77% (99) 23% (29) 17% 26%

2011 17% (144) 60% (86) 40% (58) 15% 19%

Frequent

1895 19% (127) 89% (113) 11% (14) 20% 13%

2011 17% (152) 68% (103) 32% (49) 18% 16%

Common

1895 36% (244) 83% (202) 17% (42) 35% 38%

2011 34% (294) 71% (208) 29% (86) 37% 28%

2012] Hamlin et al.—Middlesex Fells Flora 241

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native taxa that were already common in 1895, 81% were still either

common or frequent in 2011. All of the 42 introduced species reported by

Deane as common were refound (Table 5), and only 8 of the 202 native

species reported as common in 1895 could not be refound (Table 4).

Although 22% (125) of the total native taxa of 1895 could not be refound

(Table 4), nearly twice as many (41%) either increased in frequency or

stayed common. One hundred and thirty-three new native species have

been reported since the Deane Flora, 118 being extant, leading to a net

reduction of only 1.2% of native species over the last century.

Table 6 shows the breakdown of our findings according to

taxonomic group. A comparison with the same taxonomic

breakdown for the survey of the Greater Mount Holyoke Range

(GMHR; Searcy 2008) shows that despite an almost identical

number of vascular taxa reported (868 for the Fells, 863 for

GMHR), less than half the number (148 c/w 305) of introduced taxa

were found at GMHR (Searcy 2008).

Plant communities. The complex geology and terrain of the Fells

results in a diverse landscape mosaic of thirty habitats, 20 terrestrial

and 10 palustrine, including nine priority habitat communities, four

of which were not previously known to occur in the Boston Basin

Ecoregion. The plant composition of representative habitat plots is

reported by structural layer (Appendix 3).

DISCUSSION

An initial comparison of the 1895 and 2011 survey totals

indicates little net change in the number of native taxa and a

Table 4. Change in number and frequency representation of native planttaxa in Middlesex Fells, MA, over 116 years (1895–2011), shown by changes infour frequency categories. Occas. 5 occasional, Freq. 5 frequent.

FrequencyCategory

Native Species No. (%)

Total1895

Not Re-found in

2011

2011 Total by Frequency Category

Rare Occas. Freq. Common

Rare 156 62 (40%) 55 (35%) 20 (13%) 12 (8%) 7 (4%)Occasional 99 33 19 15 12 20Frequent 113 22 19 14 20 38Common 202 8 15 15 38 126 (62%)Totals 570 125 108 64 82 191

242 Rhodora [Vol. 114

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tripling of invasives (Table 2). For the 2011 survey, the figures for

introduced taxa, ranging from 40% rare to 29% common, are

indicative of new non-natives continuing to arrive (Table 3). Of the

42 non-natives that Deane reported as common, all were refound

and all but three were still either common (74%) or frequent

(Table 5). A further 70 non-natives, reported since 1895, were either

common or frequent by 2011, including several invasive plants.

Non-natives that did become established showed a greater ability to

persist. Of the non-native taxa that were reported as occasional or

more frequent in 1895 (Table 5), 59% increased further in frequency

or stayed common. This compares with only 34% of native taxa

that increased in frequency or stayed common. Of taxa that were

already common in 1895, 74% of non-native taxa stayed common

compared with 62% of native taxa (Tables 4, 5). This continual

arrival and persistence of many non-natives is corroborated by the

steady increase in overall percentage of non-natives from 16% of

total taxa in 1895, to 26% in 1993, and 35% in 2011. At the same

Table 5. Change in number and frequency representation of introducedplant taxa in Middlesex Fells, MA, over 116 years (1895–2011), shown bychanges in four frequency categories. Occas. 5 occasional, Freq. 5 frequent.

FrequencyCategory

Number of Introduced Species (%)

Total 1895

Not Re-found in

2011

2011 Total by Frequency Category

Rare Occas. Freq. Common

Rare 25 17 (68%) 3 (12%) 2 1 2Occasional 29 6 3 7 8 5Frequent 14 0 3 2 3 6Common 42 0 1 2 8 31 (74%)Totals 110 23 10 13 20 44

Table 6. A summary of numbers of taxa found in Middlesex Fells, MA,2011 survey. Numbers of families, genera, and introduced and native taxa arearranged according to taxonomic group.

Taxonomic Group

Number Number of Taxa

Families Genera Native Introduced Total

Ferns and allies 15 20 34 0 34Gymnosperms 3 9 6 11 17Dicotyledons 88 310 376 242 618Monocotyledons 25 95 147 52 199Totals 131 434 563 305 868

2012] Hamlin et al.—Middlesex Fells Flora 243

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time, the high proportion of rare, non-native taxa that were not

refound in this study (68%, compared with 40% of rare native taxa)is an indication of the ephemeral nature of some introduced taxa

(Tables 4, 5).

There are some indications that the figure of 110 (16%) non-

native taxa for the Fells in 1895 is too low. Just 25 years after

Deane’s Flora, Kidder vouchered various common weeds in the

Fells, including Saponaria officinalis and Silene vulgaris, which

Deane had reported for the Blue Hills but not for the Fells. Of the

65 vouchers made since 1895 and in the first decades of the 20thcentury, 42% (27) were non-native. On the other hand, seven of

those 27 were not known in the state until after 1895 (Sorrie 2005).

Sorrie did a thorough study of herbarium records and early floristic

publications and determined that 42% of the state’s 1317

documented non-natives should be considered ‘non-established

waifs,’ and reported an additional 17% for which ‘‘their status as

fully established members of the flora is less certain’’ (Sorrie 2005,

p. 289). Using Sorrie’s determinations, this survey’s checklistcontains 74 introduced taxa (24%) that were considered waifs or

unestablished by Sorrie, or had not been reported in the state until

after 1895. Only seven of these taxa were reported by Deane; of

those, Deane reported five as rare, and only one (Echinochloa crus-

galli) was found in our survey. Of the remaining 67 taxa, 59 were

not known in the state until after 1895. Most notable among

recently established taxa is Alliaria petiolata, first reported in the

state in 1938, described in five or six locations by Drayton in 1993,and found in all eight sectors during this survey, spreading rapidly

in many parts of the Fells (Drayton 1993; Sorrie 2005). Sorrie’s

observation that 59% of the state’s non-natives were not

established, parallels our finding that 68% of the non-natives found

as rare by Deane did not establish themselves and were not refound

by this survey (Sorrie 2005; Table 5).

In contrast, rare native taxa that have persisted as rare have

either done so in habitats that have not been altered, or they havebeen able to adapt to slow forest succession. Two species found in

our survey are protected under the Massachusetts Endangered

Species Act (MESA): Desmodium cuspidatum and Nabalus serpen-

tarius. Eleven others are watch-listed by the state: Betula nigra,

Chenopodium simplex, Coreopsis rosea, Crataegus keepii, Hottonia

inflata, Lespedeza frutescens, Polygonum tenue, Ranunculus alleghe-

niensis, R. fascicularis, R. sceleratus, and Selaginella rupestris

244 Rhodora [Vol. 114

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(Massachusetts Natural Heritage and Endangered Species Program

2011). Two other regionally rare taxa, Platanthera lacera and P.

psycodes, were found growing sympatrically in a Red Maple swamp

with a diverse overstory dominated by Swamp Oak. This habitat

was depicted as marsh on the 1895 vegetation map of the Fells

(Olmsted and Olmsted 1905). The two orchid species now grow in

shade rather than full sun, which may be reducing their ability to

flower and fruit. Deane only reported P. lacera from the western

half of the Fells, where the very small population was not re-located

by this survey. Multiple populations and habitats in a reserve are

critical in order for rare taxa to persist. The large size of the Fells

could contribute to the survival of locally rare taxa, potentially

supporting a higher level of overall diversity than would be possible

in a smaller reserve (Collinge 2009).

Newly found rare, woody taxa of interest were two species of

Crataegus, C. keepii and C. macracantha. This genus has become

rare in the Northeast due to the regrowth of early successional

habitats into mature forest since farming was abandoned in the past

century (Hoover 1961; J. Macklin, HUH, pers. comm.). The two

species are represented by single stations in or on the edge of small

clearings. Another regionally rare woody taxon is Taxus canadensis,

a large population of which was found in Sector 7. Assuming that

such large plants were not overlooked in the 1890s, it is noted that

all three taxa produce fruit attractive to birds and may have been

dispersed over long distances.

Accuracy of reporting. With any plant survey, the questions are

always: how many taxa were overlooked and how many were

misidentified? Archaux et al. (2009) tested the accuracy of field

survey work on woodland plots of 100 m2 and found that, on

average, 15.5% of woody taxa were overlooked in these surveys and

2.3% of species were either not identified or misidentified. They

found that 19.2% of herbaceous taxa were overlooked and 5.3%

were misidentified. Our study of the vouchers collected during the

1895 survey reveal an approximate 4% error rate in identifications,

comparable to Archaux et al.’s (2006) report of a 5% average error

rate. The challenging graminoid genera, Agrostis and Carex, had

the most misidentifications of the 1895 survey vouchers and there

were also some errors in the genus Viola.

There are indications that both the 1895 and the 2011 surveys

under-reported graminoids. Deane reported 124 graminoids (18

2012] Hamlin et al.—Middlesex Fells Flora 245

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introduced) and this survey reports 137 (34 introduced). Of the 106

native graminoids reported by Deane, 40 (38%) were not refound,compared with 19% of non-graminoids not refound. Thirty-eight

new native graminoids have been reported since Deane’s Flora, with

37 extant, representing 32% of total new natives, much higher than

the 18% proportion that native graminoids occupy amongst extant

total native taxa. These figures are all indicative of regular under-

reporting of graminoids.

Deane called the 1895 survey ‘Preliminary,’ and there are

indications of under-reporting, not only of graminoids, but ofwetland plants, non-native weeds, and some trees. It seems unlikely,

for example, that Carya cordiformis was really rare in 1895, as it is

now common, with numerous mature trees. L. L. Dame, who

reported on the trees, was an older botanist who lived south of the

Fells and may have had limited access to other parts of the

reservation. Access to the Fells would have been limited by the lack

of automobiles in the 1890s. There were some carriage roads and

paths within the Fells, but far fewer than a century later. Vouchersfrom the 1895 survey indicate a lack of penetration into the

reservation. In general, aquatic and wetland taxa are also frequently

under-reported, due to the difficulty in accessing their habitats. This

survey found 24 new, native, wetland taxa, a 10% increase in this

group of plants compared with a net loss of 1.2% for native taxa

overall.

In this study, only a small fraction of the approximately 1000 ha

of the land area of the Fells could be closely surveyed. However,with comprehensive plot surveys of particular habitats, it is possible

to be reasonably confident that most species were detected (T.

Rawinski, US Forest Service, pers. comm.). In the process of

surveying plots, rarities that had eluded previous general survey

efforts, such as Paronychia canadensis and Symphyotrichum patens,

were discovered and additional rare taxa were found in the vicinity

of these plots during targeted searches.

The level of expertise of the surveyors, the level of teamwork, andman-hours spent surveying all affect accuracy (Archaux et al. 2006).

A comparative study of 10 urban parks offers some evidence that

total reportings increase when more than one surveyor is involved,

and when more time is spent on the survey (Loeb 2006). In Loeb’s

study, when there was only one surveyor, the average total reports

for three parks was 354 species, compared to an average of 570

species in three other parks when there were two surveyors. In four

246 Rhodora [Vol. 114

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parks where the time spent was either three or four years, the

average total finding was 345 species, compared with an average of

659 species in three other parks where between five and ten years

were spent surveying. The parks involved ranged in size from 400 ha

to 1119 ha, all of sufficient size to potentially support the higher

numbers (Loeb 2006).

Whereas several botanists took part in the 1895 Fells survey, only

two years were invested. Giving details on his survey, Drayton (1993,

pp. 28, 29) wrote ‘‘… the data in the present study were compiled by a

single person, over the course of three field seasons, with an estimated

field time of 300–400 hours.’’ The present study involved four primary

surveyors, for ,2000 hours over nine years. Of the 155 taxa reported

as lost by Drayton and Primack (1996), 105 (68%) were refound by

our survey, 83 within their study area (i.e., Sectors 2, 3, and 4). On the

other hand, Drayton refound at least six species that had been

reported by Deane, but which this survey failed to refind (Drayton

1993). There is concern that the single station of Goodyera pubescens

has disappeared (B. Drayton, TERC, Cambridge, MA, pers. comm.

2010); and Drayton’s single stations of Cornus canadensis and

Verbascum blattaria in Sector 2 have since been eradicated by

maintenance work (B. Drayton and B.T.H., pers. obs. 2010).

Changes in taxonomy have resulted in additions to the flora over

the last century. For example, this survey reports Aronia 3floribunda

as frequent, whereas Deane did not report it. Deane was working

from the 6th edition of Gray’s Manual (Watson and Coulter 1890),

which included the purple-fruited A. 3floribunda with A. arbutifolia,

although without listing it as a synonym. As a result, he did not

report this taxon, although it is quite likely that it occurred in the

Fells at that time. Similarly, use of the 6th edition of Gray’s Manual

may have led to the overlooking of Bidens discoidea, which was not

then recorded for eastern Massachusetts. This species can be

confused with the very common B. frondosa and may have been

overlooked by the Manual authors and therefore, the 1890s

surveyors. Deane only reported ‘Polypodium vulgare,’ whereas today,

two separate species plus their hybrid are recognized. Deane’s

voucher is actually of the hybrid Polypodium 3incognitum. Similarly,

Deane reported Lycopodium obscurum var. dendroideum, the voucher

of which is L. hickeyi, but did not report L. obscurum var. obscurum.

Comparisons with other Massachusetts plant surveys. We com-

pared our findings with three other recent surveys in Massachusetts.

2012] Hamlin et al.—Middlesex Fells Flora 247

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The Greater Mount Holyoke Range, a 3100 ha ridge of hills in

central Massachusetts ,160 km west of the Fells, was the mostpromising for comparisons because both areas are mostly forest

(Searcy 2008; Searcy, Univ. Massachusetts, Amherst, MA, pers.

comm.). The much higher figure for total native taxa in the GMHR

(715 c/w 563) might be expected for a doubling in size of the area

surveyed (Searcy 2008). However, the GMRH study found just 152,

or 18%, non-native taxa compared with 305, or 35%, for the Fells,

resulting in almost identical vascular taxa totals (Searcy 2008).

Similar percentages of non-native taxa to the Fells were found whencomparing to other suburban and urban locations. A study of the

flora of the suburban town of Needham, 20 km southwest of the

Fells, totaling 3250 ha, with 1150 ha ‘‘open and undeveloped,’’

reported 32% non-native taxa with 427 native taxa (Standley 2003).

Likewise, a study of the 9740 ha of the city of Worcester, 80 km to

the west of the Fells, with less than 18% (. 1750 ha) forest, also

reported 32% non-native taxa, with a total of 648 native taxa

(Bertin 2000).Nevertheless, the total of 563 extant native taxa in the Fells

represents a high level of biodiversity for an area of 1400 ha in

suburban Boston. This biodiversity reflects the high diversity of

habitats resulting from the complex bedrock geology and topog-

raphy of the Fells. This survey reports 30 plant communities in the

Fells compared to 20 in the twice-larger Greater Mount Holyoke

Range (Searcy 2008; Appendix 3). The surprising maintenance of

native species in the Fells is shown in comparisons of net losses ofnative species over time: 24% in Needham over a period of 117 years

since 1885 (Standley 2003), 17% in Worcester over a period of

116 years from 1894 (Bertin 2000), 4.5% in GMHR over a period of

180 years (Searcy, pers. comm.), and 1.2% net loss over 116 years

for the Fells. This low net loss may indicate that an equilibrium has

been reached in the overall number of native species that can persist

in the Fells (Hahs et al. 2009). The high percentage of, and steady

increase in, non-native plants in the Fells is indicative of its urbansetting, as well as of high levels of land disturbance, more intensive

human usage, and much fragmentation.

As further evidence of the relative levels of disturbance among

rural forests (GMHR), urban forests, suburbs, and a city, the loss

of orchid species—a family widely reported to be sensitive to

disturbance—over the last century is instructive. There was an 85%

loss of native orchid species over the last century in the Needham

248 Rhodora [Vol. 114

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study, 50% in the Worcester study, 37% in the Fells study, and 28%

in the GMHR study (Bertin 2002; Searcy 2008; Standley 2003).

Ferns were also found to be particularly affected by habitat

destruction in the Needham study, with a 50% loss of species

(Standley 2003). In the 2011 Fells survey, 12 (31%) of the 39 ferns

and related taxa reported by Deane were not refound, compared

with a lower rate of 22% for vascular plants overall.

Causes of species declines and increases: habitat disturbance. The

Fells has numerous introduced taxa, in part, because of the

extensive roadway corridors fragmenting it, with grassy margins

that are more open to non-native establishment than intact native

habitats. Of the 35% of the flora that is introduced, approximately

one third occurs on these margins. Whereas the other two thirds

occur in the interior, many of these are found along fire roads and

trails, and in disturbed areas that resulted from all the road,

reservoir, and tower construction. In 1935, there were 12 miles of

trails in the Fells (Medford Historical Society 1935). In 2010, DCR

reported an estimate of 36 miles of ‘fire roads’ and 75 miles of

official and unofficial trails (DCR 2010).

The Fells is surrounded by a heavily populated urban area, with

non-native species able to enter the Fells from nearby urban

gardens. Heavy recreational activities by hikers and mountain-

bikers also contribute to the spread of invasives when trails are

widened due to trampling of the vegetation, opening the soil to

erosion and colonization. Moreover, the replacement of native

forests around the Winchester Reservoirs with non-native conifers

has caused the single largest destruction of habitat in the Fells. The

inevitable damage to the extensive shorelines of these three

reservoirs, plus the destruction of the littoral of Spot Pond at the

close of the 19th century, resulted in particular loss of native species

from these habitats. Nine of the twelve native wetland species not

refound from the 1895 survey had been reported by Deane from

these now damaged and destroyed locations.

Construction of Interstate 93 caused significant loss of rare fen

habitat that contained plants not refound by this survey, such as

Drosera intermedia, Rhododendron canadense, Sarracenia purpurea,

and Vaccinium macrocarpon. Interstate 93 also created an effective

physical barrier between the two halves of the Fells, especially for

plants with animal-dispersed seeds. The highway is a large corridor

containing almost exclusively invasive plants along its margins, the

2012] Hamlin et al.—Middlesex Fells Flora 249

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most common one being Ailanthus altissima, which is spreading

into the Fells.

Afforestation and mesophication. With forest maturation and

the restoration of some habitats, some rare forest-associates might

be expected to increase in abundance. Of the 156 rare native taxa

reported by Deane, 94 (60%) are extant (Table 4), and 41% of these

have increased in frequency. Herbaceous taxa account for 80% of

the surviving rare taxa, woody taxa being less rare and more

persistent in general. This survey found that Betula papyrifera has

increased from rare to common and Fraxinus pennsylvanica from

rare to frequent. Notable among forbs are Boehmeria cylindrica and

Chimaphila maculata, both now common. Five woodland sedges

also showed a marked increase in frequency. The maturation of the

forest and the increased precipitation due to climate change are two

possible reasons for these increases in frequency.

Shifts in upland forest composition have been documented

throughout eastern North America, from fire-dependent, early

successional taxa, such as Betula spp. and Populus spp., and mid-

successional Pinus strobus and Quercus spp., to fire-intolerant,

shade-tolerant, mid-late successional taxa such as Acer saccharum,

Carya cordiformis, Fagus grandifolia, and Prunus serotina (Clark

1993; Lorimer 2003; Rogers et al. 2008). This change is attributed to

fire suppression, which favors mesic taxa (Nowacki and Abrams

2008). In the Fells, mesic and fire-intolerant taxa such as beech have

significantly expanded their distribution and abundance in the past

century.

