United States

Department of

Agriculture

Forest Service

Northeastern

Research Station

General Technical

Report NE-320

Vegetation of Forested Uplands in

the Massabesic Experimental Forest

Alison C. Dibble

Catherine A. Rees

Paul E. Sendak

John C. Brissette
Published by: For additional copies:

USDA FOREST SERVICE USDA Forest Service

11 CAMPUS BLVD SUITE 200 Publications Distribution

NEWTOWN SQUARE PA 19073-3294 359 Main Road

Delaware, OH 43015-8640

August 2004 Fax: (740)368-0152

Visit our homepage at: http://www.fs.fed.us/ne

ALISON C. DIBBLE, research ecologist, USDA Forest Service 686 Government

Road, Bradley, Maine 04411

CATHERINE A. REES, ecologist, USDA Forest Service, 686 Government Road,

Bradley, Maine 04411

PAUL E. SENDAK, research forester, USDA Forest Service, PO Box 640, Durham,

New Hampshire 03824

JOHN C. BRISSETTE, research forester, USDA Forest Service PO Box 640,

Durham, New Hampshire 03824

Abstract

A summary of an inventory of vascular plants in the 3,700-acre Massabesic

Experimental Forest in York County, Maine. On a grid of 399 permanent sample

points, trees 4.5 inches diameter breast height or greater were tallied using variable

radius plot sampling. Sample points also served as centers of 0.01-acre plots on

which smaller trees and all other plants were recorded. We identified about 500

species and subspecies. The most common overstory trees were eastern white

pine, eastern hemlock, northern red oak, and red maple. Hemlock was the most

abundant tree seedling. Shrub density was greatest for beaked hazlenut, followed

by several viburnum species, highbush blueberry, and winterberry. The most

common herb was starflower. We found nine listed rare plants, including a federally

threatened orchid, small whorled pogonia. A geographic information system was

developed and includes inventory data, forest type, soils, fire history, topography,

roads, and streams.

Manuscript received for publication 5 December 2003
1

Introduction

The plant species composition found in a

forest is the result of a complex interaction

among climate, site characteristics, and

natural and human-caused disturbances.

This report describes the history and site

conditions of various parcels of land that

make up the 3,700-acre Massabesic

Experimental Forest (MEF) in York

County, Maine. We detail the systematic

sampling procedures used to describe the

vegetation of the forest, summarize tree

composition and distribution, shrub

density, and understory plant distribution,

and list the vascular plants found in the

various parcels.

The MEF is a unique resource for forest

ecology and management research. The

Northeastern Research Station of the

United States Department of Agriculture

Forest Service currently manages eight

experimental forests. Four are on national

forests and three are on lands owned by a

state, forest industry, or a university. The

Station owns the MEF. The property

consists of northern and southern units of

nearly equal area and an administrative site located near

the southern unit (Fig. 1). The MEF has a number of

special ecological features, including some of the largest

wetlands in northern New England that are dominated

by Atlantic white-cedar (Chamaecyparis thyoides; full

scientific names are in Appendix I). It also has a high

density of seasonal wetlands or vernal pools, and

numerous species of plants and animals that are rare or

uncommon. The history of the MEF (detailed by parcel

below) includes clearing for farms that were

subsequently abandoned to be replaced with a

successional forest. A fire that burned 150,000 acres in

southwestern Maine in 1947 destroyed 80 percent of the

timber in the forest (McConkey and Smith 1958). The

forest has grown back and is the largest single block of

federal land in southern Maine. The MEF is valued for

open space and wildlife habitat in a rapidly developing

part of the state, is popular among local residents for

some types of recreation, and is an ideal location for

demonstrating good forest management.

The forests of which the MEF is a part have been

classified in a number of ways. Braun (1950) included all

of Maine, New Hampshire, Vermont, and much of

Massachusetts as the New England Section of the

Northern Appalachian Highland Division, which in turn

is part of the hemlock—white pine—northern

hardwoods region. The predominant forest-cover type is

white pine—northern red oak—red maple and there

may be white pine—hemlock in places (Baldwin and

Ward 1980, Powell 1980). Keys and others (1995)

classify that part of Maine and adjacent New Hampshire

as the Gulf of Maine Coastal Plain subsection (221Ai) of

the Southern New England Coastal Hills and Plain

Section of the Eastern Broadleaf Forest province. The

southern unit of the MEF is near the boundary of the

Gulf of Maine Coastal Lowland subsection (221Ak) and

has some elements of that subsection, especially the

Atlantic white-cedar swamps. Eastern hemlock (Tsuga

canadensis), eastern white pine (Pinus strobus), and

northern red oak (Quercus rubra) are important species

of the dominant potential vegetation in both

subsections.

Most of the northern unit burned in the 1947 forest fire.

The area regenerated to a forest dominated today by

white pine and red oak with dense patches of sapling-size

hemlock. Plantations of white pine and mixed plantings

of white and red pine (P. resinosa) were established after

the fire by the Forest Service on some of the most

severely burned areas and on areas that were poorly

stocked before the fire. There are extensive areas of open



	


	



	


	






		

		



 

Figure 1.—Location of the Massabesic Experimental Forest,

York County, Maine.
2

water in the unit, including Tarwater Pond, which is

surrounded on three sides by mixed woods, pine

plantations, a tamarack (Larix laricina) swamp, and a

shrub heath. Except for a portion on the east side, this

11-acre pond is within the boundary of the MEF. It

drains into Roberts Pond, an impoundment that

provides extensive shoreline and shallow water, and is

part of the Saco River drainage. Cooks Brook crosses

the northern unit. Numerous small seasonal wetlands

are scattered throughout the unit.

The southern unit contains some upland areas that did

not burn in 1947 and that appear to be relatively free

of disturbance during the past 100 years. Some burned

areas on this unit were planted to white and red pine,

while the remainder regenerated to hardwoods or

mixed stands of hardwoods and conifers. Areas of

sandy loam support mature stands of white pine and

hemlock, or of white and red pine (probably planted).

Pitch pine (P. rigida) naturally occurred on an area of

loamy sand and a few small populations remain. Some

of this area was planted with white and red pine, and

in the 1960s genetic test plantings of hybrid poplars

(Populus spp.) were established there. There are

complex drainage patterns across a large portion of the

southern unit that includes the Atlantic white-cedar

swamps. One observer reported that these swamps did

not burn extensively in 1947.1

 The unit contains

numerous small ponds and seasonal wetlands, and

borders Estes Lake, an impoundment of the Mousam

River.

The current composition and structure of plant

communities found in the MEF are the result of a

complex interaction among climate, topography, soils,

natural disturbances, and human land use, including

research activities since the 1940s.