In these conditions of a darker, moister forest, the deep-forest

herb Chimaphila maculata has increased from rare to common,

while C. umbellata has increased from frequent to common. Among

forest floor saprophytes, Conopholis americana, singled out in the

introduction to Deane’s Flora as the rarest plant in the Fells, has

increased from rare to frequent, Epifagus virginiana has increased

from rare to occasional, and Monotropa uniflora from frequent to

common. We believe that, taken together, the increase in frequency

of these species is indicative of the maturing of the forest

accompanied by mesophication, where shade-tolerant species are

replacing shade-intolerant species.

One of the concurrent effects of mesophication is an increase in

the number of introduced woody taxa in the understory. One study

in Wisconsin found that introduced taxa in forest plots increased

250 Rhodora [Vol. 114

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from 13 to 31 between 1950 and 2004, or 3.3 new taxa per decade

(Rogers et al. 2008). By comparison, introduced woody taxa in the

Fells increased from nine to 50 over 116 years. At the same time,

native, woody, understory taxa have only increased from 46 to 54

taxa, such that there are now almost as many introduced taxa as

native. A high proportion of these introduced taxa are invasives

such as Euonymus alatus, Frangula alnus, and Rosa multiflora and,

as such, are increasing in abundance. The Wisconsin study found a

correlation between the abundance of invasives with a homogeni-

zation of the herb layer, resulting in more habitat generalists

(Rogers et al. 2008). Among these invasives was Alliaria petiolata,

which is now supplanting native herbs in many sites throughout the

Fells.

Another aspect of mesophication is that the composition of many

forest stands in the Fells are a mixture of xeric and mesic taxa that

do not fit the standard NHESP community classification system

(Appendix 3). Forest homogenization has been attributed to broad-

scale anthropogenic effects of agriculture and wood harvesting in

the 19th century, and fire suppression, urbanization, and introduc-

tion of invasive plants in the 20th century. Modern forest

assemblages have novel compositions that are dissilimar to the

historical colonial ones, and are less closely tied to environmental

gradients than historical forests were. In addition, each taxon has

responded to the various disturbances in individual ways (Foster

et al. 1998). Homogenized forest stands in the Fells sometimes

contain Quercus spp. as equally codominant canopy trees with

either Carya spp. or Fagus grandifolia. Moreover, many understory

trees of Acer saccharum and F. grandifolia now occur in what were

pure stands of White Pine–Oak Forest. This shift indicates that in

many areas, fire-dependent Pinus and Quercus trees are not

regenerating, and future forest stands will be more mesic. With

continued fire suppression, the climax forest in the Fells is likely to

shift toward White Pine–Oak–Beech Forest and Red Oak–Sugar

Maple Transition Forest.

The role of fire. In the 19th century, fire was notably more

widespread than in the 20th century. In 1883 alone, it was reported

that hundreds of acres had burned (Wright 1883). With the advent

of systematic fire suppression in the 1920s (Nowacki and Abrams

2008), fires in the 20th- and early 21st-century time period have

usually been dealt with rapidly, before more than a few acres had

2012] Hamlin et al.—Middlesex Fells Flora 251

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burned. The Fells have probably always experienced a high

incidence of local low-level fires, and these have often been viewed

as destructive to species (Drayton 1993).

Fires occur most frequently on dry upper slopes and summits,

which are naturally prone to, but also maintained by, fire. Fire-

dependent habitats include Rocky Summit and Rock Outcrop

communities, Pitch Pine/Scrub Oak communities, Scrub Oak

Shrublands, and Black Oak–Scarlet Oak Woodlands (Appendix

3). The very rare oak, Quercus prinus, is restricted to a rock bald

subject to frequent fires, which allows the species to persist as young

multi-stemmed trees.

On the upper slopes of Pine Hill, fire is such a common

occurrence that the vegetation consists mainly of grasses, including

Danthonia spicata, Dichanthelium spp., Piptochaetium avenaceum,

and Schizachyrium scoparium. The ash from the fire causes the

grasses to grow lushly, and the abundant grasses catch fire easily,

creating a feedback loop. The frequent fires keep woody growth in

check, providing full sunlight in which grasses thrive. Several rare

taxa occur on the circumneutral talus slope there, and benefit from

this fire regime. These include: Desmodium spp., Eupatorium

sessilifolium, Lespedeza frutescens, Nabalus serpentarius, Pycnanthe-

mum incanum, Symphyotrichum patens, and Thalictrum revolutum.

Fires also create a heterogeneous mosaic of small openings

elsewhere in the forest and in uneven-aged successional stands of

forest, particularly in the southern part of the Fells. Many small

sites have a history of repeated burns, resulting in a perpetual

dominance of early successional species of Betula and Populus spp.,

which resprout after the low-intensity fires. These forest openings

provide cover and browse for wildlife. In sites with mature White

Pine–Oak Forest, the large trees survive small ground fires and act

as seed sources for the regeneration of these trees (Whelan 1995).

Given the widespread mesophication that is occurring in parts of

the Fells, low-level fires should be seen for their benefits in

maintaining early successional diversity in mature forest habitats,

and in maintaining fire-dependent Rock Bald and Pitch Pine

communities (Nowacki and Abrams 2008).

Climate change. Emergence from the Little Ice Age in the late

1800s, plus the anthropogenic rise in global temperatures during the

last 60 years (Zielinsky and Keim 2003), will most likely have put

certain cold-loving plants under stress since the time of the 1895

252 Rhodora [Vol. 114

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survey. Ongoing research in Worcester County, MA, gives

preliminary evidence that the ranges of certain species are shifting

north, in particular Lactuca hirsuta, Moneses uniflora, Pyrola

chlorantha, Rhododendron canadense, and Viola pubescens (R.

Bertin, College of the Holy Cross, Worcester, MA, pers. comm.).

These are all species that were reported by Deane, but which were

not found by the 2011 survey.

The Northeast Climate Impacts Assessment has reported that

since 1970, the Northeast has been warming at a rate of nearly 0.28uCper decade, with winter temperatures rising faster, at a rate of 0.72uCper decade from 1970 to 2000 (Zielinsky and Keim 2003). In 2010, the

Boston area experienced its hottest year since records were first kept

in 1872, with an average temperature for the year of 12.1uC. Snow

cover now lasts for a shorter period, with spring arriving earlier.

Even in the relatively short duration of this nine-year survey, our

photographic records show steadily earlier blooming times for

many spring wildflowers, including Anemome americana, Aquilegia

canadensis, Erythronium americanum, Sanguinaria canadensis, and

Symplocarpus foetidus. This observed trend is corroborated by a

phenological study in nearby Concord, MA (Miller-Rushing and

Primack 2008). According to another study in Concord, MA (Willis

et al. 2010), this earlier bloom time favors invasives.

There has also been a steady increase in precipitation over the last

century in the area. The northeastern US has seen an 8% increase

for the 20th century overall, with the greater proportion of that

increase in the last 40 years, and higher than the regional average

for coastal areas such as eastern Massachusetts (Clean Air-Cool

Planet and Wake 2005). Intense precipitation events—more than

5 cm in 48 hours—have also increased in frequency (Karl and

Knight 1998). Two of the three wettest years recorded for the area

occurred during our survey period. This increase in precipitation is

surely a major factor in the process of mesophication. Predictions

for the future show a steady increase in both warmth and wetness

for the area (Clean Air-Cool Planet and Wake 2005).

Herbivory. Deer and insects are now also negatively impacting

plant species. For example, Deane reported both Lilium canadense

and L. philadelphicum as frequent. We were unable to find the latter

species. Although we found it in five sectors, L. canadense was

under considerable stress and it seldom flowered, due to herbivory

by invasive Scarlet Lily Beetles and browsing by White-tailed Deer.

2012] Hamlin et al.—Middlesex Fells Flora 253

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Both of these herbivores have entered the Fells within the last

50 years. Concern is growing that deer may prove to be one of the

main causes of lily species extinction (Rawinski 2008, 2010). Forest

succession may also be a contributing factor in these declines.

Conclusions. The Middlesex Fells is both unusual and valuable

as a sizeable forest reservation within a large city. Despite various

construction projects over its 117-plus years of existence, much of

the reservation is in a wild and undisturbed state. As a result, and

because of the complex geology of the area, the Fells has an

unusually large number of habitat types and a surprising number of

native plant species. Although the reservation has retained a high

level of biodiversity over the last century, many native taxa survive

in small populations in just one or two locations, making them

vulnerable to the inevitable changes of the coming century. Active

management is needed for the protection of sensitive habitats, the

reduction of fragmentation through trail closures, and invasive

plant removal programs.

ACKNOWLEDGMENTS. More information on the Fells project and

its findings is posted on the website, www.foundinthefells.com,

maintained by B.T.H. The site includes a Fells map in color, with

zoom-in feature, and will report updates on new findings.

We wish to thank Mike Nelson, Dick Stewart, and the

Massachusetts Department of Conservation and Recreation

(DCR) for permission to collect plant samples. We thank DCR’s

archivist Sean Fisher, John Gregoire of the Massachusetts Water

Resources Authority, and their librarians, Mary Lydon and

Rebecca Kenney, and Winchester’s Department of Public Works

for various points of information. We are indebted to Irina Kadis

and Alexei Zinoviev for fieldwork and help with willows, Dr. Lisa

Standley and Dr. Robert Bertin for help with graminoids, Thomas

Rawinski for fieldwork, Arthur Haines for help with Poaceae and

other groups, Dr. Christopher Campbell for help with Amelanchier,

and Dr. Douglas Goldman for help with Quercus hybrids and

Rubus. We want to thank Dr. Karen Searcy and Bryan Connolly

for much editorial assistance, and Dr. Brian Drayton for his advice

and fieldwork. We also wish to thank Dr. Judith Warnement and

Lisa DeCesare for access to the Harvard Botany Libraries, and Dr.

James Macklin, Harvard University, and Ray Angelo and Dr. Les

Mehroff, NEBC Curators, for access to their respective herbaria. We

254 Rhodora [Vol. 114

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thank Dr. Robert Loeb, Pennsylvania State University, for his

encouragement and for sharing documents with us, and Mike Ryan

and Ward Vandewege for technical assistance. In addition, Mike

Ryan, Hue Holley, and Dana Jewell of the Friends of the Middlesex

Fells are appreciated for their encouragement and historical

information on the Fells. Finally, our thanks to people who alerted

us about plants they had seen in the Fells: Anne Hamlin, Joe Hayes,

Ingeborg Jewel, Joyce Krieder, Mia McCollin, Joan Parker, and

Jason Sachs, with apologies to anyone we have missed.

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APPENDIX 1

EXCLUDED TAXA

Taxa that were excluded from Deane’s and Drayton’s floras (Deane 1896;Drayton 1993), and from our own checklist, are listed.

As a result of our study of the vouchers from the 1895 survey, several taxa indifficult groups like Carex, Viola, and the Rosaceae were found to be misidentified.In the case of Rubus canadensis, the voucher is R. flagellaris, whereas the voucher ofRosa carolina is R. palustris. Rosa carolina is retained in the checklist because ofhaving been reported by Drayton, although without a voucher this report cannotbe confirmed. The Viola blanda voucher from the 1895 survey was found to be V.pallens, which is consistent with our findings of this wetland violet. However,Kidder did voucher V. blanda in 1922 in an upland wooded area. Listed here aretaxa that we excluded from Deane’s and Drayton’s floras, including quotationsfrom Deane’s (1896) and Drayton’s (1993) floras and our reason for exclusion ofthese taxa; and cultivated taxa, indicating sector where found, that we had recordedbut excluded from Appendix 2.

(1) Exclusions from Deane’s flora

Barbarea stricta – ‘‘certainly grows here F.S.C.’’ conjecture by Collins(Deane Card Index, ca. 1896).

Botrychium lanceolatum – ‘‘collected some years ago in the Fells,’’ historical.

Botrychium matricariifolium – ‘‘collected once in the Fells’’, historical.

Brassica rapa – cultivated Turnips in field of acquired farm.

Delphinium regalis – ‘‘Persisting for years at rubbish dump near Spot Pond.’’ex cultivation.

Lycopodium clavatum – ‘‘collected in years past by G. E. Davenport.’’historical.

Rosa humilis – unknown identity, no voucher.

Rubus canadensis – Deane followed error in Gray’s Manual, ed. 6. 5

R. flagellaris.

(2) Exclusions from Drayton’s flora

Agalinis purpurea – misinterpreted by Deane, misidentification of A. paupercula.

Larix laricina – no locality reported, probable misidentification forL. decidua.

Linum usitatissimum – ‘‘Winthrop Hill,’’ probable misidentification forL. lineare.

Lonicera canadensis – no locality reported, probable misidentification.

Morus rubra – ‘‘north Pine Hill area,’’ probable misidentification for M. alba.

Nabalus albus – ‘‘Pine & Nanepashemet Hills,’’ 1895 voucher 5

N. serpentarius, not N. albus. Drayton followed Deane 1896.

Picea rubens – no locality reported, probable misidentification of P. glauca.

Ranunculus hispidus – no locality reported, out of range.

Ribes lacustre – ‘‘In three sectors,’’ out of range.

Rubus canadensis – ‘‘Bear Hill, Money Hill, Sheepfold,’’ probablyR. flagellaris, Drayton followed Deane 1896.

Senna hebecarpa – ‘‘Molly’s Spring area,’’ probable misidentification.

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Solidago speciosa – no locality reported, out of range.

Symphyotrichum drummondii – no locality reported, out of range.

Taxus baccata – probable misidentification for T. cuspidata.

Ulmus rubra – ‘‘East side of Bear Hill,’’ probable misidentification.

Viola blanda – ‘‘Silver Mine, Bear Hill,’’ 1895 voucher 5 V. pallens. Drayton

followed Deane 1896.

(3) Exclusions from 2011 list: cultivated plants with sector(s) where found

Cotinus coggygria – Sector 8.

Cotoneaster sp. – Sector 6.

Hamamelis 3intermedia – Sectors 6, 7.

Rhododendron spp. – Sectors 2, 4, 8.

Salix babylonica hybrid – Sectors 5, 8.

Spiraea 3vanhouttei – Sector 2.

Styphnolobium japonicum – Sector 5.

Tilia cordata – Sector 7.

Tulipa sp. – Sector 4.

Viburnum carlesii – Sector 7.

Viburnum opulus var. roseum – Sector 1.

APPENDIX 2

SPECIES FOUND IN THE MIDDLESEX FELLS, MA, 1894–2011

A comprehensive survey of the flora of Middlesex Fells Reservation was

made by the authors over nine years from 2003 to 2011. The area to be surveyed

was divided into eight sectors (numbered 1 to 8; Figure 1). A previous survey of

the flora of the Middlesex Fells area in 1894–1895 was reported by Walter

Deane (Deane 1896), with vouchers deposited in the New England Botanical

Club herbarium (NEBC). The 1894–1895 survey covered approximately our

Sectors 2–8. A floristic survey of approximately 40% of the Fells during 1990 to

1992 (Drayton 1993) covered our Sectors 2–4.

The combined floras from the surveys of Deane, Drayton, and our study are

listed here with additional collection information: Rep. 5 Reporter (who first

reported the taxon): DE 5 Deane, DR 5 Drayton, HA 5 Hamlin et al., MS 5

others. Frequency 5 frequency of occurrence in 1895 and 2011: X 5 Not

Found, R 5 Rare, O 5 Occasional, F 5 Frequent, C 5 Common; * 5 voucher

exists. Sectors 5 sectors in which the taxon was found in the 2011 survey. {denotes not native in the region or probably planted in the Fells; ({) indicates

that the taxon falls into the native/non-native intermediate stage, as discussed in

Methods. Although it was not reported in the Deane Flora, Poa nemoralis has

been credited to the Deane listing, as an 1894 voucher exists for this taxon. It

was presumably overlooked by Deane while compiling the Flora.

260 Rhodora [Vol. 114

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Appendix 2.

Family Taxon Rep.

Frequency Sectors

1895 2011 2011

LYCOPHYTES

HUPERZIACEAE Huperzia lucidula (Michx.) R.Trevis.

HA X R* 4

ISOETACEAE Isoetes echinosporaDurieu subsp. muricata(Durieu) A. & D. Love

DE R* R 6

LYCOPODIACEAE Diphasiastrum digitatumDill. ex A. Braun

DE R* R 8

Lycopodium hickeyi W.H.Wagner, Beitel &R.C. Moran

DE C C* 2–8

Lycopodium obscurum L. HA X C* 2–8SELAGINELLACEAE Selaginella apoda (L.)

C. MorrenHA X R* 7

Selaginella rupestris (L.)Spring

DE R R* 6

MONILOPHYTES

ASPLENIACEAE Asplenium platyneuron(L.) Britton

DE O* R* 3, 7

Asplenium trichomanes L. DE O* X –BLECHNACEAE Woodwardia virginica

(L.) Sm.DE F* F 1–4, 6

DENNSTAEDTIACEAE Dennstaedtia punctilobula(Michx.) T. Moore

DE F* C 1–8

Pteridium aquilinum(L.) Kuhn subsp.latiusculum (Desv.) Hulten

DE C C 1–8

DRYOPTERIDACEAE Dryopteris campyloptera(Kunze) Clarkson

DE R X –

Dryopteris carthusiana(Vill.) H.P. Fuchs

DE O C 1–4, 6–8

Dryopteris clintoniana(D.C. Eaton) Dowell

DE R* R 4, 6

Dryopteris cristata (L.) A.Gray

DE R* R 4

Dryopteris intermedia(Muhl. ex Willd.) A. Gray

DE O* C 1–7

Dryopteris marginalis (L.)A. Gray

DE C* C 1–8

Dryopteris 3boottii (Tuck.)Underw.

DE R* R* 1, 4

Dryopteris 3slossoniaeWherry ex Lellinger

DE R* X –

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Appendix 2. Continued.

Family Taxon Rep.

Frequency Sectors

1895 2011 2011

Dryopteris 3triploidea Wherry HA X R* 1, 4Polystichum acrostichoides

(Michx.) SchottDE C* F 2–4, 6–7

EQUISETACEAE Equisetum arvense L. DE C C 1–8Equisetum fluviatile L. DE O* X –Equisetum hyemale L. subsp.

affine (Engelm.) Calder &Roy L. Taylor

DE R* R* 4

ONOCLEACEAE Onoclea sensibilis L. DE C C 1–8OPHIOGLOSSACEAE Botrychium dissectum Spreng. DE R* X –

Botrychium virginianum(L.) Sw.

DE O* R 4

OSMUNDACEAE Osmunda claytoniana L. DE C C 1–8Osmunda regalis L. var.

spectabilis (Willd.) A. GrayDE C C 1–8

Osmundastrum cinnamomeum(L.) C. Presl

DE C C 1–8

POLYPODIACEAE Polypodium appalachianumHaufler & Windham

HA X F* 2–4, 6–7

Polypodium virginianum L. DE X C* 1–8Polypodium 3incognitum Cusik HA C* C* 1–7

PTERIDACEAE Adiantum pedatum L. DE R X –THELYPTERIDACEAE Parathelypteris noveboracensis

(L.) ChingDE C* C 1–8

Parathelypteris simulata(Davenport) Holttum

DE R R 7

Phegopteris connectilis(Michx.) Watt

DE R X –

Phegopteris hexagonoptera(Michx.) Fee

DE R* X –

Thelypteris palustris Schottvar. pubescens (G. Lawson)Fernald

DE C* C 1–8

WOODSIACEAE Athyrium angustum (Willd.)C. Presl

DE C* C 1–8

Cystopteris tenuis (Michx.)Desv.

DE F* X –

Deparia acrostichoides (Sw.)M. Kato

HA X R* 6

Gymnocarpium dryopteris (L.)Newman

DE R X –

Woodsia ilvensis (L.) R. Br. DE F X –Woodsia obtusa (Spreng.)

Torr. subsp. obtusaDE O* X –

262 Rhodora [Vol. 114

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Appendix 2. Continued.

Family Taxon Rep.

Frequency Sectors

1895 2011 2011

GYMNOSPERMS

CUPRESSACEAE Chamaecyparis thyoides (L.)Britton, Sterns & Poggenb.

DE R X –

Juniperus communis L. var.depressa Pursh

DE F* F 1, 3–4,6–8

Juniperus virginiana L. var.virginiana

DE C* C 1–8

{Thuja occidentalis L. MS X O 2–3, 7–8PINACEAE {Abies balsamea (L.) Mill. DR X R 1, 3

{Larix decidua Mill. HA X O 3–4, 8{Picea abies (L.) Karst. DE O F 2–4, 6–8{Picea glauca (Moench) Voss DR X F* 1–4, 6{Picea pungens Engelm. HA X R 2, 8{Pinus nigra Arnott HA X F 2–5, 7–8{Pinus resinosa Aiton DR X C 1–7Pinus rigida Mill. DE C C 1–8Pinus strobus L. DE C* C 1–8{Pinus sylvestris L. HA X C 1–8{Pseudotsuga menziesii

(Mirb.) FrancoHA X F 2–6, 8

Tsuga canadensis (L.) Carriere DE F C 1–8TAXACEAE Taxus canadensis Marshall HA X R* 7

{Taxus cuspidata Siebold& Zucc.

HA X C 1–8

MAGNOLIIDS

LAURACEAE Lindera benzoin (L.) Blume DE F* C 1–8Sassafras albidum (Nutt.)

NeesDE F C 1–8

MAGNOLIACEAE {Liriodendron tulipifera L. MS X R 1NYMPHAEACEAE Brasenia schreberi J.F. Gmel. DE O R 6

Nuphar variegata Durand exClinton

DE C F 1–3, 5–6

Nymphaea odorata Aiton DE O F* 2–7

TRICOLPATES

ADOXACEAE Sambucus nigra L. subsp.canadensis (L.) R. Bolli

DE C C 1–8

Viburnum acerifolium L. DE C C 1–8Viburnum dentatum L. var.

lucidum AitonDE C C 1–8

{Viburnum dilatatum Thunb. HA X R* 1–2Viburnum lentago L. DE F* F* 1, 3–7

2012] Hamlin et al.—Middlesex Fells Flora 263

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Frequency Sectors

1895 2011 2011

Viburnum nudum L. var.cassinoides (L.) Torr. & A.Gray

DE O R 7

{Viburnum opulus L. subsp.opulus

HA X O* 2–3, 7–8

AMARANTHACEAE {Amaranthus blitum L. subsp.emarginatus (Moq. exUline & Bray) Carreterovar. emarginatus

HA X R 7

{Amaranthus hybridus L.subsp. hybridus

HA X O 4, 7–8

{Amaranthus powellii S.Watson subsp. powellii

HA X R* 3

{Amaranthus retroflexus L. DE O O* 2, 4, 6, 8{Chenopodium album L. HA X C 1–4, 6–8{Chenopodium berlandieri var.

bushianum (Aellen)Cronquist

DE R* X –

{Chenopodium glaucum L.subsp. glaucum

HA X R* 3

Chenopodium simplex (Torr.)Raf.

DE R O* 3, 5–6

{Dysphania ambrosioidesMosyakin & Clemants

HA X R 6, 8

{Dysphania botrys (L.)Mosyakin & Clemants

HA X R 8

ANACARDIACEAE Rhus copallinum L. var.latifolia Engl.

DE O R* 4, 6

Rhus glabra L. DE C* C 1–8Rhus typhina L. DE C C 1–8Toxicodendron radicans (L.)

Kuntze subsp. radicansDE C C* 1–8

Toxicodendron vernix (L.)Kuntze

DE F R 7

APIACEAE {Aegopodium podagraria L. MS X R 1, 7{Aethusa cynapium L. HA X O* 4, 6–7{Carum carvi L. DE O X –Cicuta bulbifera L. DE F R* 7–8Cicuta maculata L. var. maculata DE F* F 1, 4–7{Daucus carota L. DE C C 1–8Osmorhiza claytonii (Michx.)

C.B. ClarkeDE F R 3

Osmorhiza longistylis (Torr.) DC. DE R X –

Appendix 2. Continued.

264 Rhodora [Vol. 114

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Frequency Sectors

1895 2011 2011

{Pastinaca sativa L. DE O X –Sanicula marilandica L. DE C* O 3–4, 6Sium suave Walter DE C* C 1–8

APOCYNACEAE Apocynum androsaemifolium L. DE C* C 1–8Apocynum cannabinum L. DE R* X –Asclepias exaltata L. DE R O 1, 3–4, 6Asclepias incarnata L. var.

pulchra (Ehrh. ex Willd.) Pers.DE C* F 1–6

Asclepias quadrifolia Jacq. DE O O 1, 4, 6{Asclepias speciosa Torr. HA X R 6Asclepias syriaca L. DE C C 1–8{Asclepias tuberosa L. subsp.

tuberosaHA X R 6

{Cynanchum louiseae Kartesz& Gandhi

DR X C* 1–8

{Vinca minor L. MS X C 1–8AQUIFOLIACEAE Ilex mucronata (L.) M. Powell,

Savol. & S. AndrewsDE O X –

({)Ilex opaca Aiton var. opaca HA X R* 2, 4Ilex verticillata (L.) A. Gray DE C C 1–8

ARALIACEAE {Aralia elata (Miq.) Seem. HA X R* 2Aralia hispida Vent. DE C C 1–8Aralia nudicaulis L. DE C C 1–8Aralia racemosa L. subsp.

racemosaDE O* R* 3, 7

{Eleutherococcus sieboldianus(Makino) Koidz

HA X R* 7

{Hedera helix L. HA X F* 1–4, 6–7Hydrocotyle americana L. DE C O* 5–7

ASTERACEAE {Achillea millefolium L.subsp. millefolium

DE C C 1–8

Ageratina altissima (L.) R.M.King & H. Rob. var. altissima

DE R* R* 4, 7

Ageratina aromatica (L.) Spach DE R X –Ambrosia artemisiifolia L. DE C C 1–8Anaphalis margaritacea (L.)

Benth. & Hook. f.DE F* O 1, 4, 6–7

Antennaria howellii Greene subsp.neodioica (Greene) Bayer

MS X X –

Antennaria plantaginifolia (L.)Richardson

DE C* C 1–8

{Anthemis cotula L. DE O X –{Arctium lappa L. DR X F 1, 3, 6–8{Arctium minus Bernh. DE O C* 1–8

Appendix 2. Continued.

2012] Hamlin et al.—Middlesex Fells Flora 265

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Family Taxon Rep.

Frequency Sectors

1895 2011 2011

{Artemisia ludoviciana Nutt.subsp. ludoviciana

HA X R 6

{Artemisia vulgaris L. var.vulgaris

HA X C 1–8

Bidens cernua L. DE F R* 1Bidens connata Muhl. ex Willd. DE F* C 1–7Bidens discoidea (Torr. & A.

Gray) BrittonHA X C* 1–7

Bidens frondosa L. DE C* C* 1–8Bidens vulgata Greene MS X O* 1–3, 7{Centaurea nigra L. DE O* F 1, 3–5,

7–8{Centaurea stoebe L. subsp.

micranthos (Gugler) HayekHA X C 2–8

{Cichorium intybus L. DE C C 1–8{Cirsium arvense (L.) Scop. DE F R 6, 8Cirsium pumilum (Nutt.)

Spreng. var. pumilumDE F* X –

{Cirsium vulgare (Savi) Ten. DE C F 1–2, 5–8{Coreopsis lanceolata L. DR X R 3Coreopsis rosea Nutt. MS X R* 3Doellingeria umbellata (Mill.)

Nees var. umbellataDE F* R* 1

{Echinacea pallida (Nutt.) Nutt. DR X R 6Erechtites hieraciifolia (L.)

Raf. ex DC. var.hieraciifolia

DE C C 1–8

Erigeron annuus (L.) Pers. DE F C 1–8Erigeron canadensis L. DE C C 1–8Erigeron pulchellus Michx.

var. pulchellusDE F X –

Erigeron strigosus Muhl. exWilld. var. strigosus

DE C F* 2–3, 5–7

Eupatorium perfoliatum L. DE C C 1–7Eupatorium pilosum Walter DE R* X –Eupatorium sessilifolium L. HA X R* 4Eurybia divaricata (L.) G.L.

NesomDE F C* 1–8

Eurybia macrophylla (L.) Cass. DE O* C 1–8Eurybia schreberi (Nees) Nees HA X O* 2–3, 6Eurybia spectabilis (Aiton)

G.L. NesomMS X X –

Euthamia graminifolia (L.) Nutt. DE C C 1–8

Appendix 2. Continued.

266 Rhodora [Vol. 114

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Frequency Sectors

1895 2011 2011

Eutrochium dubium (Willd. exPoir.) E.E. Lamont

DE C C* 1–7

Eutrochium purpureum (L.) E.E.Lamont var. purpureum

DE R* R* 5

{Galinsoga quadriradiata Ruiz& Pavon

MS X F 1–2, 4–7

Gnaphalium uliginosum L. DE C* F 1, 4–6, 8{Helianthus annuus L. DE R R 3, 8Helianthus divaricatus L. DE R O* 3–4, 6{Helianthus maximiliani Schrad. MS X X –{Helianthus tuberosus L. HA X R* 5–6{Hieracium aurantiacum L. MS X X –{Hieracium caespitosum Dumort. HA X C* 1–8Hieracium kalmii L. DR X C 1–8{Hieracium lachenalii K.C. Gmel. HA X R* 2Hieracium paniculatum L. DE C* C 1–8{Hieracium pilosella L. HA X O 2–5{Hieracium sabaudum L. HA X C 1–4, 6–8Hieracium scabrum Michx. DE F* C 1–7Hieracium venosum L. DE C* F 1–2, 4–6{Hieracium 3floribundum

Wimmer & Grab.HA X R* 1, 4

Hieracium 3marianum Willd. DE R X –Ionactis linariifolia (L.) Greene DE C* F 1–6Krigia virginica (L.) Willd. DE C O 1–2, 6–7Lactuca biennis (Moench)

FernaldMS X R* 1, 3

Lactuca canadensis L. DE C F 1–3, 6–8Lactuca hirsuta Muhl. ex Nutt. DE R* X –{Lactuca sativa L. MS X X –{Lactuca serriola L. HA X O* 1–3, 6{Lapsana communis L. HA X O* 3–4, 7–8{Leucanthemum vulgare Lam. DE C C 2–8{Matricaria discoidea DC. DR X F* 1, 3–4,

6, 8Mikania scandens (L.) Willd. HA X R* 1Nabalus altissimus (L.) Hook. MS X F 3–8Nabalus serpentarius (Pursh)

Hook.DE R R 4

Nabalus trifoliolatus Cass. DE O C* 1–8Oclemena acuminata (Michx.)

NesomDE C F* 1, 5–8

Packera aurea (L.) A. & D. Love DE C* O 1, 3–4, 6

Appendix 2. Continued.

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Frequency Sectors

1895 2011 2011

Packera paupercula (Michx.)W.A. Weber & A. Love

DE R X –

Pseudognaphaliumobtusifolium (L.) Hilliard &B.L. Burtt

DE F C* 1–8

{Rudbeckia hirta L. var.pulcherrima Farw.

DE F F 1–2, 6–8

{Scorzoneroides autumnalisL.) Moench subsp.autumnalis

DE C C 1–3, 5–8

{Senecio vulgaris L. HA X R 1, 6Sericocarpus asteroides (L.)

Britton, Sterns & Poggenb.DE C* F* 1, 4–6, 8

Sericocarpus linifolius (L.)Britton, Sterns & Poggenb.

DE R* R* 6

Solidago aestivalis E.P. Bicknell DE O X –Solidago altissima L. subsp.

altissimaDE C O 6–8

Solidago arguta Aiton var. arguta DE R O* 2–4, 7Solidago bicolor L. DE C C* 1–8Solidago caesia L. var. caesia DE C C* 1–8Solidago canadensis L. var.

canadensisDE C C 1–8

Solidago flexicaulis L. DE O* O 1, 3–4, 7Solidago gigantea Aiton DE F* C 1–4, 6–8Solidago juncea Aiton DE C* C 1–8Solidago nemoralis Aiton

subsp. nemoralisDE C* C 1–7

Solidago odora Aiton subsp.odora

DE F* O 1, 6, 8

Solidago puberula Nutt. var.puberula

DE C C 1–8

Solidago rugosa Mill. subsp.rugosa

DE C C 1–8

{Solidago sempervirens L.subsp. sempervirens

HA X R 5, 8

Solidago uliginosa Nutt. DE R X –Solidago ulmifolia Muhl. ex

Willd. var. ulmifoliaDE O* O* 4–7

{Sonchus arvensis L. var. arvensis HA X O 1, 3, 6–7{Sonchus asper (L.) Hill DE O F 1–4, 8{Sonchus oleraceus L. DR X C 1–3, 5–8Symphyotrichum cordifolium

(L.) G.L. NesomDE C C* 1–8

Appendix 2. Continued.

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Frequency Sectors

1895 2011 2011

Symphyotrichum dumosum(L.) G.L. Nesom

DE R* O* 5–6, 8

Symphyotrichum ericoides (L.)G.L. Nesom var. ericoides

DE R F* 1–6

Symphyotrichum laeve (L.) A.& D. Love var. laeve

DE C X –

Symphyotrichum lanceolatum(Willd.) G.L. Nesom

DE O* C* 1–3, 5–8

Symphyotrichum lateriflorum(L.) A. & D. Love

DE C F* 1–2,4–5, 7

({)Symphyotrichum novae-angliae (L.) G.L. Nesom

DR X R* 6–7

Symphyotrichum novi-belgii(L.) G.L. Nesom var.novi-belgii

DE C R* 5

Symphyotrichum patens (Aiton)G.L. Nesom var. patens

DE C R* 4

Symphyotrichum pilosum(Willd.) G.L. Nesom var.pringlei (A. Gray) G.L.Nesom

MS X C* 1–8

{Symphyotrichumprenanthoides (Muhl. exWilld.) G.L. Nesom

HA X R* 6

Symphyotrichum puniceum(L.) A. & D. Love var.puniceum

DE F F* 1–3, 6, 8

Symphyotrichum racemosum(Elliott) G.L. Nesom

DE C O* 5, 7–8

Symphyotrichum undulatum(L.) G.L. Nesom

DE C C* 1–7

{Tanacetum vulgare L. DE O C 1–8{Taraxacum laevigatum

(Willd.) DC.DE R* X –

{Taraxacum officinale G.H.Weber ex F.H. Wigg.

DE C* C 1–8

{Tragopogon pratensis L. DR X O 3–4, 6–7{Tussilago farfara L. DR X O 2–3, 5{Xanthium strumarium L. var.

strumariumDE R X –

BALSAMINACEAE Impatiens capensis Meerb. DE C C 1–8BERBERIDACEAE {Berberis thunbergii DC. MS X C 1–8

{Berberis vulgaris L. DE C C 1–8

Appendix 2. Continued.

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Frequency Sectors

1895 2011 2011

{Mahonia aquifolium (Pursh)Nutt.

HA X R 4

BETULACEAE {Alnus glutinosa (L.) Gaertn. HA X O 3–5, 7Alnus incana (L.) Moench

subsp. rugosa (Du Roi)R.T. Clausen

MS X O 3–5, 7

Alnus serrulata (Aiton) Willd. DE C* F* 2, 4–5,7–8

Betula alleghaniensis Britton DE O F 2, 4–7Betula lenta L. DE O* C 1–8({)Betula nigra L. HA X F* 2–4, 6–8Betula papyrifera Marshall DE R C* 1–8Betula populifolia Marshall DE C* C 1–8Carpinus caroliniana Walter

subsp. virginiana(Marshall) Furlow

DE R R 3–4

Corylus americana Walter DE C O 1–4Corylus cornuta Marshall

subsp. cornutaDE O C 1, 3–8

Ostrya virginiana (Mill.) K. Koch DE C C 1–8BIGNONIACEAE {Campsis radicans (L.) Seem.

ex BureauHA X R 1

{Catalpa bignonioides Walter DR X F 1–4, 6, 8BORAGINACEAE {Cynoglossum officinale L. DE R X –

{Echium vulgare L. DE R X –Myosotis laxa Lehm. DE C R 1, 4{Myosotis stricta Link ex

Roem. & Schult.HA X O* 3, 6–7

Myosotis verna Nutt. DE F* R 7–8BRASSICACEAE {Alliaria petiolata (Bieb.)

Cavara & GrandeDR X C 1–8

{Arabidopsis thaliana (L.) Heynh. HA X O* 2–3, 6–7{Barbarea stricta Andrz. MS X X –{Barbarea vulgaris W.T. Aiton DE O C 1–8{Berteroa incana (L.) DC. HA X O 3, 5–6, 8Boechera canadensis (L.) Al-

ShehbazDE R* R* 4

Boechera missouriensis(Greene) Al-Shehbaz

DE O X –

{Capsella bursa-pastoris (L.)Medik.

DE C C 1–7

Cardamine bulbosa (Schreb.ex Muhl.) Britton, Sterns &Poggenb.

DR X X –

Appendix 2. Continued.

270 Rhodora [Vol. 114

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Family Taxon Rep.

Frequency Sectors

1895 2011 2011

Cardamine concatenata(Michx.) Sw.

HA X R* 1

Cardamine parviflora L. var.arenicola (Britton) O.E.Schulz

DE F* R* 3, 7

Cardamine pensylvanicaMuhl. ex Willd.

DE C F 1–4, 6–7

{Conringia orientalis (L.)Dumort.

HA X O* 3, 5, 7–8

{Draba verna L. DR X F 1, 3–5,7–8

{Hesperis matronalis L. HA X O 4, 5, 7{Lepidium campestre (L.)

W.T. AitonHA X C* 2–8

{Lepidium didymum L. HA X R* 2{Lepidium latifolium L. HA X R* 2{Lepidium ruderale L HA X R* 3, 7Lepidium virginicum L. var.

virginicumDE C C 1–8

{Lunaria annua L. HA X R 3{Nasturtium officinale W.T.

AitonDE O F 2–5, 7

{Raphanus raphanistrum L.subsp. raphanistrum

HA X R 6

Rorippa palustris (L.) Besservar. palustris

DE C X –

{Sinapis arvensis L. HA X O* 4–5, 7{Sisymbrium officinale (L.) Scop. DE C O 5–6, 8Turritis glabra L. DR X F* 3–5, 7–8

BUXACEAE {Pachysandra terminalisSiebold & Zucc.

HA X R 2, 4

CALLITRICHACEAE Callitriche heterophylla Purshsubsp. heterophylla

DE R* O* 2, 5–6

Callitriche palustris L. DE C R 7–8CAMPANULACEAE Campanula aparinoides Pursh DE O X –

{Campanula rapunculoides L. HA X F* 1, 4–7Lobelia cardinalis L. DE F O 1, 4–5Lobelia inflata L. DE C* F* 1, 3–4,

6–8{Lobelia siphilitica L. var.

siphiliticaHA X R* 6

Lobelia spicata Lam. var.spicata

DE F X –

Triodanis perfoliata (L.) Nieuwl. DE F* F* 2–7

Appendix 2. Continued.

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Frequency Sectors

1895 2011 2011

CANNABACEAE {Humulus lupulus L. HA X R* 2CAPRIFOLIACEAE Diervilla lonicera Mill. DE C C 1–8

{Lonicera japonica Thunb. DR X O* 1, 3, 7{Lonicera morrowii A. Gray HA X F* 1, 3–7({)Lonicera sempervirens L.

var. sempervirensDE R R* 1, 4

{Lonicera tatarica L. HA X R 1{Lonicera xylosteum L. HA X R* 3–4{Lonicera 3bella Zabel HA X O* 1, 3, 5, 7Triosteum aurantiacum E.P.