Climate

Both units of the MEF are 15 to 20 miles northwest of

the Atlantic Ocean and the climate of the region is

greatly influenced by the ocean’s moderating effect. The

forest is in Plant Hardiness Zone 5a (cold hardy, -15 to

-20 °F) (Cathey 1990), while a few miles farther inland

is Zone 4b (hardy, -20 to -25 °F). The closest weather

station to the MEF with long-term data published by the

National Oceanic and Atmospheric Administration

(NOAA) is at the airport in Sanford (43°26 ́ N, 70°47 ́

W; elevation 280 ft), about 3 miles southwest of the

southern unit. NOAA uses a 30-year period, updated

each decade, to establish “normal” conditions. The

period 1961-90 defined normal or average conditions

when the inventory was conducted. The average annual

temperature for the period was 46.6 °F, during which

July was the warmest month (70.2 °F) and January the

coldest (21.5 °F) (Fig. 2). The average high temperature

was 94.7 °F and the average low was -17.1 °F. The

extremes were 101 °F on August 2, 1975 and -27 °F on

January 11, 1968. Annual precipitation for the period

averaged 46.85 inches. September was the driest month

(3.21 inches) and November the wettest (5.22 inches)

(Fig. 3). The growing season, or period between killing

1

Swett, Charles E. 1999. Personal communication. Retired,

USDA Forest Service.

Figure 2.—Monthly mean temperatures at the Sanford,

Maine airport for the 30-year period 1961-1990. The

horizontal line is the mean annual temperature (46.6 °F)

for the period. For each month, center line is normal

temperature (overall mean), upper and lower lines are

maximum and minimum means over the 30-year period.

Figure 3.—Monthly mean total precipitation at the Sanford,

Maine airport for the 30-year period 1961-1990. Vertical lines

are the 95 percent confidence limits for the period.
3

frosts (i.e., ≤ 28 °F), averaged 157 days. May 4 was the

average date of the last killing frost in the spring, and

October 8 was the first in the fall (Fig. 4).

Topography and Soils

The terrain in the MEF is flat to gently rolling with

elevation between 200-450 ft above sea level.

Noteworthy topographic features that contribute to

the diversity of habitats include sandy flats, rocky

hills, and ledge outcrops interspersed with numerous

areas of peat and muck. There are small horsebacks,

kames, possible kettleholes, and pocket wetlands that

were sculpted by the receding Wisconsin glacier about

16,000 years ago. Underlying bedrock is granite

composed especially of muscovite-biotite, though

feldspar and quartz could be more abundant.2

 Major

soil taxa in the Gulf of Maine Coastal Plain

subsection are Dystrochrepts, Udothents, and

Udipsamments (Keys et al. 1995). Soils of the MEF

are primarily stony to very stony sandy loams, ranging

to sandy soils on outwash plains. The relatively few

glacial erratics and rocky outcrops are granite.

Land Use History

Presettlement use of the area that is now the MEF is

largely unknown. It can be assumed that the Massabesic

Indians, who lived in the area when the first European

colonists arrived, probably traveled through, hunted, and

camped within the forest, especially along rivers and

streams. It is likely that these Native Americans used fire

to enhance hunting habitat in or near the area, as was

common throughout much of coastal New England into

the 1600s (Cronon 1983). Because of good harbors on

the Atlantic coast and relatively good, light textured

soils, European settlers were attracted to York County

early in the 1600s. The area was heavily forested with

both softwoods and hardwoods and lumbering was an

important industry in the region by the 1630s (Carroll

1973). Colonial and early American farms once

occupied what is now the MEF, as evidenced by old

foundations, family cemeteries, quarries, and stone

fences that are found throughout the forest. These farms

created a patchwork of fields and woodlots. After the

Civil War, farm abandonment began and continued

through the Great Depression. White pine is well

adapted to conditions following clearing and

abandonment and became the dominant tree species in

parts of the MEF, as it did elsewhere in the region.

About two-thirds of the area that is now the MEF was

purchased in the late 1800s by Benjamin Clark Jordan, a

local lumberman. He willed this property to Bates

College of Lewiston, ME in 1912 for the purpose of

establishing a forestry program. Bates College

discontinued its forestry program due to financial

hardship during the Depression. This change, combined

with the tax burden, led Bates College to sell the land in

the late 1930s (Coolidge 1963).

Establishment of the MEF

The parcels that make up the MEF were bought by the

Northeastern Forest Experiment Station (now

Northeastern Research Station) under authority of the

Weeks Act between 1937 and 1942 (McConkey and

Smith 1958). A number of old farms were purchased in

addition to the land bought from Bates College. The

land was acquired to conduct timber management

research, focusing on white pine. Because this species

was the dominant tree across previously farmed

landscapes in central and southern New England,

important management questions could be addressed in

the MEF. However, soon after the purchases were

completed, the new experimental forest was closed

during World War II. In 1946 the MEF reopened with

design of the first experiments and initiation of

treatments. As land was acquired for the MEF, a grid of

permanent plots was established across the forest.

Unfortunately, between the forest fire in 1947 and

storms in the early 1950s, most of these plot locations

were lost. 2

Leak, William B. 1999. Personal communication. Research

Forester, USDA Forest Service, Durham, NH.

Figure 4.—Dates of last spring and first fall killing frosts

(≤ 28 °F) at the Sanford, Maine airport for the years 1961-

1990. Horizontal lines are the average dates, May 4 and

October 8, respectively. The interval between is considered

the growing season and averaged 157 days for the 30-year

period. Note that the datum for the last killing frost in the

spring of 1987 is missing.
4

1947 Fire and its Aftermath

Following a prolonged drought, wildfires erupted across

Maine in October 1947 before leaf fall had completed in

southern and coastal sections. Statewide, more than

200,000 acres burned, resulting in the death of 16

people and the loss of entire villages. About 3,025 acres

of the MEF burned. In some places the fire crowned,

causing stand-replacing disturbance; in other areas only

part of the forest floor was consumed. The fire skipped

some areas entirely. Immediately following the fire, a

map of the burn was prepared with a report on quantity

and size of remaining timber (USDA Forest Service

1947). Many of the trees killed in the fire, especially in

the northern unit, were harvested the following year.

Nearly 4.5 million board feet of timber was salvaged

(McConkey and Smith 1958). The fire killed only some

of the trees outright, but it reduced stand density and

left many surviving trees weakened, leaving the stands

more vulnerable to subsequent disturbances.

Consequently, a substantial number of trees blew down

in the gale of November 1950 and during Hurricane

Carol in 1954. Another 1.75 million board feet of

mostly white pine was salvaged. After the fire and

storms, 990 acres had merchantable or potentially

merchantable stands, and 580 acres were considered

“nonproductive land” (USDA Forest Service 1955). Of

these “nonproductive” areas 330 acres were considered to

be in need of planting and much of this total was

thought to require site preparation. In the 1950s,

plantations of white and red pine were established

primarily for rehabilitation, though some were designed

with research objectives.

The Research Program, 1946 to 1996

Experiments from the early years also have influenced

the current structure and composition of the forest. In

1950, a program of management studies was started in

the unburned portion of the MEF, based on a design

similar to other silvicultural studies initiated throughout

the Forest Service around that time. The goal of these

Compartment Management studies was to compare an

array of long-term management treatments at the stand

level, and thus to evaluate quantity and quality of timber

yields, changes in growing stock conditions, and costs of

labor and materials. Treatments included diameter-limit

cutting, shelterwood, and patch and strip clearcutting

(McConkey and Smith 1958).

Other research started in the 1950s included artificial

regeneration of white and red pine by planting and

direct seeding (Graber 1965, 1968); chemical pest

control, primarily for white pine weevil (Pissodes strobi

Peck) (McConkey and Swett 1967); woodlot

management; and early stand management techniques,

such as release using herbicides and precommercial

thinning.3

 Some of the burned areas in the northern unit

regenerated to stands dominated by paper birch (Betula

papyrifera). To provide management prescriptions for

paper birch stands, an experiment was established to

study the effects of thinning, with and without fertilizer

application (Safford 1989). Except for an occasional

small salvage operation, there has been no commercial

harvesting in the MEF since the mid-1960s. Harvesting

of firewood, including standing hardwood trees, was

allowed by permit for many years, but most of that

activity was confined to a narrow band along forest

roads. Firewood harvesting was halted by the early

1980s. The last Northeastern Research Station employee

located in the MEF retired in 1983.

Forest genetics experiments have been an important

extension of the artificial regeneration research in the

MEF. In 1959-60 four white pine provenance studies

were planted on old fields owned by Chadbourne

Lumber Company. These lands border the northern unit

and their use made clearing additional land in the MEF

unnecessary. Genetics studies in the southern unit

evaluated weevil resistance in selected half-sib progeny of

western white pine (P. monticola) and local eastern white

pine.

Soil processes were studied at the Littlefield farm on Ida

Jim Road. This farm was in operation from at least

1842, perhaps earlier, until the 1920s, then was

abandoned, and burned in the 1947 fire. During World

War II, it was under cultivation for a few years. Soil pH,

concentrations of total carbon (C) and nitrogen (N),

C:N ratio, and percentage organic matter were

influenced more by agricultural history of the landscape

than by the 1947 fire (Soulia 1997).

Currently, there are no studies in the MEF that require

manipulating plant community composition or

structure. Ongoing research includes owl surveys in the

southern unit and aquatic insect ecology and stream

chemistry monitoring of Cooks Brook in the northern

unit. Since 1998, the Massabesic Experimental Forest

Education Project partners have been involved in

developing and promoting the MEF as a location for

conservation education and demonstration. These

partners include the Forest Service’s Northeastern

Research Station, Maine Forest Service, the town of

Alfred Conservation Commission, the York County Soil

and Water Conservation District, the Small Woodlands

3

Massabesic Rehabilitation Journal. 1956-1959. On file at

USDA Forest Service, Northeastern Research Station, Louis C.

Wyman Forestry Sciences Laboratory, Durham, NH. Typed log.
5

Owners Association of Maine (SWOAM), LaValley

Lumber Company, and the Threshold to Maine

Resource and Conservation District. The MEF

Education Project developed interpretative trails on the

forest with accompanying brochures and conducts tours

for schools and other local organizations. Among these

partners, the Maine Forest Service and SWOAM

strongly advocate a Forest Service research program

focusing on the needs of private owners of smaller forest

lots.

The Inventory, 1997-2000

The MEF mission is to provide a location for

conducting long-term ecological and management

research, and for demonstrating sustainable forest

management in central New England. The primary goal

is to provide research results that develop a better

understanding of the effects of management practices on

forest structure, composition, and function. A secondary

goal is to provide a setting where ecological principals

and management techniques can be presented to

professional and educational groups and the public.

Given the location and history of the forest, and the

interest of stakeholders in managing small forest

properties, the MEF is an ideal outdoor laboratory for

conducting ecology and management research on oldfield sites typical of New England. To provide the critical

knowledge needed by landowners and land managers,

experiments that harvest overstory trees must be

designed and implemented. Such experiments change

the composition and structure of the overstory and

understory. Before a comprehensive research program

can be put into effect, it is necessary to know the

composition and structure of the plant communities that

will be altered when the manipulative experiments are

conducted.

Thus in 1997 we developed a plan to obtain a floristic

description of the MEF and we tested aspects of data

collection. The plan called for a grid of permanent

sample plots across the upland portions of the forest and

a protocol to measure herbaceous and woody plants.

Anecdotal observations of non-vascular plants, lichens,

fungi, and some species of animals also were included.

However, the primary focus was on upland vascular

plants because we were interested in conducting

manipulative experiments on well-drained sites, not

wetlands. We established permanent plots and took

measurements over three field seasons, 1998-2000.

Objectives

The inventory was designed to: 1) gather information

needed to plan experiments in upland habitats that will

answer questions about regeneration, individual tree and

stand growth, productivity, and wildlife habitat; 2)

provide information about forest structure and

composition that could serve as a baseline for comparing

the effects of change due to management practices,

natural disturbance, and ecological succession; and 3)

develop a geographic information system (GIS) database

that can be queried to locate areas that meet defined

criteria, such as tree size or species composition, so that

experiments can be established in the most appropriate

areas of the forest.

Methods

Failing in our attempt to relocate the original plot

system, we established a new grid of plots. To facilitate

the inventory, we used a pre-existing set of 10 arbitrary

parcels plus the administrative site (Figs. 5 and 6). Parcel

boundaries were based primarily on land tenure and to

some extent on municipal boundaries, roads, and natural

features. Consequently, parcels differ in size, topography,

and soils. Parcels are administrative designations rather

than management compartments or forest stands. They

are described below and summarized in Table 1. All soils

information is from Flewelling and Lisante (1987) and

U.S. Department of Agriculture, Natural Resources

Conservation Service, MUIR Database (USDA Natural

Resources Conservation Service 1994). Wetland

information is from Cowardin et al. (1979) and

National Wetlands Inventory (USDI Fish and Wildlife

Service 2002).

Prior to the start of any field work, we familiarized

ourselves with the rare plant list for Maine, which is an

administrative list maintained by the Maine Natural

Areas Program (Maine Department of Conservation

1999), and a checklist that provided a county-by-county

listing of herbarium records (Campbell et al. 1995), so

that we could better document rare plant populations

while collecting field data.

Description of parcels

Northern Unit

Parcel N1 (36 acres) is a narrow, isolated parcel north of

Maine Route 5 in Lyman (Fig. 5). The west boundary is

Cooks Brook, which is in the Saco River drainage. An

abandoned railroad bed crosses the parcel at the south

end near Route 5. Forested wetlands associated with

Cooks Brook bound the parcel at the southwest with

another small wetland near the center of the parcel. At

the north end of the parcel there is a stand of mature

northern hardwoods with sugar maple (Acer saccharum),

yellow birch (Betula alleghaniensis), a few white ash

(Fraxinus americana), red oak, hemlock, and white pine.

Within the recent past, the private woodlot west of this

stand was clearcut to the MEF boundary. Soils are
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Figure 5.—Location of permanent sample points in the northern unit, Massabesic Experimental Forest,

and parcels used for summarizing data.
7

mostly stony, very fine sandy loams, well to somewhat

excessively drained Skerry and Hermon, with a small

proportion of poorly drained Brayton and some

excessively drained, gravelly, loamy sand (Colton).