Bicknell var. aurantiacumDE O X –

CARYOPHYLLACEAE {Agrostemma githago L. var.githago

DE R* X –

{Arenaria serpyllifolia L. DE O* X –{Atocion armeria (L.) Raf. HA X O 4–5, 8{Cerastium fontanum Baumg.

subsp. vulgare (Hartm.)Greuter & Burdet

DE C F 1, 3,5–6, 8

{Dianthus armeria L. DR X C 1–8{Dianthus barbatus L. HA X R 8{Dianthus deltoides L. HA X R 3{Lychnis coronaria (L.) Murray HA X R* 5{Lychnis flos-cuculi L. subsp.

flos-cuculiHA X R 6

Moehringia lateriflora (L.) Fenzl DE O X –Paronychia canadensis (L.)

Alph. WoodDE R* R* 4, 7

{Saponaria officinalis L. MS X C 1–2, 4–8{Scleranthus annuus L. DE C F 1, 3, 5–7Silene antirrhina L. DE O O* 3–4, 7{Silene latifolia Poir. subsp. alba

(Mill.) Greuter & BurdetHA X C 1–8

{Silene vulgaris (Moench)Garcke subsp. vulgaris

MS X C 1–6, 8

{Spergularia rubra (L.) J. &K. Presl

DE C C 1–8

{Stellaria graminea L. DE O O 3, 6–7{Stellaria media (L.) Vill.

subsp. mediaDE C C 1–8

CELASTRACEAE {Celastrus orbiculatus Thunb. DR X C 1–8Celastrus scandens L. DE F X –{Euonymus alatus (Thunb.)

SieboldDR X C 1–8

{Euonymus europaeus L. HA X F 1–5, 8

Appendix 2. Continued.

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Frequency Sectors

1895 2011 2011

{Euonymus fortunei (Turcz.)Hand.-Mazz.

HA X F* 1–2, 4,6–8

{Euonymus hamiltonianus Wall.var. maackii (Rupr.) Kom.

HA X R* 8

{Euonymus sachalinensis (F.Schmidt) Maxim.

HA X R* 2

CISTACEAE Crocanthemum bicknellii(Fernald) Barnh.

MS X X –

Crocanthemum canadense (L.)Britton

MS X C 1–6, 8

Lechea intermedia Leggett exBritton

DE F* O 1, 3–4, 6

Lechea mucronata Raf. DE C R 7Lechea tenuifolia Michx. DE C* C* 1–8

CLETHRACEAE Clethra alnifolia L. DE C C 1–8CLUSIACEAE Hypericum boreale (Britton)

E.P. BicknellMS X X –

Hypericum canadense L. DE C* F* 2–6, 8Hypericum ellipticum Hook. DE F* X –Hypericum gentianoides (L.)

Britton, Sterns & Poggenb.DE C F 1–4, 6–7

Hypericum majus (A. Gray)Britton

HA X R 2, 8

Hypericum mutilum L. DE C* F* 1–3, 5–6{Hypericum perforatum L. DE C C 1–8Hypericum punctatum Lam. DE R* X –Triadenum virginicum (L.) Raf. DE F C 1–8

CONVOLVULACEAE ({)Calystegia sepium (L.) R. Br. MS X F 2–6, 8{Convolvulus arvensis L. DR X O 1, 3, 7Cuscuta gronovii Willd. ex

J.A. Schult.DE O F* 1, 3–4,

6–8CORNACEAE Cornus alternifolia (L. f.) Small DE C C 1–4, 6–8

Cornus amomum Mill. DE C C 1–8Cornus canadensis L. DE O R 1Cornus florida L. DE O* C 1–8Cornus racemosa (Lam.)

MoldenkeDE C* F* 1–3, 6, 8

Cornus rugosa (Lam.) Rydb. DE R* R 4, 8{Cornus sericea (L.) Holub,

cultivarHA X R* 2

CRASSULACEAE {Hylotelephium telephium (L.)H. Ohba

DE O O 1, 5, 8

{Sedum acre L. HA X R 4{Sedum sarmentosum Bunge HA X R 6

Appendix 2. Continued.

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Frequency Sectors

1895 2011 2011

{Sempervivum tectorum L. HA X R 5CUCURBITACEAE Echinocystis lobata (Michx.)

Torr. & A. GrayDE R R* 7

DROSERACEAE Drosera intermedia Hayne DE F* X –Drosera rotundifolia L. HA X R* 2

ELATINACEAE Elatine minima (Nutt.) Fisch.& C.A. Mey.

DE R* R* 3, 6

ERICACEAE Arctostaphylos uva-ursi (L.)Spreng.

DE R F* 1–2, 4–6

Chamaedaphne calyculata (L.)Moench

DE F R* 7

Chimaphila maculata (L.) Pursh DE R C 1–8Chimaphila umbellata (L.)

W.P.C.Barton subsp.cisatlantica (S.F. Blake)Hulten

DE F* C 1–8

Eubotrys racemosa (L.) Nutt. DE F* F* 3–8Gaultheria procumbens L. DE C C 1–4, 6–8Gaylussacia baccata

(Wangenh.) K. KochDE C C 1–8

Gaylussacia frondosa (L.)Torr. & A. Gray ex Torr.

DE O* C* 1–8

Kalmia angustifolia L. var.angustifolia

DE C C 1–8

Kalmia latifolia L. MS X F 2–4, 6–7{Leucothoe fontanesiana

(Steud.) SleumerHA X R 3

Lyonia ligustrina (L.) DC. var.ligustrina

DE F* C 1–8

Moneses uniflora (L.) A. Gray DE R X –Monotropa hypopithys L. DE F* F* 1–2, 5–7Monotropa uniflora L. DE F C 1–8Pyrola americana Sweet DE F* C* 1–7Pyrola chlorantha Sw. DE R X –Pyrola elliptica Nutt. DE C* C* 1–7Rhododendron canadense (L.)

Torr.DE O X –

Rhododendron viscosum (L.)Torr.

DE C C 1–8

Vaccinium angustifolium Aiton DE C C 1–8Vaccinium corymbosum L. DE C* C 1–8Vaccinium macrocarpon Aiton DE C X –Vaccinium pallidum Aiton DE C C 1–8

Appendix 2. Continued.

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1895 2011 2011

EUPHORBIACEAE Acalypha gracilens A. Gray DE R* R 8Acalypha rhomboidea Raf. DE F C 1–8{Euphorbia cyparissias L. DE F* R 6{Euphorbia esula L. HA X O 5–6, 8Euphorbia maculata L. DE R* F 1–2, 5–8

FABACEAE {Albizia julibrissin Durazz. HA X R 4{Amorpha fruticosa L. MS X R 8Amphicarpaea bracteata (L.)

FernaldDE F C 1–7

Apios americana Medik. DE F C 1–8Baptisia tinctoria (L.) R. Br.

ex Aiton f.DE C* F 1–4, 6

{Cladrastis kentukea(Dum.-Cours.) Rudd

HA X R* 6–7

Desmodium canadense (L.) DC. MS X R* 6Desmodium cuspidatum (Muhl.

ex Willd.) DC. ex LoudonDE R* R* 4

Desmodium glutinosum (Muhl.ex Willd.) Alph. Wood

DE R O 2, 4, 6

Desmodium nudiflorum (L.) DC. DE F* R 1, 6Desmodium obtusum (Muhl.

ex Willd.) DC.DE R* X –

Desmodium paniculatum (L.) DC. DE F* C* 1–8Desmodium perplexum B.G.

Schub.DE R O* 3–4, 6

Desmodium rotundifolium DC. DE O O* 3–4, 6Desmodium 3humifusum

(Muhl. ex Bigelow) BeckHA X R 4, 6

{Genista tinctoria L. HA X R* 6{Gleditsia triacanthos L. DR X C 1–2, 4–8{Laburnum anagyroides Medik. HA X R* 5{Lathyrus latifolius L. HA X R 3Lespedeza capitata Michx. DE F O 3–4, 6Lespedeza frutescens (L.)

Hornem.DE R R* 4

Lespedeza hirta (L.) Hornem.subsp. hirta

DE F* C 1–7

Lespedeza procumbens Michx. DE O* R* 4Lespedeza violacea (L.) Pers. DE C* F* 1, 3–6, 8Lespedeza virginica (L.) Britton DE O* O* 1–2, 4Lespedeza 3nuttallii Darl. DE R* X –{Lotus corniculatus L. HA X C 2–8{Lupinus polyphyllus Lindl.

var. polyphyllusHA X R 6, 8

Appendix 2. Continued.

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Frequency Sectors

1895 2011 2011

{Medicago lupulina L. DE C F* 1–2, 4–5,7–8

{Medicago sativa L. subsp.sativa

HA X O* 5–6, 8

{Melilotus albus Medik. DR X C 1–6, 8{Melilotus officinalis (L.) Lam. HA X O 1, 3, 6{Robinia hispida L. HA X R 2{Robinia pseudoacacia L. DE O F 1, 4–7{Securigera varia (L.) Lassen HA X C* 1–8{Trifolium arvense L. DE C C 1–8{Trifolium aureum Pollich DE F F* 1, 3–4,

6–8{Trifolium campestre Schreb. HA X F* 2–5, 7–8{Trifolium dubium Sibth. HA X R 3, 5{Trifolium hybridum L. DE F F 1, 3, 5–8{Trifolium pratense L. DE C C 1–8{Trifolium repens L. DE C C 1–8{Vicia cracca L. subsp. cracca DE O C 1–8{Vicia sativa L. DE R X –{Vicia tetrasperma (L.) Schreb. DE R O* 2, 5, 7{Wisteria sinensis (Sims) DC. HA X R 2, 8

FAGACEAE Castanea dentata (Marshall)Borkh.

DE F F 1–2, 4,6–7

Fagus grandifolia Ehrh. DE O C 1–8{Fagus sylvatica L. HA X O 2–3, 8Quercus alba L. DE C C 1–8Quercus bicolor Willd. DE C C 1–7{Quercus cerris L. MS X X –Quercus coccinea Muenchh. DE O C 1–8Quercus ilicifolia Wangenh. DE C C 1–8{Quercus palustris Muenchh. HA X R* 5, 8Quercus prinoides Willd. DE O X –Quercus prinus L. HA X R* 4Quercus rubra L. DE C C* 1–8Quercus velutina Lam. DE F C 1–8Quercus 3saulii C.K. Schneid. MS X R* 4–5

GENTIANACEAE Bartonia virginica (L.)Britton, Sterns & Poggenb.

MS X R* 4–5

Gentianopsis crinita (Froel.) Ma MS X X –GERANIACEAE Geranium bicknellii Britton DE R* R* 6

Geranium carolinianum L. DE O* R* 3–4Geranium maculatum L. DE C C 1–8Geranium robertianum L. DE F* F 1, 3–4,

6–7

Appendix 2. Continued.

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Frequency Sectors

1895 2011 2011

HALORAGACEAE {Myriophyllum heterophyllumMichx.

HA X O* 2, 4–5

Myriophyllum humile (Raf.)Morong

DE R O 5–6, 8

Proserpinaca palustris L. DE O* F* 2–6HAMAMELIDACEAE Hamamelis virginiana L. DE C* C 1–8HIPPOCASTANACEAE {Aesculus hippocastanum L. HA X R 7HYDRANGEACEAE {Hydrangea paniculata Siebold HA X R* 1–2

{Philadelphus coronarius L. HA X C 1–6, 8{Philadelphus 3virginalis Rehder HA X R 6

JUGLANDACEAE Carya cordiformis (Wangenh.)K. Koch

DE R C* 2–8

Carya glabra (Mill.) Sweet DE F* C 1–8Carya ovata (Mill.) K. Koch DE C C 1–8Carya tomentosa (Poir. in

Lam.) Nutt.HA X F* 1–6

Juglans cinerea L. DE O X –LAMIACEAE {Ajuga reptans L. HA X O 2, 4, 6–7

{Galeopsis bifida Boenn. HA X R* 3, 7{Glechoma hederacea L. DE R C 1–8Hedeoma pulegioides (L.) Pers. DE C O* 1–2, 4, 7{Lamium galeobdolon (L.) L.

subsp. galeobdolonHA X R* 2

{Lamium purpureum L. HA X O* 2–3, 6–7{Leonurus cardiaca L. DE O F* 1–3, 5, 8Lycopus americanus Muhl. ex

W. BartonDE C F* 1, 5–8

Lycopus uniflorus Michx. DE C* C 1–8Lycopus virginicus L. HA X O 3–4, 6Mentha canadensis L. DE C* O 1, 4–6{Mentha 3piperita L. HA X R* 5{Monarda fistulosa L. HA X R* 6{Nepeta cataria L. DE O R 2{Origanum vulgare L. HA X R 2({)Prunella vulgaris L. DE C* C 1–8({)Pycnanthemum incanum

(L.) Michx. var. incanumHA X R* 4, 6

{Pycnanthemum torrei Benth. HA X R* 6Scutellaria lateriflora L. DE F* C 1–8Teucrium canadense L. DR X X –{Thymus pulegioides L. HA X R* 2Trichostema dichotomum L. DE C F 1, 3–4,

6–8

Appendix 2. Continued.

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Frequency Sectors

1895 2011 2011

LENTIBULARIACEAE Utricularia gibba L. DE R R* 5–6Utricularia macrorhiza Le Conte DE R R 5, 7Utricularia purpurea Walter DE R X –Utricularia radiata Small MS X O 2–5

LINACEAE {Linum perenne L. HA X R* 3Linum virginianum L. DE F X –

LINDERNIACEAE Lindernia anagallidea(Michx.) Pennell

MS X F 2–4, 6, 8

Lindernia dubia (L.) Pennell DE R* R 3, 6LYTHRACEAE Decodon verticillatus (L.) Elliott DE R O 2, 4, 7

{Lythrum salicaria L. DR X C 1–8MALVACEAE {Malva moschata L. MS X X –

{Malva neglecta Wallr. DE O* R 6, 8Tilia americana L. DE F C 1–7

MELASTOMATACEAE Rhexia virginica L. HA X R* 2–3MENYANTHACEAE Nymphoides cordata (Elliott)

FernaldHA X R* 5

MOLLUGINACEAE {Mollugo verticillata L. HA X C 1–8MORACEAE {Morus alba L. HA X C 1–5, 7–8MYRICACEAE Comptonia peregrina (L.) Coult. DE C* C* 1–8

Morella caroliniensis (Mill.)Small

DE C F 1–3, 6, 8

Myrica gale L. DE O X –MYRSINACEAE Lysimachia ciliata L. DE R* R* 2–3

Lysimachia hybrida Michx. DE O X –{Lysimachia nummularia L. DE R R 1Lysimachia quadrifolia L. DE C C 1–8Lysimachia terrestris (L.)

Britton, Sterns & Poggenb.DE C C 2–8

Lysimachia thyrsiflora L. DE R O* 4–7Trientalis borealis Raf. DE C* C 1–4, 6–8

NYSSACEAE Nyssa sylvatica Marshall DE O C 1–8OLEACEAE {Forsythia suspensa (Thunb.)

VahlDR X C 1–8

Fraxinus americana L. DE C C 1–8Fraxinus nigra Marshall DE R O 2, 4–5, 7Fraxinus pennsylvanica Marshall DE R F 1, 3, 5–8{Ligustrum obtusifolium

Siebold & Zucc.HA X O 3–5

{Ligustrum vulgare L. DE F O 1, 4, 7{Syringa vulgaris L. DE O O 1, 5, 8

ONAGRACEAE Chamerion angustifolium (L.)Holub subsp. circumvagum(Mosq.) Kartesz

DE C* X –

Appendix 2. Continued.

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Frequency Sectors

1895 2011 2011

Circaea canadensis (L) Hillsubsp. canadensis

DE O C 1–8

Epilobium ciliatum Raf. DE F* C* 1–8Epilobium coloratum Biehler HA X R* 3–4{Epilobium hirsutum L. HA X R 6Epilobium leptophyllum Raf. DE C X –Ludwigia palustris (L.) Elliott DE C* C 1–8Oenothera biennis L. DE C C 1–5, 7–8Oenothera perennis L. DE C X –

OROBANCHACEAE Agalinis paupercula (A. Gray)Britton

DE C* C 1–6, 8

Agalinis tenuifolia (Vahl) Raf. DE C O* 2, 4–6Aureolaria flava (L.) Farw.

var. flavaDR X O 4–5, 8

Aureolaria pedicularia (L.) Raf. DE F C 1–6, 8Aureolaria virginica (L.) Pennell DE F C* 1, 3–8Conopholis americana (L.) Wallr. DE R* F 3–4, 6–8Epifagus virginiana (L.)

W.P.C. BartonDE R O* 5–8

Melampyrum lineare Desr. DE C C 1–8Orobanche uniflora L. DE O* O 1, 7–8Pedicularis canadensis L.

subsp. canadensisDE F R 1, 6

OXALIDACEAE Oxalis stricta L. DE C C 1–8PAPAVERACEAE Capnoides sempervirens (L.)

Borkh.DE C* C 1–8

{Chelidonium majus L. DE F* C 1–8Sanguinaria canadensis L. DE F* F 3–8

PENTHORACEAE Penthorum sedoides L. DE O O* 2, 4, 6PHRYMACEAE Mimulus ringens L. DE C R 4, 7PHYTOLACCACEAE Phytolacca americana L. var.

americanaDE O C 1–8

PLANTAGINACEAE Chelone glabra L. DE C C 1, 3–8Gratiola aurea Pursh DE F* F 2–4, 6, 8{Linaria vulgaris Mill. DE C C 1–8Nuttallanthus canadensis (L.)

D.A. SuttonDE C C 1–8

{Penstemon digitalis Nutt. exSims

HA X R* 5–6

{Plantago aristata Michx. DR X R* 3{Plantago lanceolata L. DE C C 1–8{Plantago major L. var. major DE C C 1–8Plantago rugelii Decne. DE C* C* 1–7

Appendix 2. Continued.

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Family Taxon Rep.

Frequency Sectors

1895 2011 2011

{Veronica arvensis L. DE R* F 1, 4–5,7–8

{Veronica officinalis L. DE R* C 1–8Veronica peregrina L. HA X R* 3, 8Veronica scutellata L. DE C O* 4–7{Veronica serpyllifolia L. DE C F* 1, 4–6, 8{Veronicastrum virginicum

(L.) Farw.HA X R 6

PLATANACEAE Platanus occidentalis L. DE O O* 1, 4–5POLYGALACEAE Polygala polygama Walter DE C* R* 4, 6

Polygala sanguinea L. DE F* O* 1–4POLYGONACEAE {Fagopyrum esculentum Moench DE R X –

{Fallopia convolvulus (L.) A.Love

DE F* C 1–8

{Fallopia japonica (Houtt.)Ronse Decr. var. japonica

DR X C 1–8

Fallopia scandens (L.) Holub DE O F 1–4, 6Persicaria amphibia (L.) S.F.

Gray var. emersa (Michx.)J.C. Hickman

MS X R* 7

Persicaria arifolia (L.) Haroldson DE F* C 1–4, 6–8Persicaria careyi (Olney) Greene HA X O* 2–3, 6({)Persicaria hydropiper (L.)

OpizDE C* O* 1, 3, 6

Persicaria hydropiperoides(Michx.) Small

DE F* C* 1–2, 4–8

Persicaria lapathifolia (L.)S.F. Gray

DE O* R 4, 8

{Persicaria longiseta (Bruijn)Kitag.

HA X O* 1, 4, 6

{Persicaria maculosa S.F. Gray DE C C* 1–8Persicaria pensylvanica (L.)

G. MazaDR X O* 3, 5, 7

Persicaria punctata (Elliott)Small

DE R R 5–6

Persicaria sagittata (L.) H. Gross DE C F 2, 4–8Persicaria virginiana (L.) Gaertn. HA X R* 6–7{Polygonum aviculare L. HA X C* 1–8Polygonum tenue Michx. HA X R* 6{Rumex acetosa L. DE R X –{Rumex acetosella L. subsp.

pyrenaicus (Pourr. exLapeyr.) Akeroyd

DE C C 1–8

{Rumex crispus L. subsp. crispus DE C C 1–2, 4–8

Appendix 2. Continued.

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Frequency Sectors

1895 2011 2011

{Rumex obtusifolius L. subsp.obtusifolius

DE F* C 1–8

PORTULACACEAE {Portulaca oleracea L. DE C F 2, 4, 6–8PRIMULACEAE Hottonia inflata Elliott DE O* R* 4, 7RANUNCULACEAE Actaea pachypoda Elliott DE R* R 7

Actaea rubra (Aiton) Willd. DE R X –Anemone americana (DC.) H.

HaraDE F* O 3, 5, 7

Anemone quinquefolia L. var.quinquefolia

DE C* C 1–8

Anemone virginiana L. DE F* R 6Aquilegia canadensis L. DE C* F 1–4, 6–7Caltha palustris L. DE C* F 1, 3–4,

6–7Clematis occidentalis

(Hornem.) DC. subsp.occidentalis

MS X X –

Clematis virginiana L. DE F R 3, 7Coptis trifolia (L.) Salisb. DE R R 1Ranunculus abortivus L. DE C* R 3{Ranunculus acris L. DE C F 1–3,

5–6, 8Ranunculus allegheniensis

BrittonDE O* R 3–4

Ranunculus aquatilis L. var.diffusus Withering

DE R X –

{Ranunculus bulbosus L. DE C O 3, 5, 8Ranunculus fascicularis Muhl.

ex BigelowDE F* R 3

{Ranunculus ficaria L. HA X O 2, 6–7Ranunculus flabellaris Raf. DE F R* 4, 6Ranunculus micranthus Nutt. DE O* X –Ranunculus recurvatus Poir.

var. recurvatusDE R* R* 5

{Ranunculus repens L. DE C C 1–8Ranunculus sceleratus L. var.

sceleratusHA X O* 4, 7–8

Thalictrum dioicum L. DE C* F 1–4, 7–8Thalictrum pubescens Pursh DE C* C 1–8Thalictrum revolutum DC. DE R* R 4, 6Thalictrum thalictroides (L.)

A.J. Eames & B. BoivinDE F* R 1

RHAMNACEAE Ceanothus americanus L. DE C* R 2–3{Frangula alnus Mill. HA X C 1–8{Rhamnus cathartica L. DE F C 1, 3–8

Appendix 2. Continued.

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Family Taxon Rep.

Frequency Sectors

1895 2011 2011

ROSACEAE Agrimonia gryposepala Wallr. DE C* F 1–3, 5, 7Amelanchier canadensis (L.)

Medik.DE C* F 1–2, 5–8

Amelanchier intermedia Spach DE C F 2–3, 5–7Amelanchier spicata (Lam.)

K. KochHA X O* 2, 6–7

Aronia arbutifolia (L.) Pers. DE C* F* 1, 5–8Aronia melanocarpa (Michx.)

ElliottDE C C 1–8

Aronia 3floribunda (Lindl.)Spach

HA X F* 1–3, 5–7

Crataegus keepii Sarg. HA X R* 6Crataegus macracantha Lodd.

ex LoudHA X R* 4

{Crataegus monogyna Jacq. HA X R* 8{Cydonia oblonga Mill. DE R X –Fragaria vesca L. subsp.

americana (Porter) StaudtDE R* X –

Fragaria virginiana Duchesne DE C C 1–4, 6–8Geum canadense Jacq. var.

canadenseDE F* C 1–8

Geum rivale L. DE O X –Geum virginianum L. DE F X –{Kerria japonica (L.) DC. HA X R* 8{Malus baccata (L.) Borkh. HA X F 3–8{Malus floribunda Siebold ex

Van HoutteHA X O 1, 4, 7

{Malus pumila Mill. DE R O 1, 3, 6{Malus sieboldii (Regel) Rehd. HA X C 1–7{Potentilla argentea L. DE C C* 1, 3–8Potentilla canadensis L. DE C* C 1–8{Potentilla norvegica L. DE C C 1–6, 8{Potentilla recta L. MS X C 1–8Potentilla simplex Michx. MS X C 1–8{Prunus avium L. DE O O* 2, 4, 6–7{Prunus cerasus L. DE R X –Prunus pensylvanica L. f. var.

pensylvanicaDE F* F 2, 4–8

Prunus pumila L. var.susquehanae (Willd.) H. Jaeger

DE O X –

Prunus serotina Ehrh. var.serotina

DE C* C 1–8

Prunus virginiana L. var.virginiana

DE C* C 1–8

Appendix 2. Continued.

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1895 2011 2011

{Pyrus communis L. DE O R 4{Rhodotypos scandens

(Thunb.) MakinoHA X R* 3, 7

Rosa carolina L. DR X X –{Rosa eglanteria L. DE O X –{Rosa multiflora Thunb. ex

MurrayDR X C 1–8

Rosa palustris Marshall DE C* C 1–8{Rosa rugosa Thunb. HA X R 4Rosa virginiana Mill. DE F F* 2–3, 5–8Rubus allegheniensis Porter MS X C* 1–8Rubus arenicola Blanch. MS X X –Rubus flagellaris Willd. DE C C* 1–8Rubus hispidus L. DE C C 1–6, 8Rubus idaeus L. subsp.

strigosus (Michx.) FockeDE C C 1–5, 7–8

Rubus occidentalis L. DE C* C 1–8Rubus pensilvanicus Poir. DE C F 1–5{Rubus phoenicolasius Maxim. HA X R* 2Rubus pubescens Raf. DE F O* 1, 3–5Rubus setosus Bigelow HA X R* 3, 6{Sorbus aucuparia L. DE F* C 1–8Spiraea alba Du Roi var.

latifolia (Aiton) DippelDE C C 1–8

{Spiraea japonica L. f. var.fortunei (Planch.) Rehd.

HA X F 1–2, 4,6–7

Spiraea tomentosa L. DE C F 2–6RUBIACEAE Cephalanthus occidentalis L. DE F* C 1–8

Galium aparine L. DE F C 1–4, 6–8Galium asprellum Michx. DE F O 3–4, 6, 8Galium circaezans Michx. DE F* F* 1, 3–5, 7Galium lanceolatum Torr. DE F* R* 6{Galium mollugo L. HA X F 1, 3–6, 8Galium palustre L. MS X C* 1–8Galium pilosum Aiton var.

pilosumDE R* X –

Galium tinctorium (L.) Scop. DE C* R* 3–4Galium trifidum L. subsp.

trifidumHA X O* 2–3, 5

Galium triflorum Michx. DE O* R 3Houstonia caerulea L. DE C R 2, 8Houstonia longifolia Gaertn. DE O* X –Mitchella repens L. DE C* C 1–8

Appendix 2. Continued.

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Frequency Sectors

1895 2011 2011

SALICACEAE {Populus deltoides W. Bartramex Marshall var. deltoides

HA X F* 2–6, 8

Populus grandidentata Michx. DE F* C 1–8Populus tremuloides Michx. DE C F 1–3, 5,

7–8{Populus 3jackii Sarg. DE R X –{Salix atrocinerea Brot. HA X F 2–6, 8Salix bebbiana Sarg. DE C X –Salix discolor Muhl. DE C O* 2–3, 5Salix eriocephala Michx.

subsp. eriocephalaDE O C 2–8

Salix humilis Marshall DE F R 2–3Salix lucida Muhl. subsp. lucida DE O X –Salix nigra Marshall DR X F 1, 3–5, 8Salix occidentalis Walter DE O X –Salix pedicellaris Pursh DE O X –Salix petiolaris Smith DE R* R 2, 4{Salix purpurea L. DE R X –Salix sericea Marshall DE R* R 2, 4Salix tristis Aiton DE O* R 2{Salix alba 3fragilis L. HA X R 3–4Salix 3conifera Wangenh. DE R* X –

SANTALACEAE Comandra umbellata (L.)Nutt. subsp. umbellata

DE C C 1–8

SAPINDACEAE ({)Acer negundo L. MS X C 1–4, 6–8{Acer palmatum Thunb. HA X O 2–4, 6Acer pensylvanicum L. HA X O 2, 7–8{Acer platanoides L. MS X C 1–8Acer rubrum L. DE C* C 1–8Acer saccharinum L. MS X O 1–2, 5, 8Acer saccharum Marshall

subsp. saccharumDE C* C 1–4, 6–8

{Acer tataricum L. subsp.ginnala (Maxim.) Wesmael

HA X O 2–3, 8

SARRACENIACEAE Sarracenia purpurea L. subsp.purpurea

DE F X –

SAXIFRAGACEAE Chrysosplenium americanumSchwein. ex Hook.

DE F* F 1, 3–4,6–7

Micranthes pensylvanica (L.)Haw.

DE F X –

Micranthes virginiensis(Michx.) Small

DE C R 2–3

Ribes hirtellum Michx. DE O O* 1, 3–4, 7

Appendix 2. Continued.

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1895 2011 2011

{Ribes nigrum L. HA X R 1{Ribes rubrum L. DE R* R 1, 6Ribes triste Pallas DE R X –

SCROPHULARIACEAE Scrophularia lanceolata Pursh HA X R 2, 6Scrophularia marilandica L. DR X R* 6–7{Verbascum blattaria L. DR X R 3, 7{Verbascum thapsus L. DE C C 1–8

SIMAROUBACEAE {Ailanthus altissima (Michx.)Small

HA X C* 1–8

SOLANACEAE {Nicotiana sp. HA X R 4–5Physalis heterophylla Nees

var. heterophyllaHA X R* 3

{Solanum carolinense L. var.carolinense

MS X F 2–3, 5–7

{Solanum dulcamara L. DE C C 1–8Solanum ptycanthum Dunal MS X C 1–2, 4–8

STYRACACEAE {Halesia carolina L. HA X R* 3ULMACEAE Celtis occidentalis L. HA X O* 2–4

Ulmus americana L. DE C C 1–8{Ulmus glabra Huds. HA X R 7{Ulmus procera Salisb. HA X R* 6

URTICACEAE Boehmeria cylindrica (L.) Sw. DE R C 1–8Pilea pumila (L.) A. Gray var.

pumilaDE R O* 2, 4, 6

{Urtica dioica L. subsp. dioica HA X O* 1, 4, 7Urtica dioica L. subsp. gracilis

(Aiton) SelanderDE C R 3

VERBENACEAE {Verbena bracteata Lag. & Rodr. HA X R* 8Verbena hastata L. var. hastata DE O O 1, 3, 6, 8Verbena urticifolia L. var.

urticifoliaDE F C 1–8

VIOLACEAE Viola blanda Willd. MS X X –Viola cucullata Aiton MS X F 1–6Viola labradorica Schrank DE R X –Viola lanceolata L. subsp.

lanceolataDE C* F 2–6, 8

Viola pallens (Banks ex DC.)Brainerd

DE C* C 1–7

Viola pedata L. DE C* O 1, 4, 6Viola pubescens Aiton DE R X –Viola sagittata Aiton var. ovata

(Nutt.) Torr. & A. GrayDE C C 1–8

Viola sororia Willd. DE C* C 1–8Viola 3primulifolia L. DE R* X –

Appendix 2. Continued.

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Family Taxon Rep.

Frequency Sectors

1895 2011 2011

VITACEAE {Ampelopsis glandulosa var.brevipedunculata (Maxim.)Momiy.

MS X F 3, 5–8

Parthenocissus quinquefolia(L.) Planch.

DE C C 1–8

Vitis aestivalis Michx. DE O* C* 1–2, 4–8Vitis labrusca L. DE C C 1–4, 6–8Vitis riparia Michx. HA X R 4

MONOCOTS

ACORACEAE {Acorus calamus L. DE C R 7ALISMATACEAE Alisma subcordatum Raf. DE R F 1, 4–7

Sagittaria graminea Michx.var. graminea

DE O* R* 3

Sagittaria latifolia Willd. DE F O* 1, 5, 7–8ALLIACEAE Allium canadense L. var.

canadenseDE R* O 1, 3–4

Allium tricoccum Aiton DR X X –{Allium vineale L. HA X C 1, 3–8

AMARYLLIDACEAE {Galanthus nivalis L. subsp.nivalis

HA X R 2

{Narcissus pseudonarcissus L. HA X O 2, 6–7ARACEAE Arisaema triphyllum (L.)

Schott subsp. triphyllumDE C* C 1–8

Arisaema triphyllum subsp.stewardsonii (Britton)Huttleston

MS X X –

Calla palustris L. DE R R 6Lemna minor L. DE C C 1–2, 4–8Peltandra virginica (L.) Raf.

ex SchottMS X F* 3, 5–8

Spirodela polyrrhiza (L.)Schleid.

DE F X –

Symplocarpus foetidus (L.)Salisb. ex W.P.C. Barton

DE C* C 1–8

Wolffia brasiliensis Wedd. HA X R* 3ASPARAGACEAE {Asparagus officinalis L. DE O X –

{Convallaria majalis L. MS X C 1–6, 8Maianthemum canadense Desf. DE C C 1–8Maianthemum racemosum (L.)

Link subsp. racemosumDE C C 1–8

Polygonatum pubescens(Willd.) Pursh

DE C* C 1–8

Appendix 2. Continued.

286 Rhodora [Vol. 114

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Family Taxon Rep.

Frequency Sectors

1895 2011 2011

COLCHICACEAE Uvularia perfoliata L. DE R X –Uvularia sessilifolia L. DE C C 1–8

COMMELINACEAE {Commelina communis L. HA X C 1–8{Tradescantia virginiana L. HA X O* 5–7

CYPERACEAE Bulbostylis capillaris (L.)Kunth ex C.B. Clarke

HA X F* 2–4, 6–7

Carex albicans Willd. exSpreng. var. albicans

HA X O* 4–5, 7

Carex albicans var. emmonsii(Dewey ex Torr.) J. Rettig

DE R* R* 1

Carex annectens (E.P.Bicknell) E.P. Bicknell

DE O* F* 2–6, 8

Carex argyrantha Tuck. DE O* F* 1, 3, 6–8Carex atlantica L.H. Bailey

var. atlanticaDE R* X –

Carex blanda Dewey DE R* O* 1, 4–5, 7Carex bromoides Schkuhr ex

Willd. subsp. bromoidesDE R* R* 4, 6

Carex brunnescens (Pers.)Poir. var. brunnescens

HA X F* 1–2, 5–7

Carex canescens L. subsp.disjuncta (Fernald) Toivonen

DE C* X –

Carex cephalophora Muhl. exWilld.

DE R* C* 1–8

Carex comosa Boott DE O* C* 2–8Carex conoidea Schkuhr ex Willd. DE R* X –Carex crinita Lam. DE F* F* 1–2, 4–7Carex cryptolepis Mack. DE R X –Carex cumulata (L.H. Bailey)

FernaldMS X X –

Carex debilis Michx. var.rudgei L.H. Bailey

DE R* F* 1–5, 8

Carex digitalis Willd. var.digitalis

DE R* R 6

Carex echinata Murr. subsp.echinata

DE O* X –

Carex folliculata L. HA X R* 1Carex gracillima Schwein. DE O* F 1–6Carex intumescens Rudge DE O X –Carex lasiocarpa Ehrh. subsp.

americana (Fernald) D.Love & J.-P. Bernard

DE O X –

Carex laxiculmis Schwein.var. laxiculmis

DE R* R 5

Appendix 2. Continued.

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Family Taxon Rep.

Frequency Sectors

1895 2011 2011

Carex laxiflora Lam. DE F* F* 3–6, 8Carex leptalea Wahlenb.

subsp. leptaleaDE F X –

Carex leptonervia (Fernald)Fernald

HA X R* 3, 5

Carex lucorum Willd. ex Linkvar. lucorum

HA X R* 5

Carex lupulina Muhl. ex Willd. DE O* F* 1–5, 8Carex lurida Wahlenb. DE C* F 1–3, 5–7Carex nigra (L.) Reichard DE R* X –Carex normalis Mack. HA X R* 3, 7Carex pallescens L. DE O* X –Carex pellita Muhl. HA X R* 1Carex pensylvanica Lam. DE C* C* 1–8Carex platyphylla Carey DE R X –Carex projecta Mack. HA X R* 6Carex radiata (Wahlenb.) Small DE R R* 5–6Carex retroflexa Muhl. ex Willd. DE R* R* 4, 6Carex rosea Schkuhr ex Willd. DE R* F* 1, 3–7Carex scabrata Schwein. DE R* X –Carex scoparia Schkuhr ex

Willd. var. scopariaDE C* C* 1–2, 4–8

Carex seorsa Howe DE R* R* 5–6Carex sparganioides Muhl. ex

Willd.DE O X –

{Carex spicata Huds. DE R* X –Carex stipata Muhl. ex Willd.

var. stipataDE F* C 2–8

Carex straminea Willd. exSchkuhr

DE R* R 5

Carex stricta Lam. DE F F* 1–2, 5–8Carex swanii (Fernald) Mack. DE R C* 1–8Carex tenera Dewey DE R X –Carex umbellata Schkuhr ex

Willd.DE C* C* 1–8

Carex utriculata Boott DE R* R* 7Carex vestita Willd. HA X R* 1Carex virescens Muhl. ex Willd. DE R* R* 1, 7Carex vulpinoidea Michx. HA X O* 1, 4, 6–7Cyperus dentatus Torr. DE R* X –Cyperus diandrus Torr. DE R* X –{Cyperus esculentus L. var.

leptostachyus Boeck.HA X O* 3–4, 7

Appendix 2. Continued.

288 Rhodora [Vol. 114

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Family Taxon Rep.

Frequency Sectors

1895 2011 2011

Cyperus lupulinus (Spreng.)Marcks subsp. macilentus(Fernald) Marcks

HA X F* 3–8

Cyperus strigosus L. DE R* F* 1–3, 5, 8Dulichium arundinaceum (L.)

Britton var. arundinaceumDE O* F* 2–6

Eleocharis acicularis (L.)Roem. & Schult.

DE R* F* 2–6

Eleocharis engelmannii Steud. DE R X –Eleocharis flavescens (Poir.)

Urban var. olivacea (Torr.)Gleason

DE R* R* 2, 6

Eleocharis obtusa Schult. DE C* O* 1, 3, 6Eleocharis palustris (L.)

Roem. & Schult.DE R* O* 4–7

Eleocharis tenuis (Willd.) Schult. DE F X –Eriophorum gracile W.D.J.

KochDE R* X –

Eriophorum virginicum L. DE R* X –Fimbristylis autumnalis (L.)

Roem. & Schult.DE R* O* 2–3, 5

Rhynchospora alba (L.) Vahl DE F* X –Rhynchospora capitellata

(Michx.) VahlDE F* R* 3

Schoenoplectus purshianus(Fernald) M.T. Strong

DE R* X –

Schoenoplectustabernaemontani (C.C.Gmel.) Palla

HA X F* 2–7

Scirpus atrovirens Willd. HA X C* 1–8Scirpus cyperinus (L.) Kunth DE O* C 1–7Scirpus microcarpus J. & C.

PreslDE R X –

Scirpus pungens Vahl HA X R* 6ERIOCAULACEAE Eriocaulon aquaticum (Hill)

DruceDE O X –

HEMEROCALLIDACEAE {Hemerocallis fulva (L.) L. DR X C 1–8HOSTACEAE {Hosta sp. HA X F 1, 3–5,

7–8HYACINTHACEAE {Chionodoxa forbesii Baker HA X R 7

{Hyacinthoides non-scripta(L.) Chouard ex Rothm.