Hydric soils include small areas of Naumburg sand and

Chocorua peat.

Parcel N2 (total area 979 acres, area sampled 881 acres)

is located at the northern end of the unit (Fig. 5) in the

towns of Lyman, Hollis, and Dayton. Topography is

varied with elevation ranging to the highest point in the

MEF at about 450 ft above sea level, with large and

small forested wetlands, vernal pools, and the swell and

swale undulations left by glaciation. Cooks Brook flows

easterly across the northern part of the parcel, eventually

to the Saco River. The southern boundary includes

frontage on Roberts Pond, which is an impoundment of

Tarwater Stream, and wetlands associated with that

body. Small pockets of scrub shrub and forested

wetlands are found throughout this parcel.

A flat, sandy outwash area where Chicken Farm Road

extends east to the MEF boundary was abandoned

farmland that was harvested prior to the 1947 fire. This

area, which burned intensively in 1947, was “mostly

brush land prior to the fire, with a fair scattering of

white pine saplings.”3

 A salvage harvest was conducted

here following the fire. The area was converted to pine

plantations in the 1950s. The riparian zone of Cooks

Brook broadens to the east of CC Road, the main northsouth forest road, with a ravine up to 70 ft deep and

about 500 ft wide. Some pitch pine and bear oak (Q.

ilicifolia) are present near CC Road, which may be

remnants of the presettlement forest. A 0.5-acre jack

pine (P. banksiana) plantation was established in 1951

near a borrow pit off Chicken Farm Road. The Civilian

Conservation Corps, after which CC Road was named,

established this pit in the 1930s and the deposits

probably were used to build the road system in the unit.

In 1954, about 4 acres were rototilled and in 1955 white

and red pine were planted. The red pine was replanted in

1956 due to poor survival. West of the access road to the

borrow pit is an area that was treated with an aerial spray

of 2,4,5-trichlorophenoxy acetic acid (2,4,5-T) in 1954

and planted with white pine, but details of this planting

are not included in the MEF Journal.3

 Most of the

planted strip between Chicken Farm Road and the

southern boundary of the parcel was treated with an

aerial application of 2,4,5-T in 1956. On the southern

boundary of the treated area, 7 acres were planted with

white pine in 1958.

In the southeast part of this parcel are stone walls, dug

wells, and cellar holes from earlier farms. This area,

Table 1.—Total area, area sampled, and numbers of sample points and meander transects in

the inventory of upland vascular plants in the Massabesic Experimental Forest, by parcel.

Parcel Total area Area sampled Sample points Transects

acres

N1 36 36 11 10

N2 979 881 170 176

N3 335 91 22 25

N4 318 35 11 12

 Total,

 northern unit 1,668 1,043 214 223

S1 167 37 13 13

S2 167 107 22 24

S3 1,336 593 114 135

S4 58 54 17 12

S5 36 25 8 8

S6 192 36 15 12

S7 (Admin site) 52 0 0 0

 Total,

 southern unit 2,008 852 189 204

Total 3,676 1,895 403 427
8

which is known locally as Thirteen Cellar Holes, is

located on a forested hill that is most easily accessed

from the settlement at Roberts Pond. Soils support a

small population of sugar maple and shagbark hickory

(Carya ovata); the former is unusual in the MEF and the

latter is known only from this one locale. An inholding

on the east side of the parcel is near the summit of the

highest promontory in the MEF. A narrow ravine among

granitic ledges seems to have remained unburned when

the 1947 fire consumed the humus layer on the slope to

the west of the ravine.

Soils in Parcel N2 are predominantly stony, fine sandy

loams, including somewhat excessively drained Hermon

and poorly drained Brayton. Loamy sands are also

important components including well-drained Adams,

moderately well-drained Croghan, and small amounts of

excessively drained Colton. A small amount of granitic

rock outcrop has been mapped. Hydric soils include

small amounts of Naumburg sand, and Chocorua and

Sebago peats. Some large glacial erratics are present but

infrequent.

Parcel N3 (total area 335 acres, sampled area 91 acres) is

located along Jellerson Road in Lyman (Fig. 5). It is

bounded on the north by a wetland associated with the

Tarwater Pond outlet, which feeds into Roberts Pond. A

large scrub shrub wetland to the north and east (labeled

“heath” on the USGS quadrangle) extends onto private

land. Forested wetlands that are typically dominated by

red maple (Acer rubrum) extend to the south and west.

The sampled area burned in 1947, and pine was salvaged

post-fire. Rehabilitation efforts included small

plantations of white and red pine south of the woods

road from Jellerson Road to Tarwater Pond in 1948 and

1949. Part of the area was sprayed with 2,4,5-T in 1955

and then white pine was planted in 1956. Moose Point

Road crosses the parcel in the southeastern portion. Soils

are mostly loamy sands including well-drained Adams

and moderately well-drained Croghan. Hydric soils

fringe the sampled areas and include peat types such as

Chocorua, Vassalboro, and Sebago with a small amount

of Naumburg sand.

Parcel N4 (total area 318 acres, sampled area 35 acres) is

divided into east and west portions by Jellerson Road

(Fig. 5). The south boundary includes part of Tarwater

Pond and its associated drainage. An inholding of private

land is in the southeast portion. Parcel N4 burned in

1947. In 1957 it was treated with an aerial spray of

2,4,5-T and then a part of the area was planted with

white pine. Reforestation of the area east of Jellerson

Road appears to have been successful, but efforts on the

west side were not. Due to boulders, stumps, ledge

outcrops, and boggy spots on the west side, this area was

considered “not plantable.”3

 The soil of this parcel is

mapped as Hermon, a stony, very fine sandy loam. Small

pockets of scrub shrub wetlands occur throughout the

parcel, and a forested wetland is southwest of the

sampled area.

Southern Unit

Parcel S1 (total area 167 acres, sampled area 37 acres) is

north of Old Kennebunk Road in Alfred (Fig. 6). Old

agricultural fields are now covered with second growth

pine-oak in a rolling topography with many stone walls.

Nearly all the soils in the sampled area are stony, very

fine sandy loams, and include Hermon, which is

somewhat excessively drained, and Brayton, which is

poorly drained. The three plots on the western side of

this parcel are in a forested wetland and are underlain by

Naumburg sand, a hydric soil. There is a large sedge

meadow along Old Kennebunk Road, the hydrology for

which was probably influenced by the building of the

road through the wetland. This wetland is noteworthy

because despite its proximity to a residential area, it does

not contain such invasive plants as purple loosestrife

(Lythum salicaria L.), common reed grass (Phragmites

australis [Cav.] Trin. Ex Steud.), or exotic shrubs, such as

smooth buckthorn (Frangula alnus).

Parcel S2 (total area 167 acres, sampled area 107 acres) is

an isolated parcel on the north side of Old Kennebunk

Road in Lyman (Fig. 6). Second growth pine-oak

dominates the uplands. Topography is especially diverse,

with a steep 40-ft rocky ledge, rolling hills, and

numerous small wetlands including vernal pools. The

largest wetland (7.4 acres) is a red maple swamp that

drains to the northwest into Carlisle Brook, and thence

to the Kennebunk River. Old stone walls, several cellar

holes, and a woods road were established in the 1800s or

possibly earlier. This suggests that the upland soils were

plowed or pastured. A family cemetery is adjacent to the

southeast corner of the parcel. Prior to 1947, there was a

stand of pine presumably planted around 1925 in the

northwest corner of the parcel and an adjacent stand of

natural pine, possibly on an old field, that grew on the

slope along Old Kennebunk Road. In both areas, the

1947 fire damaged most of the pine. Four acres

northeast of the woods road and along Old Kennebunk

Road were planted with white and red pine in 1948. The

remainder of the frontage along Old Kennebunk Road

was treated with a basal spray of 2,4,5-T, followed by

cutting stems within 1 foot of pine regeneration in 1954.

A 15-acre area north of the woods road was part of a test

aerial application of dilute 2,4,5-T amine. Poor coverage

made treatment relatively ineffective. Soils are mostly

stony, very fine sandy loams including a large proportion
9

  

























  

 





 

 













































































      































































 



























































 



















 

 

   



































  





 



 









 







 











	
	




	






		















 









 


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Figure 6.—Location of permanent sample points in the southern unit, Massabesic Experimental

Forest, and parcels used for summarizing data.
10

of Hermon (somewhat excessively drained) with smaller

amounts of Skerry (moderately well-drained), Becket

(well-drained), and Brayton (poorly drained). Hydric

soils occur in small amounts, including Sebago and

Chocorua peats. Rock outcrops occur in a few places

and feature some plant species usually not found

elsewhere in the MEF.

Parcel S3 (total area 1,363 acres, sampled area 593 acres)

is located in Alfred and Lyman (Fig. 6) on the north side

of Estes Lake, an impoundment of the Mousam River.

Access is from various local roads: Whichers Mills Road

(southern portion), Mouse Lane (west), the unpaved Ida

Jim Road, named after members of the Littlefield family

who owned a farm (western portion), Old Kennebunk

Road (northern portion), and the unpaved Government

Road (eastern portion). Vegetation ranges from mature

white pine and hemlock to red maple swamp. Topography

is highly varied. Wetlands are numerous and include

patches of forested wetland, streams, scrub shrub, vernal

pools, and a swell and swale system of glacial outwash

ridges, kames, and possible kettleholes. The largest

wetland is an Atlantic white-cedar swamp complex that

bounds the sampled area to the east and northeast.

Uplands east of Ida Jim Road burned in 1947. White

and red pines were planted in 1948 on a 6-acre site of

former farmland south of Bermed Road (which is

perpendicular to and extending east from Ida Jim Road).

Records show that the western portion of this site was a

cultivated field in 1946; the remainder was old field.3

Numerous small plantations of hybrid poplars were

planted in 1948 and 1950 in addition to a 2.5-acre

plantation of white and red pine established in 1948. At

the end of Bermed Road, thinning was conducted on 23

acres in the early 1950s, but hurricanes blew down many

of the remaining trees in 1954. This area was planted

with red pine in 1956 and an additional 12 acres in the

vicinity were treated with an aerial application of 2,4,5-

T. Experiments using varying amounts of 2,4,5-T were

conducted in 1957 in these same areas to either control

hardwoods or prepare areas for planting. An application

of 2,4,5-T was made to a 5-acre area at the northernmost

boundary along Ida Jim Road to release pine seedlings

overtopped by birch and red maple. A natural stand (0.7

acres) on the north side of Bermed Road was thinned in

1957 and then sprayed with DDT in 1958 to control

white pine weevil. At the corner of Government Road

and Whichers Mills Road, 3.5 acres of hardwood

saplings were treated with a basal stem application of

2,4,5-T, then planted with white and red pine in 1955.

The 1947 fire burned 130 acres northeast of the

intersection of Ida Jim Road and Whichers Mills Road

and killed white pine smaller than 4-6 inches diameter at

breast height (dbh). In 1956, trees remaining in the

stand were released to provide a future seed source. On

the south side of Whichers Mill Road, Scotch pine (P.

sylvestris) was planted to compare growth rate and form

of selected families. Western white pine (P. monticola)

and Himalayan blue pine (P. wallichiana) were planted

nearby to compare growth rates and weevil resistance

with eastern white pine.

The majority of soils in the sampled area are Hermon

(somewhat excessively drained) and Skerry (moderately

well-drained); both are stony, very fine sandy loams.

These are generally found on the swells. Brayton, a

poorly drained, stony, very fine sandy loam, is generally

found in the swales. Present in smaller amounts are:

Allagash, a well-drained, very fine sandy loam; Adams, a

well-drained loamy sand; and Madawaska, a somewhat

poorly drained, fine sandy loam. Hydric soils include

small amounts of Chocorua peat, Sebago peat, and

Naumburg sand.

Parcel S4 (total area 58 acres, sampled area 54 acres) is in

Alfred (Fig. 6) on the south side of Whichers Mills Road

and the northern shore of Estes Lake. The topography is

gently sloping to the south and there are a few small,

forested wetlands. There is evidence of granite quarry

activity presumably from the 1800s. A 5-acre plantation

of white pine was established on the western boundary.

This area is described as a “...rather open, merchantable

stand of white pine burned in 1947. There were

indications of an earlier fire about 1920 - consistent

rather long butt scars on the larger trees.”3

 Soils are

mostly mapped as rock outcrop with a small proportion

of Croghan, which is a moderately well-drained loamy

sand. Brayton, a poorly drained, stony, very fine sandy

loam, can be found in areas of lower elevation. Wetlands

are on two hydric soils: Chocorua peat and Naumburg

sand.

Parcel S5 (total area 36 acres, sampled area 25 acres) is in

Alfred (Fig. 6) on the north side of Whichers Mills Road

and is crossed by Shackford Farm Road, an unpaved

woods road that proceeds north to Old Kennebunk

Road. Topography is moderately rolling with some

forested wetlands at the northwest corner of the parcel

and along the southeast boundary. A stream flows

southeast and crosses Shackford Farm Road at an old

homestead, then cuts deeply into the slope before

entering a scrub shrub wetland. Natural stands are

mostly pine-oak with hemlock and a small area of pitch

pine on a ledge to the southeast. Prior to the 1947 fire,

the parcel was stocked with 60-year-old white pine and a

band of hardwood less than 10 inches dbh running

north to south through the center of the parcel. Except

for some pine in the northeast corner, most of the parcel

burned in 1947. Many of the surviving trees blew down
11

in the hurricanes of 1950 and 1954. In 1948, white and

red pines were planted on 1 acre near the northwest

corner of the parcel along Whichers Mills Road. Five

acres were released in 1954 by cutting competing trees

and treating stumps with 2,4,5-T. In the plantation,

hardwoods were girdled and ammate crystals applied to

the wounds. A gypsy moth (Lymantria dispar [L.])

infestation in 1954 defoliated the red and white pines,

slowing growth but not killing the trees. Further east on

Whichers Mills Road, all hardwoods and pulpwood were

salvaged on 4 acres and stumps were treated with

ammate (2,4,5-T) in 1955. Three of the treated acres

were planted with red pine in 1956. Soils in this parcel

have been mapped as mostly rock outcrop with some

Adams, which is a well-drained loamy sand, and a small

amount of Brayton, a poorly drained, stony, very fine

sandy loam. Hydric soils include a large proportion of

Naumburg sand and some Chocorua peat.