HA X R 4

{Muscari botryoides (L.) Mill. HA X R 4{Ornithogalum umbellatum L. MS X R 1, 6

Appendix 2. Continued.

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Family Taxon Rep.

Frequency Sectors

1895 2011 2011

{Othocallis siberica (Haw. exAndr.) Speta

HA X O 2, 4, 7

HYDROCHARITACEAE Elodea canadensis Michx. HA X R* 2, 5Elodea nuttallii (Planch.) H.

St. JohnMS X X –

HYPOXIDACEAE Hypoxis hirsuta (L.) Coville DE C* F 1–6IRIDACEAE {Iris pseudacorus L. HA X O 3–5, 8

Iris versicolor L. DE C C* 1–2, 4–8Sisyrinchium atlanticum E.P.

BicknellDE F* X –

Sisyrinchium montanum Greene MS X O* 3–4, 6, 8JUNCACEAE Juncus acuminatus Michx. DE O* O* 2–3, 5, 8

Juncus articulatus L. DE R* X –Juncus brevicaudatus

(Engelm.) FernaldDE R* X –

Juncus bufonius L. HA X R* 6Juncus canadensis J. Gay ex

LaharpeDE R* O* 2, 5–6, 8

Juncus effusus L. DE C C 1–8Juncus greenei Oakes & Tuck. HA X C* 2–8Juncus marginatus Rostk. DE R* R* 3Juncus pelocarpus E. Mey. DE O* F* 2–4, 6, 8Juncus tenuis Willd. DE O* C 1–8Luzula multiflora (Ehrh.) Lej.

subsp. multifloraDE C* C 1–8

LILIACEAE Erythronium americanum Ker-Gawl. subsp. americanum

DE R R* 1, 4

Lilium canadense L. DE F F 1–4, 6Lilium philadelphicum L. DE F X –Medeola virginiana L. DE C C 1–7

MELANTHIACEAE Trillium cernuum L. DE F* O 1–4{Trillium grandiflorum

(Michx.) Salisb.HA X R 7

Veratrum viride Aiton subsp.viride

DE F O 1, 3–4

NAJADACEAE Najas flexilis (Willd.) Rostk.& W.L.E. Schmidt

DE R* R* 5

Najas gracillima (A. Braun)Magnus

DE R X –

ORCHIDACEAE Corallorhiza maculata (Raf.)Raf.

DE O X –

Corallorhiza trifida Chatelain DE R R* 7Cypripedium acaule Aiton DE F C 1–8{Epipactis helleborine (L.) Crantz HA X F* 2–4, 6–8

Appendix 2. Continued.

290 Rhodora [Vol. 114

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Family Taxon Rep.

Frequency Sectors

1895 2011 2011

Goodyera pubescens (Willd.)R. Br.

DE O X –

Platanthera hookeri (Torr. exA. Gray) Lindl.

DE R X –

Platanthera lacera (Michx.)G. Don

DE R R* 5

Platanthera psycodes (L.)Lindl.

DE R R* 5

Spiranthes cernua (L.) Rich. DE C* O* 1, 3, 5–6Spiranthes lacera (Raf.) Raf.

var. laceraDE F* X –

POACEAE {Agrostis capillaris L. DE R* X –{Agrostis gigantea Roth HA X R* 3, 6Agrostis hyemalis (Walter)

Britton, Sterns & Poggenb.DE F* X –

Agrostis perennans (Walter)Tuck.

DE C* C* 1–7

Agrostis scabra Willd. HA X R* 2, 4{Agrostis stolonifera L. DE O* C* 2–8Alopecurus aequalis Sobol. DE R* X –{Alopecurus geniculatus L. DE R* X –{Alopecurus pratensis L. DE F R* 2Andropogon virginicus L. var.

virginicusHA X R* 8

Anthoxanthum nitens (Weber)Y. Schouten & Veldkampsubsp. nitens

DE R* X –

{Anthoxanthum odoratum L. DE O O 1, 3, 5, 7Aristida dichotoma Michx.

var. dichotomaHA X O* 2–4, 6

{Arrhenatherum elatius (L.) J.& C. Presl

MS X R* 2, 5

{Avena sativa L. MS X X –Brachyelytrum aristosum

(Michx.) Trel.HA X O* 1–2, 4, 6

{Bromus arvensis L. HA X R* 3Bromus pubescens Muhl. ex

Willd.DE O* R* 3–4

{Bromus secalinus L. DE R* X –{Bromus tectorum L. HA X F* 1, 3, 5–8Calamagrostis canadensis

(Michx.) P. Beauv.DE F O* 1, 5, 7–8

Calamagrostis cinnoides(Muhl.) W.P.C. Barton

DE R* R* 6

Appendix 2. Continued.

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Family Taxon Rep.

Frequency Sectors

1895 2011 2011

{Calamagrostis epigejos (L.)Roth var. georgica (K.Koch) Ledeb.

HA X R 4

Cinna arundinacea L. DE O* C* 1–7{Dactylis glomerata L. DE C C 1–8Danthonia compressa Austin

ex PeckDE O* R* 5, 7

Danthonia spicata (L.) P.Beauv. ex Roem. & Schult.

DE C* C* 1–8

Deschampsia flexuosa (L.) Trin. MS X F* 3–8Dichanthelium acuminatum

(Sw.) Gould & C.A. Clarksubsp. columbianum (Scribn.)Freckmann & Lelong

DE R X –

Dichanthelium acuminatumsubsp. fasciculatum (Torr.)Freckmann & Lelong

DE R* C* 1–8

Dichanthelium acuminatumsubsp. implicatum (Scribn.)Freckmann & Lelong

DE R F* 2, 4–6, 8

Dichanthelium acuminatumsubsp. spretum (Schult.)Freckmann & Lelong

DE O X –

Dichanthelium clandestinum(L.) Gould

HA X C* 1–8

Dichanthelium commutatum(Schult.) Gould subsp.ashei (G. Pearson ex Ashe)Freckmann & Lelong

DE R* R* 4

Dichanthelium depauperatum(Muhl.) Gould

HA X R* 4, 6

Dichanthelium dichotomum(L.) Gould

DE O* F* 1–2,4–6, 8

Dichanthelium latifolium (L.)Harvill

DE O* O* 4, 6–7

Dichanthelium linearifolium(Scribn.) Gould

DE R* F* 2–3, 6–8

Dichanthelium oligosanthes(Schult.) Gould subsp.scribnerianum (Nash)Freckmann & Lelong

HA X R* 2, 5

Dichanthelium sphaerocarpon(Elliott) Gould

DE O* F* 2–3, 5–7

{Digitaria sanguinalis (L.) Scop. HA X F* 2–3, 6–8

Appendix 2. Continued.

292 Rhodora [Vol. 114

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Family Taxon Rep.

Frequency Sectors

1895 2011 2011

{Echinochloa crus-galli (L.) P.Beauv.

DE F O 1, 3–5

Echinochloa muricata (P. Beauv.)Fernald var. muricata

HA X O* 3–4, 6

Elymus hystrix L. DE R O* 2–4, 7{Elymus repens (L.) Gould DE F* C 1–6, 8Elymus trachycaulus (Link)

Gould ex Shinners subsp.glaucus (Pease & Moore)Cody

DE R* X –

Eragrostis capillaris (L.) Nees HA X R* 4, 6{Eragrostis pilosa (L.) P.

Beauv. var. pilosaHA X F* 2–4, 6–7

Eragrostis spectabilis (Pursh)Steud.

HA X C* 1, 3–8

{Festuca filiformis Pourr. HA X O* 1, 6–8{Festuca ovina L. MS X O* 3, 5–6({)Festuca rubra L. HA X O* 3, 5, 8Festuca subverticillata (Pers.)

AlexeevDE R* X –

Glyceria acutiflora Torr. DE O* R 2Glyceria borealis (Nash) Batch. DE R* X –Glyceria canadensis (Michx.)

Trin.DE F F 2–3, 5,

7–8Glyceria grandis S. Watson

var. grandisHA X R* 1, 7

Glyceria striata (Lam.)Hitchc.

DE F* C* 1–7

{Holcus lanatus L. MS X R* 4Leersia oryzoides (L.) Sw. HA X C* 1–2, 4–8Leersia virginica Willd. HA X F* 1–5, 7{Lolium perenne L. HA X F* 1–5, 8Muhlenbergia frondosa (Poir.)

FernaldHA X O* 2–3, 6

Muhlenbergia schreberi J.F.Gmel.

MS X R* 2

Muhlenbergia sobolifera(Muhl. ex Willd.) Trin.

HA X R* 4

Muhlenbergia sylvatica Torr.ex A. Gray

DE O X –

Muhlenbergia tenuiflora(Willd.) Britton, Sterns &Poggenb.

DE O* X –

Appendix 2. Continued.

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Family Taxon Rep.

Frequency Sectors

1895 2011 2011

Muhlenbergia uniflora (Muhl.)Fernald

DE R* X –

Oryzopsis asperifolia Michx. DE R* X –Panicum capillare L. subsp.

capillareHA X R* 3, 6

Panicum dichotomiflorum Michx. HA X O* 1, 3–4, 6Panicum rigidulum Bosc ex Nees DE O* O 3–5, 8Paspalum setaceum Michx.

var. muhlenbergii (Nash)D.J. Banks

DE O* X –

({)Phalaris arundinacea L. HA X F* 2–6, 8{Phleum pratense L. DE C* C* 1–8{Phragmites australis (Cav.)

Trin. ex Steud. subsp.australis

HA X C 1–8

{Phyllostachys dulcis McClure HA X R* 1Piptatherum racemosum (Sm.)

EatonDE R* X –

Piptochaetium avenaceum (L.)Parodi

MS X R* 4, 6

{Poa annua L. DE C C 1–8{Poa compressa L. DE C C 1–8{Poa nemoralis L. DE O F* 1, 3–5, 7Poa palustris L. DE C* F* 1–6{Poa pratensis L. DE C* F 2–4, 6–8{Poa pratensis L. subsp.

angustifolia (L.) Arcang.MS X R* 3, 5

{Schedonorus pratensis(Huds.) P. Beauv.

DE C* C* 1, 3–8

Schizachyrium scoparium(Michx.) Nash var.scoparium

DE C C 1–8

{Setaria faberi Herrm. HA X F* 2–4, 6–8{Setaria pumila (Poir.) Roem.

& Schult. subsp. pumilaDE O O* 3, 5–6

{Setaria viridis (L.) P. Beauv.var. viridis

DE O* F* 1–6

Sphenopholis nitida (Biehler)Scribn.

DE R* X –

Torreyochloa pallida (Torr.)Church var. pallida

DE F* R 5

{Tridens flavus (L.) Hitchc.var. flavus

HA X R* 6

{Vulpia myuros (L.) C.C. Gmel. HA X R* 5

Appendix 2. Continued.

294 Rhodora [Vol. 114

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APPENDIX 3

PLANT COMMUNITIES OF THE MIDDLESEX FELLS

The natural communities of the Fells are listed in two groups: palustrine (10)

and terrestrial (22). Each title denotes the NHESP classification community

name and NHESP ‘‘element rank’’ from Critically Imperiled in Massachusetts:

S1, to Secure: S5. The S1 to S3 habitats are called ‘‘Priority Natural

Communities,’’ which are considered to be exemplary and uncommon, and to

require monitoring for their conservation. This system is based on the species

element ranking developed for the Natural Heritage system by The Nature

Conservancy (Swain and Kearsley 2001). Six community types are recognized

here which were not included in the NHESP classification, for which a name

was assigned and is shown in brackets. Two new community types were found,

which were recognized and named by NHESP.

For each community, there are four entries: 1) Community name and

NHESP rank, followed by site description; 2) Study site location and elevation

in the Fells; 3) Life forms and Wetland Indicator Status (WIS). This includes

Family Taxon Rep.

Frequency Sectors

1895 2011 2011

PONTEDERIACEAE Pontederia cordata L. DE C O 3, 5, 7POTAMOGETONACEAE Potamogeton amplifolius Tuck. HA X R* 2–3

Potamogeton bicupulatusFernald

MS X R* 4

{Potamogeton crispus L. HA X R 6Potamogeton epihydrus Raf. DE O R* 3, 5Potamogeton foliosus Raf.

subsp. foliosusHA X O 2–3, 7

Potamogeton natans L. DE R R 3Potamogeton perfoliatus L. HA X R 5–6Potamogeton pulcher Tuck. HA X R* 2, 6Potamogeton pusillus L. HA X R 8Potamogeton spirillus Tuck. DE R* X –Potamogeton vaseyi J.W. Robbins DE R X –Stuckenia pectinatus (L.) Boerner HA X R 6

SMILACACEAE Smilax herbacea L. DE F C 1–8Smilax rotundifolia L. DE C C 1–8

TYPHACEAE Sparganium americanum Nutt. HA X C* 1–7Sparganium emersum Rehmann DE F X –Typha angustifolia L. HA X F* 1, 4–5,

7–8Typha latifolia L. DE C C 1–8

XYRIDACEAE Xyris difformis Chapm. var.difformis

DE F X –

Appendix 2. Continued.

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the number of taxa in each life form, total taxa, and number of taxa native orintroduced, followed by the number of taxa in each WIS category, from the driest

to the wettest: Upland ‘UPL’, Facultative Upland ‘FACU’, Facultative ‘FAC’,Facultative Wetlands ‘FACW)’ and Obligate Wetlands ‘OBL’ (USDA, NRCS2011b), reflecting the overall hydrology of the plot; and 4) A list of taxa present in

each of the tree, shrub, and herb layers. The tree layer is equivalent to thecombined Canopy and Sub-canopy terms used in the NHESP classification.

Palustrine Communities

Acidic Graminoid Fen (S3). An herbaceous community dominated by sedges,

with standing water year round from groundwater and/or surface inflow. Thishabitat is rare in the Fells.

Study site: East of Doleful Pond, in Sector 7, Stoneham, 42u28960N, 71u59220W,46 m elevation.

Life forms: Trees-0, Shrubs-3, Vines-0, Forbs-3, Graminoids-1, Ferns-0. Totaltaxa: 7, 6 Native (86%), 1 Introduced (14%). WIS: UPL-0, FACU-0, FAC-0,FACW-2, OBL-5.

Shrub Layer: Chamaedaphne calyculata, Rosa palustris, Spiraea tomentosa.Herb Layer: Carex utriculata, Decodon verticillatus, Lythrum salicaria,

Polygonum hydropiperoides.

Alluvial Red Maple Swamp (S3). These seasonally flooded, flat, closed-canopyareas with silt muck soils are commonly found along the larger streams in theFells. Red Maple Swamps can also experience overbank flooding from streams,

but these occur in undrained basins with no inlet or outlet, whereas the alluvialvariant has direct hydrological connections with the stream. This connectioncan result from being confluent when there are no streambanks, or when there

are openings in the bank that act as both inlets during flooding, and outletssubsequent to flooding.

Study site: Straight Gully Brook downstream of Bellevue Pond on the southside of Pine Hill, Medford, 42u259510N, 71u69230W, 21 m.

Life forms: Trees-3, Shrubs-5, Vines-0, Forbs-6, Graminoids-2, Ferns-2. Totaltaxa: 18, 16 Native (89%), 2 Introduced (11%). WIS: UPL-0, FACU-2, FAC-2,FACW-10, OBL-4.

Tree Layer: Acer platanoides, A. rubrum, Ulmus americana. Shrub Layer:

Lindera benzoin, Rosa multiflora, Sambucus nigra var. canadensis, Toxicoden-

dron radicans, Viburnum dentatum var. lucidum. Herb Layer: Arisaema

triphyllum, Cardamine pensylvanica, Chelone glabra, Impatiens capensis,Mimulus ringens, Onoclea sensibilis, Osmundastrum cinnamomeum, Phalaris

arundinacea, Poa palustris, Symplocarpus foetidus.

Deep Emergent Marsh (S4). These marshes commonly occur along the marginsof ponds, lakes and slow moving streams. They have standing water, 0.5–3 mdeep, and muck soils, with tall herbaceous plants dominating.

Study site: Doleful Pond, in Sector 7, Stoneham, 42u28940N, 71u59370W, 44 m.

Life forms: Trees-0, Shrubs-0, Vines-0, Forbs-4, Graminoids-2, Ferns-1. Totaltaxa: 7, 5 Native (71%), 2 Introduced (29%). WIS: UPL-0, FACU-0, FAC-0,

FACW-2, OBL-5.

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Herb Layer: Cicuta bulbifera, Eleocharis palustris, Lythrum salicaria, Phrag-mites australis, Sagittaria latifolia, Thelypteris palustris var. pubescens, Typhalatifolia.

[Deep Emergent/Submersed Lake/Pond Community]. There are several kettleponds, in addition to the reservoirs, with floating-leaved plants. Differentsubmersed plants occur in deeper more open water, depending on the pH,alkalinity, and nutrients in the water column.Study site: Shiner Pool in Sector 6, Medford 42u269490N, 71u059250W, 66 m.Life forms: Trees-0, Shrubs-0, Vines-0, Forbs-5, Graminoids-0, Ferns-0. Totaltaxa: 5, 5 Native (100%), 0 Introduced (0%). WIS: UPL-0, FACU-0, FAC-0,FACW-0, OBL-5.Herb Layer: Brasenia schreberi, Nuphar variegata, Nymphaea odorata,Potamogeton pulcher, Utricularia gibba.

Inland Acidic Pondshore/Lakeshore (S4). Sandy shoreline areas are mostlysmall in area, whereas muddy pond shores occur where brooks enter ponds.Herbaceous plants dominate, with graminoids being very common. This habitatis uncommon partly due to the use of rip-rap around the reservoirs, and alsobecause many shorelines are bedrock or wooded banks.Study site 1: Spot Pond, Stoneham, 42u27952.30N, 71u6910W, 47 m; a sandybeach in the Straw Point cove on the north shore.Life forms: Trees-0, Shrubs-2, Vines-0, Forbs-15, Graminoids-2, Ferns-0. Totaltaxa: 19, 17 Native (90%), 2 Introduced (10%). WIS: UPL-0, FACU-1, FAC-4,FACW-10, OBL-4.Shrub Layer: Frangula alnus, Salix nigra. Herb Layer: Agalinus paupercula, Apiosamericana, Bidens frondosa, Cyperus strigosus, Eupatorium perfoliatum, Euphorbiamaculata, Euthamia graminifolia, Gratiola aurea, Hypericum majus, Lindernia dubiavar. anagallidea, Lycopus americanus, Mollugo verticillata, Panicum rigidulum,Persicaria maculosa, Portulaca oleracea, Rubus hispidus, Viola lanceolata.Study site 2: Fells Reservoir, Basin 2, Stoneham, 42u26950.90N, 71u591.20W,79 m; a muddy pond shore on the northwest side of the reservoir.Life forms: Trees-0, Shrubs-0, Vines-0, Forbs-3, Graminoids-4, Ferns-0. Totaltaxa: 7, 7 Native (100%), 0 Introduced (0%). WIS: UPL-0, FACU-0, FAC-0,FACW-1, OBL-6.Herb Layer: Eleocharis acicularis, E. obtusa, E. palustris, Eupatoriumperfoliatum, Gratiola aurea, Lindernia dubia var. anagallidea, Schoenoplectuspungens.