Parcel S6 (total area 192 acres, sampled area 36 acres) is

in Alfred (Fig. 6) along Old North Berwick Road, and is

mostly flat with a scrub shrub and forested wetland

complex to the northwest, including small remnants of

long-dead Atlantic white-cedar. Logging was conducted

between 1930 and 1940, and regrowth burned in 1947.

A 3.5-acre area on the north side of Old North Berwick

Road was planted with white and red pine in 1948. A

strip of jack pine was planted on the north side of Old

North Berwick Road in 1951. Seventeen acres north of

that road were planted with white and red pine in 1955.

The red pine was replanted in 1956. A 12-acre site north

of these plantations was planted with white pine in

1958, but survival was poor. South of Old North

Berwick Road, 20 acres were planted with white pine in

1958. Nearby, some burned areas have regenerated to

pitch pine. Soils in this parcel are Adams and Croghan,

which are mostly well-drained and moderately welldrained loamy sands. Smaller areas of two hydric soils,

Chocorua peat and Naumburg sand, have also been

mapped in this parcel.

Parcel S7 (52 acres) is the administrative site for the

MEF. It is on the north side of Maine Route 111 in

Lyman (Fig. 6). It includes Forest Service buildings that

once served as MEF headquarters. Now state and county

agencies occupy the buildings. Topography is rolling,

with a few patches of forested wetlands and seasonal

streams that flow into Carlisle Brook. Vegetation is pineoak on former farm fields, with stone walls and cellar

holes. Invasive shrubs and vines are more common here

than elsewhere in the MEF. The area “...north of office

planted with white and red pine in 1949 and 1954...a

few Austrian pine and Scotch pine were also planted in

1955.” 3

 In 1956, white pine was planted on 2 acres west

of the office and near Route 111. Soils in this parcel are

stony, very fine sandy loams. Hermon, excessively welldrained, is the greatest proportion, followed by Brayton,

which is poorly drained, and there is a small amount of

moderately well-drained Skerry. There are a few small

forested wetlands here and there in the parcel. One small

scrub shrub wetland is mapped near the south boundary

at Route 111. No sampling points were established in

this parcel.

Field Data Collection

Field data were collected during the summers of 1998-

2000 using methods developed in a reconnaissance

survey in 1997. Variables are listed in Table 2. Common

and scientific names for trees follow Little (1979).

Names for all other plants follow Haines and Vining

(1998) and unpublished updates supplied by these

authors.

Grid of permanent sample points

Wetlands are extensive in some parcels, but sampling for

this inventory was mostly in uplands because

manipulative experiments will be conducted on higher

ground. Thus sampling covered one small parcel

completely (N1), but for most parcels only a portion of

the total area was sampled (Table 1). A parcel may

contain one or more stands and may eventually be

divided into one or more research compartments. A 330

x 660 ft (5 x 10 chain) grid was established wherever

sufficient upland forest was available. The grid was laid

out from an east-west baseline established for each parcel

at an easily identified corner, typically monumented with

a pre-existing granite post or painted boulder (Figs. 5

and 6). Crews of three to five people used staff compass

and tape to survey each sampling point, based on true

north. We monumented each point with an 18-inch

stainless steel stake topped with a yellow plastic cap.

Sampling procedures

Each sampling point served as a permanent location for

variable radius plot sampling (VRP) (Schreuder et al.

1993) and as the center for a 0.01-acre fixed-radius plot

(radius = 11.77 ft). We used VPR to tally trees 4.5 inches

dbh or greater with a 10 basal area factor (BAF) prism.

We identified trees to species, measured their dbh with a

diameter tape to the nearest 0.1 inch, and recorded each

as live or dead. Any dead snag 4.5 ft tall or more that we

encountered using the prism was tallied as a dead tree if

its dbh was 4.5 inch or greater.

In the 0.01-acre plots, trees between 0.5 and 4.5 inches

dbh were tallied by species, diameter, and as live or dead.

Woody stems were counted by species if taller than 1 ft

but less than 0.5 inch dbh. Tree and shrub seedlings were

identified to species and scored as “present” if less than 1
12

ft. For example, if at least one red oak seedling less than1

ft was found, that species was noted as present in the

plot. Abundance of each plant species was estimated in

the 0.01-acre plot using percentage cover classes of

Witham et al. (1993). This involved visualizing each plot

as a cylinder rising through the canopy and classifying

the relative abundance for each species within the

cylinder. For species that occupied less than 5 percent of

the plot, three classes were used: 1) Rare, 5 or fewer

individuals; 2) Occasional, numerous individuals but

not common; 3) Common, occurred more or less

throughout the plot but with coverage less than 5

percent. For species that occupied 5 percent or more of

the plot, classification was in four additional classes: 4)

5-25 percent; 5) 26-50 percent; 6) 51-75 percent; and 7)

76-100 percent. Height also was classified. If a woody

stem was shorter than 0.8 ft, it was assigned to class 1,

otherwise it was estimated and assigned to a class as

follows: 2) 0.8-3.4 ft; 3) 3.5-17 ft; 4) 18-34 ft; and 5)

taller than 34 ft.

We also tallied any species not inside the plot but

outside and within sight of it (about 30 - 50 ft radius,

depending on homogeneity of habitat) as being in the

vicinity of the sample point. These off-plot data were

used to develop a more complete plant list for the MEF,

not for any quantitative analyses.

A disturbance category was determined for each sample

point based on evidence within or near the plot (i.e., 50-

ft radius). A plot was classified as “Agriculture” if there

were stone walls or foundation holes, “Harvest” if there

were cut stumps, “Fire” if there were fire scars and/or

charcoal (with no further classification, i.e., recent vs.

presettlement fire), “Plantation”, or “None” if there was

no obvious disturbance. We did not include evidence of

wind throw, though we noticed this on occasion.

Meander transects

Although we assume that data from VRP sampling and

fixed-area plots are representative of the upland

vegetation in the MEF, we wanted a more complete

picture of the flora of the MEF. To augment the VRP

and plot inventory, a series of transects was established

along the north-south lines between sample points.

Between one fixed plot and the next (approximately 307

ft), all vascular plants were recorded within a transect

width of about 15 ft for the herb stratum and about 100

ft for the tree stratum. Transects were not fixed in width.

If a feature, such as a vernal pool, rocky outcrop, stream,

or glade, could be seen from a transect, we visited the

area and recorded all plant species. We included

wetlands in these meander transects, while passing from

one upland plot area to another. Abundances for all

species were assigned according to methods of Palmer et

al. (1995). Like the off-plot data, information collected

along meander transects was used to develop a more

complete plant list and was not used in any quantitative

analysis.

Table 2.—Variables measured or recorded in the inventory of upland vascular plants in the

Massabesic Experimental Forest.

Sample method Taxa and strata sampled Variable Measurement units

Variable radius plot trees ≥ 4.5 inch dbh name species

dbh nearest 0.1 inch

status live or dead

Fixed area plot trees ≥ 0.5 dbh <4.5 inch name species

dbh nearest 0.1 inch

status live or dead

trees < 0.5 inch dbh but ≥ 1 ft tall name species

stem count number

trees < 1 ft name species

abundance class

shrubs ≥ 1 ft name species

stem count number

shrubs < 1 ft name species

abundance class

nonwoody plants name species when possible

abundance class

Meander transect all name species when possible
13

GIS database development

To facilitate use of forest vegetation data gathered in the

inventory with other available spatial data, we developed

a geographic information system (GIS). By referencing

VRP, fixed plot, and transect inventory data with the

geographic location from which it was obtained, the

vegetation data can be viewed and analyzed in the

context of other geographic datasets, such as soils,

hydrology, topography, and disturbance history. Many

such datasets exist and are available from various federal,

state, or private sources. The GIS for the MEF is made

up of the following layers: a base map with boundaries,

aerial photos, roads and trails, wetlands, streams, ponds,

rivers, drainage divides, soils, 1947 fire map, forest cover

types, permanent sample points, meander transects, and

most of the data we gathered in the field.

Digital, raster based, U. S. Geological Survey 7.5

minute, 1:24,000 scale quadrangles were used as base

maps (USDI Geological Survey 2000). Forest boundary

layers were created as vector files and registered on the

base map. True color aerial photographs were taken in

June 1999 at a scale of 1:20,000. We obtained both

photographs and digital (1320 dots per inch) stereo pair

images from J.W. Sewall Company, Old Town, ME. We

created a mosaic of the images to provide complete

coverage of the two units. We used the Universal

Transverse Mercator (UTM), North Atlantic Datum 83

(NAD83) projection for the photo mosaics and the

boundaries so that they could be used most easily with

data layers obtained from the Maine Office of

Geographic Information Systems (MEGIS).

We obtained the National Wetlands Inventory (NWI)

layer (USDI Fish and Wildlife Service 2002) through

MEGIS. We created fields to translate the wetland codes

provided so that wetlands could be sorted by system,

class, and subclass (Cowardin 1979). This allows the user

to easily identify polygons representing, for example:

palustrine (system), forested (class), broad-leaved

deciduous (subclass) wetlands. Data layers containing

roads and streams as vectors came from MEGIS. Roads

were categorized by type from interstate to trail. Stream

segments were categorized as perennial or intermittent in

the attribute tables. We obtained polygon data layers

containing rivers, ponds, and drainage divides from

MEGIS. In addition, we incorporated vector files

containing coordinates taken with the use of a

geographic positioning system (GPS) receiver on some

of the trails and woods roads in the MEF.

Two other data layers were added to the GIS. The soils

data were obtained as a polygon layer from the USDA

Natural Resources Conservation Service, National Map

Unit Interpretation Record (MUIR) Database (USDA

Natural Resources Conservation Service 1994).

Attributes available include soil texture, soil erodability

factors, hydric soil ratings, seasonal ponding, likelihood

of flooding, water table depth, bedrock depth, and site

indices of common trees. A map of southwestern Maine

showing an estimate of fire severity and remaining

timber type and size class was prepared immediately

following the fire of 1947 (USDA Forest Service 1947).

Portions that featured the MEF were digitized and

categorized by the type of fire sustained—ground,

crown, or none. Despite its historical interest, we

consider the resolution of the 1947 fire map to be too

coarse to provide a detailed description of fire in the

MEF.

We created the forest-cover type layer based on data

collected in the upland areas, then combined with NWI

data to determine wetland cover types. For each sample

point we determined percent basal area (dbh ≥ 0.5 inch)

of each tree species. Using an inverse distance-weighted

method, we interpolated percentage cover for each major

tree species across the forest. For each 10 m2 pixel we

determined a cover type by calculating percentage basal

area from the interpolated values of the major tree

species. Forest-cover types defined for the upland

portions of the MEF follow the types recognized by

Forest Inventory and Analysis (FIA).4 Because part of our

tree data was collected using VRP sampling, rather than

exclusively in fixed area plots as is done by FIA, we used

percentage basal area of each species as its stocking value

rather than determine stocking as specified by FIA

methodology.

Single species types included white pine, hemlock, red

pine, and Atlantic white-cedar when those species

accounted for more than 50 percent of the basal area at a

sample point. Where softwoods dominated multiplespecies composition, the white pine—hemlock forest

type was most common. If hardwoods exceeded 50

percent total basal area, and the percentage of white pine

plus red oak exceeded 25 percent, then the forest-cover

type was classified as pine—oak. For samples where pine

was not present with oak, cover type was classified as

either red maple or red oak, depending on whether

either species was more than 50 percent of the basal area.

For the few points where red oak did not meet this

4

Arner, S.L., Woudenberg, S.; Water, S. [and others].

Unpublished. National algorithms for determining stocking

class, stand size class, and forest type for forest inventory and

analysis plots. [Documentation available from the

Northeastern Research Station, Forest Inventory and Analysis,

11 Campus Blvd., Newtown Square, PA 19073].
14

criterion but white oak (Quercus alba) was a significant

component, the oaks were combined. If all oak species

combined met the 50 percent threshold, they were

classified as red oak. One mixed-hardwood type was

present in the MEF: the maple—American beech (Fagus

grandifolia)—yellow birch type. For some sample points,

we added paper birch or sweet birch (Betula lenta) to the

percentage basal area of maple (mostly red), beech, and

yellow birch to determine this type. Because plot

locations were biased toward the upland areas and the

inventory avoided many of the wetlands, we

incorporated the national wetlands inventory cover types

into the upland coverage we had created and allowed the

NWI-derived types to take precedence in the wetlands.

Approximate locations of sample points and transects

were established in the GIS by using coordinates from

the base map and boundary layer to determine the origin

of the grid for each parcel. Inventory data can be

accessed from the GIS by unit, parcel, or plot.

Data summary and analyses

Data from trees between 0.5 and 4.5 inches dbh

collected in the fixed plots and from larger trees by VRP

sampling, were summarized using forest inventory

software (Brann and Solomon 2001). Number of trees,

basal area, and volume/acre were summarized by species

and for all species for each parcel. The inventory

software calculated volume based on regional height to

diameter relationships by species contained in the

inventory software because no tree height measurements

were taken as part of the MEF inventory. Abundance of

seedlings taller than 1 ft but less than 0.5 inch dbh was

summarized as seedlings/acre by species, by parcel. We

summarized results of the inventory of live trees (≥ 0.5 in

dbh) in a series of tables (Tables 3-12) and figures (Figs.

9-18) by 1-inch diameter classes by parcel. Species were

ranked in order by number of trees/acre in the tables.

Diameter distributions were plotted in stacked bar

graphs by species. We used the four most abundant

species and an “all other species” category, and inset the

smallest diameter classes (1 - 4 inches) in each of the

larger graphs (> 4 inches).