Red Maple Swamp (S5). This common forest occurs in undrained basins, onhillside seeps, around vernal pools, and on pond shores. See also Alluvial RedMaple Swamp for this variant.Study site: Far northern end of the pond in Wright’s Park in a wetlands mosaic,Medford, 42u269420N, 71u69W, 43 m.Life forms: Trees-7, Shrubs-8, Vines-1, Forbs-22, Graminoids-1, Ferns-3. Totaltaxa: 42, 42 Native (100%) 0 Introduced (0%). WIS: UPL-3, FACU-8, FAC-10,FACW-17, OBL-4.Tree Layer: Acer rubrum, Betula lenta, Fraxinus americana, Quercus bicolor.Shrub Layer: Corylus cornuta, Clethra alnifolia, Hamamelis virginiana, Lindera

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benzoin, Nyssa sylvatica, Parthenocissus quinquefolia, Tilia americana, Toxico-

dendron radicans, Ulmus americana, Vaccinium corymbosum, Viburnum denta-

tum var. lucidum, V. lentago. Herb Layer: Aralia nudicaulis, Arisaema

triphyllum, Chelone glabra, Cinna arundinacea, Eupatorium perfoliatum, Eurybia

divaricata, Eutrochium purpureum, Geranium maculatum, Iris versicolor,

Maianthemum canadense, M. racemosum, Nabalus altissimus, Oclemena

acuminata, Onoclea sensibilis, Osmunda regalis var. spectabilis, Osmundastrum

cinnamomeum, Platanthera lacera, P. psycodes, Polygonatum pubescens,

Ranunculus recurvatus, Rubus hispidus, R. pubescens, Scutellaria lateriflora,

Smilax herbacea, Symphyotrichum lanceolatum, Viola pallens.

Shallow Emergent Marsh (S4). Like Deep Emergent Marshes, these marshes

commonly occur along the margins of ponds, lakes, and slow moving streams.

They have standing water less than 0.15 m, deep, muck soils, and short,

herbaceous vegetation dominated by graminoids.

Study site: Straight Gully Brook, downstream of the Alluvial Red Maple

Swamp plot, Medford, 42u259510N, 71u69160W, 47 m.

Life forms: Trees-0, Shrubs-0, Vines-0, Forbs-10, Graminoids-6, Ferns-0. Total

taxa: 16, 15 Native (93%), 1 Introduced (7%). WIS: UPL-0, FACU-0, FAC-0,

FACW-1, OBL-15.

Herb Layer: Alisma subcordatum, Asclepias incarnata subsp. pulchra, Calama-

grostis canadensis, Carex crinita, C. stipata, Cicuta maculata, Dulichium

arundinaceum, Galium palustre, Leersia oryzoides, Ludwigia palustris, Lysima-

chia terrestris, Lythrum salicaria, Proserpinaca palustris, Scirpus cyperinus, Sium

suave, Sparganium americanum.

Shrub Swamp (S5). Found on pond and stream margins and in basins, these

often contain vernal pools with few and small trees. This habitat is common in

the numerous hollows between the ridges.

Study site: Large basin depression between the Skyline and Reservoir Trails

north of Gerry Hill, Stoneham, 42u2792.40N, 71u6931.40W, 63 m.

Life forms: Trees-3, Shrubs-11, Vines-1, Forbs-2, Graminoids-1, Ferns-0. Total

taxa: 18, 16 Native 89 (%), 2 Introduced (11%). WIS: UPL-0, FACU-0, FAC-6,

FACW-9, OBL-3.

Tree Layer: Acer rubrum, Salix nigra. Shrub Layer: Alnus serrulata,

Cephalanthus occidentalis, Clethra alnifolia, Frangula alnus, Ilex verticillata,

Lyonia ligustrina, Quercus bicolor, Rhododendron viscosum, Rosa palustris, Salix

discolor, Smilax rotundifolia, Spiraea alba var. latifolia, Vaccinium corymbosum.

Herb Layer: Bidens sp., Scirpus cyperinus, Solanum dulcamara.

Wet Meadow (S4). A seasonally flooded graminoid community found along a

few streams in the Fells and at the base of dams. Some are maintained by

annual mowing, while other wet meadows are unmowed, with woody plants

beginning to take over.

Study site: Small meadow on the corner of Ravine and Woodland Roads

adjacent to where Spot Pond Brook flows under Woodland Road, Stoneham,

42u27919.50N, 71u59210W, 44 m.

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Life forms: Trees-3, Shrubs-3, Vines-1, Forbs-14, Graminoids-5, Ferns-3. Total

taxa: 29, 18 Native (62%), 11 Introduced (38%). WIS: UPL-5, FACU-7, FAC-

8, FACW-6, OBL-3.

Shrub Layer: Acer rubrum, Celastrus orbiculatus, Frangula alnus, Fraxinus

americana, Rhus hirta, Rosa multiflora, Toxicodendron radicans. Herb Layer:

Barbarea vulgaris, Carex comosa, C. stipata, C. vulpinoidea, Cirsium vulgare,

Daucus carota, Dichanthelium clandestinum, Epilobium ciliatum, Equisetum

arvense, Eupatorium perfoliatum, Euthamia graminifolia, Glechoma hederacea,

Juncus effusus, Leucanthemum vulgare, Lobelia inflata, Lythrum salicaria,

Onoclea sensibilis, Rubus occidentalis, Selaginella apoda, Solidago gigantea, S.

rugosa, Veronica officinalis.

Woodland Vernal Pool (S3). These wetlands occur in depressions in upland

forests and in wetlands basins, usually drying out in the summer. There are

estimated to be well over 100 vernal pools, making it one of the most common

habitats in the Fells, and an unusual concentration in such an area for eastern

Massachusetts (Lynn Harper, Natural Heritage and Endangered Species

Program, Massachusetts Division of Fisheries and Wildlife, pers. comm.).

Study site: Southeast side of Bear Hill, Stoneham, 42u279500N, 71u69160W, 61 m.

Life forms: Trees-0, Shrubs-4, Vines-0, Forbs-7, Graminoids-0, Ferns-2. Total

taxa: 13, 13 Native (100%), 0 Introduced (0%). WIS: UPL-0, FACU-0, FAC-4,

FACW-6, OBL-3.

Shrub Layer: Ribes hirtellum, Sambucus nigra subsp. canadensis, Toxicodendron

radicans, Viburnum dentatum var. lucidum. Herb Layer: Arisaema triphyllum,

Cardamine pensylvanica, Equisetum arvense, Hydrocotyle americana, Impatiens

capensis, Osmundastrum cinnamomeum, Packera aurea, Persicaria arifolia, Viola

cucullata.

Terrestrial Communities

Black Oak–Scarlet Oak Forest/Woodland (S3/S4). A fire-maintained, succes-

sional community with an open canopy occurring on dry, well-drained, rocky

upper slopes and summits, often as part of rock bald mosaics. A common

habitat that often occurs in small stands and intergrades with other mid-slope

oak forest habitats.

Study site: South of the Cross Fells Trail east of the juncture with Middle Road,

Medford, 42u269 180N, 71u69520W, 50 m.

Life forms: Trees-7, Shrubs-3, Vines-1, Forbs-2, Graminoids-1, Ferns-1. Total

taxa: 15, 14 Native (93%), 1 Introduced (7%). WIS: UPL-4, FACU-8, FAC-3,

FACW-0, OBL-0.

Tree Layer: Betula populifolia, Populus grandidentata, Quercus coccinea. Shrub

Layer: Comptonia peregrina, Frangula alnus, Pinus strobus, Quercus velutina,

Sassafras albidum, Smilax rotundifolia, Vaccinium angustifolium, V. pallidum.

Herb Layer: Carex pensylvanica, Cypripedium acaule, Lysimachia quadrifolia,

Pteridium aquilinum subsp. latiusculum.

[Blast Rock Tailings]. These large but uncommon habitats were formed by the

dumping of highway construction rock-blast debris in forests. The habitat is

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extremely xeric and nutrient-poor, with dense stands of invasive plants, early-

successional trees, and native rock-bald plants.

Study site: Between Quarry Road and the Skyline Trail near the east/west gap in

Little Pine Hill, Medford, 42u26930N, 71u6917.80W, 67 m.

Life forms: Trees-10, Shrubs-3, Vines-3, Forbs-12, Graminoids-1, Ferns-0.

Total taxa: 29, 16 Native (55%), 13 Introduced (45%). WIS: UPL-14, FACU-

12, FAC-3, FACW-0, OBL-0.

Tree Layer: Acer platanoides, A. saccharum, Betula lenta, B. papyrifera, B.

populifolia, Populus tremuloides, Prunus serotina. Shrub Layer: Frangula alnus,

Lonicera sp., Pinus strobus, Rhus typhina, Rosa multiflora, Rubus flagellaris, R.

occidentalis, Toxicodendron radicans. Herb Layer: Artemisia vulgaris, Centaurea

stoebe subsp. micranthos, Cynanchum louiseae, Danthonia spicata, Erechtites

hieraciifolius, Fragaria virginiana, Hieracium pilosella, Hypericum perforatum,

Leucanthemum vulgare, Oenothera biennis, Potentilla simplex, Rhamnus

cathartica, Solidago puberula, Verbascum thapsus.

Circumneutral Talus Forest (S3). This community, consisting of slopes of

jumbled rocks, occurs in many scattered locations in the Fells.

Study site 1: Glade on the north side of St. Denis Road near the bottom of the

south side of Pine Hill, Medford, 42u25952.80N, 71u6921.40W, 28 m.

Life forms: Trees-7, Shrubs-2, Vines-1, Forbs-18, Graminoids-7, Ferns-0. Total

taxa: 35, 30 Native (86%), 5 Introduced (14%). WIS: UPL-19, FACU-13, FAC-

3, FACW-0, OBL-0.

Tree Layer: Carya glabra, Quercus rubra. Shrub Layer: Betula lenta, Carya

glabra, Euonymus alatus, Fraxinus americana, Juniperus virginiana, Ligustrum

vulgare, Malus baccata, Quercus alba, Q. rubra, Vitis aestivalis. Herb Layer:

Agalinis tenuifolia, Agrostis perennans, Antennaria plantaginifolia, Aralia

nudicaulis, Carex pensylvanica, Desmodium 3humifusum, D. paniculatum, D.

rotundifolium, Dichanthelium clandestinum, D. dichotomum, Elymus hystrix,

Eurybia divaricata, Fallopia convolvulus, Fragaria virginiana, Galium circaezans,

Helianthus divaricatus, Lespedeza frutescens, L. procumbens, L. violacea,

Maianthemum racemosum, Quercus alba, Schedonorus pratensis, Solidago

arguta, S. bicolor, Symphyotrichum patens, S. undulatum.

Study site 2: Southeast side of Bear Hill, Stoneham, 42u279500N, 71u69190W,

67 m.

Life forms: Trees-5, Shrubs-6, Vines-2, Forbs-14, Graminoids-0, Ferns-1. Total

taxa: 28, 19 Native (68%), 9 Introduced (32%). WIS: UPL-10, FACU-14, FAC-

4, FACW-0, OBL-0.

Tree Layer: Acer saccharum, Carya ovata, Celtis occidentalis, Fraxinus

americana. Shrub Layer: Celastrus orbiculatus, Ligustrum vulgare, Partheno-

cissus quinquefolia, Rhamnus cathartica, Rhodotypos scandens, Rosa multiflora,

Rubus allegheniensis, R. occidentalis, Toxicodendron radicans. Herb Layer:

Alliaria petiolata, Circaea canadensis, Cynanchum louiseae, Dryopteris margin-

alis, Eurybia divaricata, Geranium robertianum, Impatiens capensis, Maianthe-

mum racemosum, Micranthes virginiensis, Polygonatum pubescens, Ranunculus

fascicularis, Sanguinaria canadensis, Solanum dulcamara, Solidago caesia,

Symphyotrichum cordifolium.

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[Cultural Forest]. Evergreen plantations in the Fells are unmanaged and, withthe exception of the non-native overstory, are not very different from nativeforests. These planted trees have spread only sparsely into adjacent areas, andwill likely decline and be succeeded by native trees.

Study site: North Dam Road on the north end of the North Reservoir,Stoneham, 42u27957.50N, 71u69590W, 45 m.

Life forms: Trees-8, Shrubs-4, Vines-1, Forbs-4, Graminoids-1, Ferns-1. Totaltaxa: 19, Native 16 (84%), Introduced 3 (16%). WIS: UPL-2, FACU-11, FAC-6, FACW-0, OBL-0.

Tree Layer: Pinus resinosa, Quercus rubra. Shrub Layer: Acer rubrum, Caryaovata, Frangula alnus, Fraxinus americana, Hamamelis virginiana, Pinus strobus,Pseudotsuga menziesii, Quercus alba, Smilax rotundifolia, Toxicodendronradicans, Viburnum acerifolium. Herb Layer: Aralia nudicaulis, Carex pensyl-vanica, Chimaphila maculata, Maianthemum canadense, Monotropa uniflora,Pteridium aquilinum var. latiusculum.

Cultural Grasslands. Grassy areas occur scattered throughout the Fells. Someare fields and roadsides that were seeded with non-native grasses, while otherswere created simply by mowing the existing vegetation. These man-made areasare fragmented, disturbed edges dominated by non-native grasses and weedyforbs. The NHESP classification for this habitat focuses on sites occurring onsandplains, including airports, cemeteries, pastures, and hayfields, none ofwhich apply to the Fells.

Study site: Just north of gate 25 in a 2 by 50 m strip adjoining the sidewalk onRoute 28, Stoneham, 42u27911.50N, 71u6915.80W, 55 m.

Life forms: Trees-4, Shrubs-2, Vines-0, Forbs-45, Graminoids-14, Ferns-0.Total taxa: 65, 32 Native (49%), 33 Introduced (51%). WIS: UPL-39, FACU-19, FAC-3, FACW-4, OBL-0.

Tree Layer: Ailanthus altissima, Quercus rubra. Herb Layer: Achillea millefolia,Agrostis stolonifera, Alliaria petiolata, Amaranthus powellii, Ambrosia artemi-siifolia, Antennaria plantaginifolia, Artemisia vulgaris, Asclepias syriaca,Barbarea vulgaris, Bulbostylis capillaris, Capnoides sempervirens, Carya glabra,Cerastium fontanum subsp. vulgare, Euphorbia maculata, Chelidonium major,Chenopodium album, C. simplex, Chicorium intybus, Erigeron canadensis,Cyperus esculentus, Daucus carota, Digitaria sanguinalis, Echinochloa muricata,Elymus repens, Eragrostis capillaris, E. spectabilis, Erechtites hieraciifolius,Eurybia divaricata, Euthamia graminifolia, Fraxinus pensylvanica, Geraniumcarolinianum, Hieracium sabaudum, Lepidium campestre, L. virginicum, Leu-canthemum vulgare, Linaria vulgaris, Melilotus alba, Mollugo verticillata,Nuttallanthus canadensis, Oenothera biennis, Oxalis stricta, Panicum capillare,P. dichotomiflorum, Persicaria maculosa, Phytolacca americana, Plantagolanceolosa, P. major, Poa pratensis, Polygonum aviculare, Pseudognaphaliumobtusifolium, Rhus typhina, Rosa multiflora, Rumex acetosella, Scorzoneroidesautumnalis, Setaria faberi, S. pumila, S. viridis, Silene antirrhina, Taraxacumofficinalis, Trifolium arvense, T. hybridum, Verbascum thapsus, Vicia cracca.

Dry Rich Acidic Oak Forest (S4). A mixed deciduous forest on south-facingslopes with an open understory and diverse herbaceous layer. This habitat onlyoccurs in the Medford Dike.

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Study site: Pine Hill, Medford, 42u26940N, 71u69220W, 52 m.

Life forms: Trees-7, Shrubs-1, Vines-0, Forbs-12, Graminoids-3, Ferns-0. Total

taxa: 23, 22 Native (96%), 1 Introduced (4%). WIS: UPL-11, FACU-11, FAC-

1, FACW-0, OBL-0.

Tree Layer: Acer saccharum, Betula lenta, Fraxinus americana, Ostrya virginiana,

Quercus rubra. Shrub Layer: Carya ovata, Tilia americana, Viburnum acerifolium.

Herb Layer: Agrostis perennans, Alliaria petiolata, Aralia nudicaulis, Carex

pensylvanica, C. rosea, Chimaphila maculata, Eurybia divaricata, Maianthemum

canadense, M. racemosum, Paronychia canadensis, Polygonatum pubescens,

Solidago arguta, S. caesia, Symphyotrichum cordifolium, S. undulatum.

Forest Seep Community (S4). A moist habitat of slopes where there are springs

and seeps, occurring in upland forests, with a canopy derived from and similar

to the surrounding forest. It is characterized by an abundance of ferns and

wetland taxa. The NHESP classification describes this community as occurring

on slopes within Northern Hardwood Forests.

Study site: Bear Hill, Stoneham, 42u279540N, 71u69220W, 88 m.

Life forms: Trees-4, Shrubs-7, Vines-2, Forbs-10, Graminoids-0, Ferns-3. Total

taxa: 26, 18 Native (69%), 8 Introduced (31%). WIS: UPL-6, FACU-12, FAC-

6, FACW-2, OBL-0.

Tree Layer: Fraxinus americana, Ostrya virginiana, Quercus rubra. Shrub Layer:

Acer saccharum, Berberis thunbergii, Celastrus orbiculatus, Euonymus alatus,

Frangula alnus, Hamamelis virginiana, Rhamnus cathartica, Rosa multiflora,

Toxicodendron radicans, Vitis labrusca. Herb Layer: Amphicarpaea bracteata,

Arisaema triphyllum, Athyrium felix-femina var. angustum, Epipactis helleborine,

Eurybia divaricata, Geranium maculatum, Nabalus altissimus, Osmundastrum

cinnamomeum, Polygonatum pubescens, Polystichum acrostichoides, Solanum

dulcamara, Solidago caesia, Uvularia sessilifolia.

Hemlock–White Pine–Oak Forest (S5). These sparse, low-diversity forest

stands occur on well-drained slopes of low fertility, on north and east-facing

slopes in the Fells. With a shift in composition toward the Oaks, this would be

called Oak-White Pine-Hemlock Forest.

Study site: Molly9s Spring Road, Winchester, 42u26.540N, 71u79240W, 76–82 m.

Life forms: Trees-8, Shrubs-2, Vines-0, Forbs-4, Graminoids-1, Ferns-0. Total

taxa: 15, 15 Native (100%), 0 Introduced (0%). WIS: UPL-2, FACU-10, FAC-

3, FACW-0, OBL-0.

Tree Layer: Acer rubrum, A. saccharum, Betula lenta, Carya ovata, Pinus

strobus, Quercus coccinea, Q. rubra, Tsuga canadensis. Shrub Layer: Hamamelis

virginiana, Kalmia latifolia. Herb Layer: Aralia nudicaulis, Carex pensylvanica,

Maianthemum canadense, Mitchella repens, Monotropa uniflora.

Hickory–Hop Hornbeam Forest/Woodland (S2). An open canopy community

with few shrubs and a continuous herb layer of graminoids. The most extensive

stands have occurred historically on the upper east and south sides of Bear Hill.

Other stands occur on the east side of the hill south of Bear Hill.

Study site: Bear Hill, Stoneham, 42u279430N, 71u69150W, 70 m.

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Life forms: Trees-2, Shrubs-0, Vines-0, Forbs-2, Graminoids-1, Ferns-0. Totaltaxa: 5, 4 Native (80%), 1 Introduced (20%). WIS: UPL-3, FACU-2, FAC-0FACW-0, OBL-0.

Tree Layer: Carya glabra, Ostrya virginiana. Herb Layer: Carex pensylvanica,Galium mollugo, Polygonatum pubescens.

Mixed Oak Forest (S5). These dry forests can have all four upland oaks, Black,Red, Scarlet, and White. Understories are open, with Lowbush Blueberry andBlack Huckleberry shrubs and Bracken Fern. These habitats commonlyintergrade with stands of White Pine-Oak and Oak-Hickory.Study site: Wright’s Park, Medford, 42u269270N, 71u5950W, 52 m.Life forms: Trees-10, Shrubs-4, Vines-0, Forbs-2, Graminoids-0, Ferns-0. Totaltaxa: 16, 15 Native (94%), 1 Introduced (6%). WIS: UPL-5, FACU-7, FAC-4,FACW-0, OBL-0.Tree Layer: Quercus alba, Q. coccinea, Q. rubra, Q. velutina. Shrub Layer: Acerrubrum, Amelanchier sp., Betula lenta, Carya glabra, Frangula alnus, Gaylussaciabaccata, Prunus serotina, Sassafras albidum, Vaccinium pallidum, Viburnumacerifolium. Herb Layer: Chimaphila maculata, Maianthemum canadense.

Northern Hardwoods–Hemlock–White Pine Forest (S5). This forest occursrarely on north- and east-facing slopes, with deciduous trees mixed withevergreens. Stands such as in the Fells are considered historically rare in theBoston Basin Ecoregion.Study site: North of gate 39, Virginia Wood Trail, Stoneham, 42u279120N,71u79150W, 87 m.Life forms: Trees-7, Shrubs-2, Vines-2, Forbs-13, Graminoids-1, Ferns-2. Totaltaxa: 27, 24 Native (89%), 3 Introduced (11%). WIS: UPL-6, FACU-12, FAC-7, FACW-2, OBL-0.

Tree Layer: Acer saccharum, Betula alleghaniensis, Fagus grandifolia, Fraxinusamericana, Prunus serotina, Tsuga canadensis. Shrub Layer: Frangula alnus,Parthenocissus quinquefolia, Toxicodendron radicans, Viburnum acerifolium,Vitis aestivalis. Herb Layer: Aralia nudicaulis, A. racemosa, Chelidonium majus,Dennstaedtia punctilobula, Dryopteris carthusiana, Epifagus virginiana, Eurybiadivaricata, Geum canadense, Impatiens capensis, Leersia virginica, Maianthemumcanadense, M. racemosum, Persicaria maculosa, P. virginiana, Phytolaccaamericana, Rubus allegheniensis.

Oak–Hickory Forest (S4). This forest occurs on upper slopes, grading into allthe oak forest habitats.Study site 1: Wamoset Hill, Crystal Spring Road, Stoneham, 42u279300N,71u5930W, 29 m.Life forms: Trees-11, Shrubs-5, Vines-1, Forbs-10, Graminoids-1, Ferns-0.Total taxa: 28, 20 Native (71%), 8 Introduced (29%). WIS: UPL-7, FACU-12,FAC-7, FACW-2, OBL-0.

Tree Layer: Acer platanoides, A. rubrum, Carya ovata, Ostrya virginiana,Quercus rubra. Shrub Layer: Amelanchier sp., Berberis vulgaris, Euonymus alata,Frangula alnus, Fraxinus americana, Malus sp., Prunus serotina, Rhamnuscathartica, Smilax rotundifolia, Tilia americana, Toxicodendron radicans,Viburnum acerifolium. Herb Layer: Alliaria petiolata, Arisaema triphyllum,

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Carex sp., Circaea canadensis, Eurybia divaricata, Geum canadense, Impatiens

capensis, Maianthemum racemosum, Solanum dulcamara, Solidago caesia, Viola

sororia.

Study site 2: Route 28/Route 93 highway island, Medford, 42u269450N,

71u69140W, 57 m.

Life forms: Trees-5, Shrubs-2, Vines-0, Forbs-7, Graminoids-1, Ferns-1. Total

taxa: 16, 16 Native (100%), 0 Introduced (0%). WIS: UPL-3, FACU-11, FAC-

2, FACW-0, OBL-0.

Tree Layer: Acer rubrum, A. saccharum, Carya glabra, Quercus velutina. Shrub

Layer: Prunus serotina, Vaccinium angustifolium, Viburnum acerifolium. Herb

Layer: Aralia nudicaulis, Carex pensylvanica, Maianthemum canadense, M.

racemosum, Monotropa uniflora, Polygonatum pubescens, Pteridium aquilinum

subsp. latiusculum, Solidago caesia, Uvularia sessilifolia.

Open Oak Forest/Woodland (S3). A successional oak community maintained

by frequent anthropogenic fires, with low-statured trees forming a very open

canopy on gradual slopes. The canopy composition varies from the model

depending on the matrix forest the community occurs in, this one being in an

Oak-Hickory Forest. The shrub layer consists of tree regrowth and shrubs

typical of rock outcrops, which often occur within the community. The herb

layer is continuous with forbs, graminoids, ferns, and mosses. This community

was newly recognized by NHESP in 2011.

Study site: Jerry Jingle Road, Melrose, 42u269260N, 71u049560W, 70 m.

Life forms: Trees-5, Shrubs-5, Vines-0, Forbs-4, Graminoids-3, Ferns-1. Total

taxa: 18, 18 Native (100%), 0 Introduced (0%). WIS: UPL-9, FACU-9, FAC-0,

FACW-0, OBL-0.

Tree Layer: Quercus alba, Q. rubra, Q. velutina. Shrub Layer: Carya ovata,

Comptonia peregrina, Rhus typhina, Sassafras albidum, Vaccinium angustifolium,

V. pallidum, Viburnum acerifolium. Herb Layer: Danthonia spicata, Deschampsia

flexuosa, Lespedeza capitata, Pteridium aquilinum subsp. latiusculum, Rubus

flagellaris, Schizachyrium scoparium, Solidago bicolor, S. puberula.

Pitch Pine–Oak Forest/Woodland (S5). A fire-dependent, open-to-closed

canopy community on dry, acidic, low nutrient soils, occurring adjacent to

Pitch Pine/Scrub Oak Communities in the Fells. The shrub layer is dense to

patchy with ericaceous shrubs, scrub oak, and catbrier, and the herb layer is

sparse.

Study site: MIT Observatory Hill, Malden, 42u269270N, 71u059040W, 82 m.

Life forms: Trees-8, Shrubs-6, Vines-1, Forbs-2, Graminoids-1, Ferns-1. Total

taxa: 19, 18 Native (95%), 1 Introduced (5%). WIS: UPL-6, FACU-9, FAC-4,

FACW-0, OBL-0.

Tree Layer: Acer rubrum, Betula populifolia, Pinus rigida, P. strobus, Populus

grandidentata, Quercus alba, Q. coccinea, Q. velutina. Shrub Layer: Comptonia

peregrina, Frangula alnus, Gaylussacia baccata, Quercus ilicifolia, Smilax

rotundifolia, Vaccinium angustifolium, V. pallidum. Herb Layer: Carex

pensylvanica, Cypripedium acaule, Gaultheria procumbens, Pteridium aquilinum

subsp. latiusculum.

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Red Oak–Sugar Maple Transition Forest (S4). Mesic, mid-slope stands with amixed deciduous and evergreen tree layer. This habitat is becoming morecommon, due to Hemlock die off and mesophication.Study site: Bear Hill, Stoneham, 42u279580N, 71u069220W, 70 m.

Life forms: Trees-7, Shrubs-2, Vines-0, Forbs-5, Graminoids-1, Ferns-0. Totaltaxa: 15, 14 Native (93%), 1 Introduced (7%). WIS: UPL-3, FACU-9, FAC-3,FACW-0, OBL-0.Tree Layer: Acer saccharum, Betula lenta, Pinus strobus, Quercus rubra, Tsugacanadensis. Shrub Layer: Acer saccharum, Frangula alnus, Prunus serotina,Quercus rubra, Toxicodendron radicans, Viburnum acerifolium. Herb Layer:Aralia nudicaulis, Carex pensylvanica, Eurybia divaricata, Maianthemumcanadense, Mitchella repens, Polygonatum pubescens.

Ridgetop Pitch Pine/Scrub Oak Community (S2). Pitch Pine has increasedsignificantly in the past century, now occurring in over two dozen placesthroughout the Fells. It occurs mainly on plutonic granitic, or volcanicrhyolitic, hilltop domes, often as part of rock-bald mosaics.

Study site: North end of Whip Hill, Stoneham, 42u279490N, 71u5911.60W, 76 m.

Life forms: Trees-5, Shrubs-5, Vines-1, Forbs-0, Graminoids-1, Ferns-0. Totaltaxa: 12, 12 Native (100%), 0 Introduced (0%). WIS: UPL-5, FACU-5, FAC-2,FACW-0, OBL-0.

Tree Layer: Pinus rigida, Quercus alba, Q. coccinea, Q. velutina. Shrub Layer:Aronia melanocarpa, Gaylussacia baccata, Juniperus communis var. depressa,Pinus strobus, Q. ilicifolia, Vaccinium angustifolium, V. pallidum. Herb Layer:Carex pensylvanica.

Rocky Summit and Rock Outcrop Communities (S3-S4). A unique feature ofthe Fells is its many, large, and widely distributed rock balds. Sun-heated,exposed rocks and thin soils create dry conditions susceptible to fire. The baldsare a mosaic of microhabitats, occurring in seral stages from bare, exposed rockcolonized by lichens and mosses, to graminoid patches, ericad and Scrub Oakshrubberies, Ridgetop Pitch Pine Forest stands, stunted woodland patches ofBlack Oak-Scarlet Oak Forest on acidic rock, and Red Cedar in circumneutralareas. Some plants, such as Rock Harlequin, are restricted to rock balds, andPleatleaf Knotweed and Rock Spikemoss, to circumneutral rocks.

Study site: Entire open summit of Nanepashemet Hill, Winchester, 42u279120N,71u79150W, 87 m.

Life forms: Trees-9, Shrubs-5, Vines-2, Forbs-12, Graminoids-10, Ferns-1.Total taxa: 39, 37 Native (95%), 2 Introduced (5%). WIS: UPL-17, FACU-14,FAC-8, FACW-0, OBL-0.

Tree Layer: Quercus alba, Q. coccinea, Q. velutina. Shrub Layer: Amelanchierspicata, Aralia hispida, Aronia melanocarpa, Betula populifolia, Carya glabra,Comptonia peregrina, Frangula alnus, Populus grandidentata, Prunus pensylvan-ica, P. serotina, Quercus ilicifolia, Smilax rotundifolia, Spiraea alba var. latifolia.Herb Layer: Apocynum androsaemifolium, Arctostaphylos uva-ursi, Aristidadichotoma, Bulbostylis capillaris, Capnoides sempervirens, Carex pensylvanica, C.umbellata, Comandra umbellata, Danthonia spicata, Dichanthelium acuminatumsubsp. fasciculatum, Digitaria sanguinalis, Erechtites hieraciifolius, Helianthe-mum canadense, Hypericum gentianoides, Juncus greenei, J. tenuis, Lechea

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tenuifolia, Pteridium aquilinum subsp. latiusculum, Rubus allegheniensis, R.flagellaris, Schizachyrium scoparium, Solidago bicolor, S. puberula.

Scrub Oak Shrubland (S1). This rare community occurs on ridge tops, anddepends on fire to keep trees from overgrowing the dominant Scrub Oak. Earlysuccessional species such as Black and Gray Birch often occur along with Blackand Scarlet Oaks, all of which are multi-stemmed from repeated fire events. Ifthe trees exceed 25% of the cover then the community would be classified asBlack Oak-Scarlet Oak Woodland. Ericad shrubs of Lowbush Blueberry andHuckleberry can be codominant when the Scrub Oak is not dense. There is anherbaceous layer of graminoids, mosses, and lichens.

Study site: North side of Gerry Hill, Medford, 42u26959.80N, 71u69290W, 73 m.

Life forms: Trees-6, Shrubs-4, Vines-0, Forbs-5, Graminoids-2, Ferns-1. Totaltaxa: 18, 17 Native (93%), 1 Introduced (7%). WIS: UPL-7, FACU-9, FAC-2,FACW-0, OBL-0.

Tree Layer: Quercus alba. Shrub Layer: Aronia melanocarpa, Betula populifolia,Comptonia peregrina, Pinus nigra, P. strobus, Prunus pensylvanica, Quercusilicifolia, Q. velutina, Vaccinium angustifolium. Herb Layer: Carex pensylvanica,Comandra umbellata, Danthonia spicata, Gaultheria procumbens, Lecheatenuifolia, Polygala sanguinea, Pteridium aquilinum subsp. latiusculum, Solidagopuberula.

[Successional Birch–Poplar Woodland]. This is the most common successionalcommunity, as it is a seral stage for all the different oak forests. Gray Birch andBigtooth Poplar are the characteristic and dominant trees, germinating on baresoils after fires, and resprouting from tree bases and roots with recurring fires.As Hemlock stands die off from Wooly Adelgid infestations, Black Birchsaplings often dominate in pure stands, while in other areas there is a mix ofdeciduous hardwoods. Herbaceous plants benefit from the increased sunlightand occur in large patches. This common successional habitat is not included inthe NHESP classification.

Study site 1: Near gate 40, Ravine Road, Stoneham, 42u27990N, 71u5960W, 37 m.

Life forms: Trees-9, Shrubs-1, Vines-3, Forbs-6, Graminoids-0, Ferns-1. Totaltaxa: 20, 14 Native (70%), 6 Introduced (30%). WIS: UPL-5, FACU-12, FAC-3, FACW-0, OBL-0.

Tree Layer: Tsuga canadensis. Shrub Layer: Acer saccharum, Ailanthusaltissima, Betula lenta, Celastrus orbiculatus, Frangula alnus, Fraxinus america-na, Prunus avium, P. serotina, Quercus rubra, Rubus allegheniensis, Vitisaestivalis. Herb Layer: Aralia hispida, Dryopteris intermedia, Fallopia japonica,Maianthemum canadense, Mitchella repens, Phytolacca americana, Solanumdulcamara, Symphyotrichum cordifolium.Study site 2: Quarter Mile Pond, in Sector 5, Medford, 42u269450N, 71u59480W,58 m.

Life forms: Trees-10, Shrubs-6, Vines-1, Forbs-9, Graminoids-1, Ferns-1. Totaltaxa: 28, 27 Native (96%), 1 Introduced (4%). WIS: UPL-10, FACU-11, FAC-7, FACW-0, OBL-0.

Tree Layer: Pinus rigida. Shrub Layer: Acer rubrum, Amelanchier sp., Betulalenta, B. populifolia, Gaylussacia baccata, Frangula alnus, Populus grandidentata,Prunus serotina, Quercus alba, Q. ilicifolia, Rhus typhina, Sassafras albidum,

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Smilax rotundifolia, Toxicodendron radicans, Vaccinium pallidum. Herb Layer:Aralia hispida, A. nudicaulis, Carex pensylvanica, Chimaphila maculata,Erechtites hieraciifolius, Nabalus trifoliolatus, Phytolacca americana, Pteridiumaquilinum subsp. latiusculum, Solidago nemoralis, S. puberula, S. rugosa.

Sugar Maple–Oak–Hickory (S2). This community occurs on a southeast-facing,concave, mesic, middle slope with downslope enrichment, deep soils with twofeet of black loam topsoil, and a thin surface organic layer. Several springs arisewithin the site, creating small intermittent streams that feed a vernal pool. Thecanopy composition fits the model of Rich Mesic Forest, being dominated bySugar Maple and Bitternut Hickory with some Ash, Basswood, and Elm, butlacks the typical rich herbaceous layer. As such, it has been recognized byNHESP as an eastern variant, and given a new community name in theClassification.

Study site: Southeast side of Bear Hill, Stoneham, 42u279500N, 71u69180W, 65 m.

Life forms: Trees-7, Shrubs-2, Vines-0, Forbs-12, Graminoids-2, Ferns-3. Totaltaxa: 26, 25 Native (96%), 1 Introduced (4%). WIS: UPL-7, FACU-14, FAC-2,FACW-3, OBL-0.

Tree Layer: Acer saccharum, Carya cordiformis, Fagus grandifolia, Fraxinusamericana, Ostrya virginiana, Tilia americana, Ulmus americana. Shrub Layer:Cornus alternifolia, Ribes hirtellum. Herb Layer: Alliaria petiolata, Anemoneamericana, A. quinquefolia, Aralia racemosa, Arisaema triphyllum, Athyriumfelix-femina var. angustum, Bromus pubescens, Carex rosea, Circaea canadensis,Eurybia divaricata, Geranium maculatum, Maianthemum racemosum, Osmorhizaclaytonii, Parathelypteris noveboracensis, Polystichum acrostichoides, Ranuncu-lus allegheniensis, Sanicula marilandica.

[White Pine Forest (S5)]. These uncommon forests have persisted since 1894 asclosed canopy stands. There has been no selective logging to maintain thesestands, nor are they post-agricultural and, as such, they do not fit SuccessionalWhite Pine Forest as defined by the NHESP classification. The plot is in a mid-aged stand, likely regenerating from the Hurricane of 1938.

Study site: Pond Street, Stoneham, 42u279270N, 71u059100W, 40 m.

Life forms: Trees-5, Shrubs-3, Vines-1, Forbs-4, Graminoids-0, Ferns-1. Totaltaxa: 14, 13 Native (92%), 1 Introduced (8%). WIS: UPL-1, FACU-8, FAC-5,FACW-0, OBL-0.

Tree Layer: Fagus grandifolia, Pinus strobus. Shrub Layer: Carya glabra,Frangula alnus, Prunus serotina, Smilax rotundifolia, Toxicodendron radicans,Vaccinium angustifolium, Viburnum acerifolium. Herb Layer: Aralia nudicaulis,Athyrium felix-femina var. angustum, Lysimachia quadrifolia, Maianthemumcanadense, Mitchella repens.

White Pine–Oak Forest (S5). The most common forest community of the Fells,with variable proportions of White Pine and Oak. There is a variant resultingfrom the continued expansion of Beech, which is favored by fire suppression,into White Pine-Oak Forests from the small pockets that existed in 1896. Beechforms dense thickets from root sprouts that exclude most other plants and,thus, has lower diversity than most of the forest stands, mostly with only theparasitic Beechdrops as an associate.

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Study site 1: West side of the Cross Fells Trail west of its juncture with MiddleRoad, Medford, 42u269170N, 71u69590W, 43 m.Life forms: Trees-8, Shrubs-2, Vines-0, Forbs-7, Graminoids-1, Ferns-0. Totaltaxa: 18, 17 Native (95%), 1 Introduced (5%). WIS: UPL-6, FACU-11-, FAC-1,FACW-0, OBL-0.Tree Layer: Pinus strobus, Quercus coccinea, Q. velutina. Shrub Layer: Betulalenta, Carya ovata, Prunus serotina, Quercus alba, Sorbus aucuparia, Vacciniumangustifolium, V. pallidum. Herb Layer: Aralia nudicaulis, Carex pensylvanica,Chimaphila maculata, Cypripedium acaule, Lysimachia quadrifolia, Maianthe-mum canadense, M. racemosum, Rubus allegheniensis.Study site 2: East side of Hemlock Pool Path, ca. 100 m south of the RockCircuit Trail, Medford, 42u269330N, 71u59190W, 56 m.Life forms: Trees-8, Shrubs-4, Vines-0, Forbs-13, Graminoids-1, Ferns-1. Totaltaxa: 27, 27 Native (100%), 0 Introduced (0%). WIS: UPL-8, FACU-17, FAC-2, FACW-0, OBL-0.Tree Layer: Betula alleghaniensis, Fagus grandifolia, Ostrya virginiana, Pinusstrobus, Quercus alba, Q. rubra. Shrub Layer: Carya ovata, Corylus cornuta,Hamamelis virginiana, Ostrya virginiana, Sassafras albidum, Vaccinium angusti-folium, Viburnum acerifolium. Herb Layer: Aralia nudicaulis, Carex pensylvan-ica, Chimaphila maculata, C. umbellata var. cisatlantica, Conopholis americana,Epifagus virginiana, Eurybia divaricata, E. macrophylla, Geranium maculatum,Lysimachia quadrifolia, Maianthemum racemosum, Monotropa uniflora, Para-thelypteris noveboracensis, Pedicularis canadensis, Solidago caesia.

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