We summarized data for plants other than trees by

frequency, or percentage of plots by parcel or unit in

which a species occurred, regardless of height class.

Plants recorded in the vicinity of sample plots and in the

meander transects are included only as “present” in the

overall list of plants (Appendix I). We used SAS ver. 8.2

— especially PROC MEANS, PROC FREQ, and

PROC GCHART (SAS Institute Inc. 1999) — to

summarize the understory plants and create most figures.

Results and Discussion

We collected and summarized data from 399 VRP

points and their corresponding 0.01-acre plots. Of these,

211 plots are in the northern unit (Fig. 5) and 188 are in

the southern unit (Fig. 6). An additional four plots were

monumented but not incorporated in the data

summaries for various reasons (Table 1). In addition,

plants recorded from 223 meander transects in the

northern unit and 204 in the southern unit were

included in the overall plant lists for the parcels

(Appendix I). We observed about 500 plant taxa,

including subspecies. The number is approximate

because some herbaceous plants could not be

conclusively identified from the material at hand. A

single plot contained as few as eight or as many as 67

species.

We found six forest cover types, discerned from GIS, in

the uplands. This represents more than 2,500 acres (Figs.

7 and 8). The most common of these is the white pine

type with more than 900 acres, followed by pine-oak

with more than 800 acres and pine-hemlock with

slightly more than 700 acres. The types differ between

the two units in terms of tree species composition and

forest structure. The northern unit contains much more

pine-oak and scrub shrub than does the southern unit.

Atlantic white-cedar swamp, a type that is rare in Maine,

is relatively abundant in the southern unit with a total of

160 acres. Total acreage of plantations is a matter of

interpretation because many of the early attempts at

establishing plantations failed.

Vascular Plant Diversity

The inventory resulted in identification of 464 vascular

plants in the MEF representing 82 families and 229

genera (Appendix I). We found very few invasive, exotic

plants. In disturbed habitats in southern Maine, there

can be dense local populations of non-native species,

such as Japanese barberry (Berberis thunbergii), Asian

honeysuckles (Lonicera spp.), Oriental bittersweet

(Celastrus orbiculata), Russian olive (Elaeagnus

angustifolia L.), autumn olive (Elaeagnus umbellata

Thunb.), and purple loosestrife. These are troublesome

because they displace native species, alter wildlife feeding

behavior, offer different wildlife cover than native

vegetation, have an overall adverse impact on

biodiversity, and could affect fuel loads (Richburg et al.

2001). We interpret the low frequency of exotic weedy

species in the MEF as an important indication of

ecosystem integrity in spite of intense agricultural

activity on some parcels since the 1600s and

establishment of plantations with non-native tree species.
15

Figure 7.—Forest cover types, northern unit, Massabesic Experimental Forest.

The inventory includes 12 species of coniferous trees and

32 species of deciduous trees that were part of, or could

potentially grow into, the overstory. In the understory,

there are nine tree species that are unlikely to be found

in the overstory except on unusual sites or following

uncommon disturbances, including black gum (Nyssa

sylvatica), shagbark hickory (Carya ovata), sugar maple

(Acer saccharum), black cherry (Prunus serotina), eastern

hophornbeam (Ostrya virginiana), and American

hornbeam (Carpinus caroliniana). The understory

vegetation includes 75 species of woody shrubs and

subshrubs, 189 species of broad-leaved herbaceous

plants, 47 species of grasses, 75 sedges and rushes, 39

species of ferns, horsetails, and clubmosses, and nine

species of vines (Appendix I).

To get a sense of the diversity of vascular plants in the

MEF, this inventory can be compared with the entire
16

    

	




















	







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Figure 8.—Forest cover types, southern unit, Massabesic Experimental Forest.
17

state of Maine where 139 families with 699 genera and

2,096 species of vascular plants have been identified

(Haines and Vining 1998). The flora of Maine is not

static and the numbers of taxa change over time.

Furthermore, we sampled only half the area of the MEF

in this inventory and think more plant species certainly

occur within its boundaries. We think at least 25 percent

of the vascular plant taxa that occur in Maine are found

in the MEF.

Overstory Trees

White pine, hemlock, red oak, and red maple are the

most abundant species in most parcels. Red maple was

found on 94 percent of plots, white pine on 86 percent,

and red oak on 84 percent. Gray birch (Betula

populifolia) was found in all parcels, and paper birch in

all but one. We found dense stands of young hemlock

especially on slopes in stands that had burned in 1947.

These thickets probably regenerated on intensely burned

sites within a few years of the fire. The hemlock density

data reflect these thickets but do not reveal their

patchiness.

Parcel Summaries

Parcel N1 — The area sampled (36 acres) has 1,139 live

trees/acre, basal area of 137.4 ft2

/acre, and volume of

3,539 ft3

/acre (Table 3). This parcel contains about onethird softwood species by basal area. Important

softwoods include white pine and hemlock in terms of

number of trees and basal area, and important

hardwoods include red maple and red oak. These four

species account for slightly more than three-quarters of

the basal area. The larger trees in this parcel are red oak

and white pine as shown by the relatively small number

of trees (11.5 percent of total) compared to high timber

volume (50.5 percent of total). This is also shown in

Figure 9, which plots the number of trees/acre by 1-inch

diameter classes. Without disturbance, red oak and white

pine could gradually be replaced by red maple, hemlock,

and American beech (which are lumped into “Other” in

Fig. 9) in Parcel N1.

Parcel N2 — This is the largest parcel in the northern

unit, 979 acres, and second largest in the MEF. The area

sampled (881 ac) has 1,843 live trees/acre, basal area of

Table 3.—Number of trees and basal area/acre for all live trees ≥ 0.5 inch dbh and gross

volume/acre for all live trees ≥ 4.5 inch dbh by species and species group in Parcel N1,

northern unit, Massabesic Experimental Forest.

Species Number of Basal area Percent Volume

trees basal area

ft2

/acre ft3/acre

Softwoods

Eastern hemlock 174 21.8 15.9 459

Eastern white pine 93 27.5 20.0 788

Balsam fir 5 0.9 0.6 17

Subtotal 272 50.2 36.5 1,264

Hardwoods

Red maple 346 28.7 20.9 692

American beech 266 8.0 5.8 165

Yellow birch 81 9.9 7.2 209

Paper birch 42 5.0 3.6 93

Northern red oak 39 28.4 20.6 998

White oak 36 1.7 1.2 —

Sweet birch 27 1.0 0.7 —

Moose maple 9 <0.1 <0.1 —

Gray birch 9 <0.1 <0.1 —

White ash 6 2.7 2.0 71

Sugar maple 4 0.9 0.6 19

Quaking aspen 2 0.9 0.6 28

Subtotal 867 87.2 63.5 2,275

Total 1,139 137.4 100.0 3,539
18

155.2 ft2

/acre, and volume of 3,246 ft3/acre (Table 4).

This parcel is slightly more than half softwood species by

basal area. Important softwoods are white pine and

hemlock, in terms of numbers of trees and basal area.

Important hardwoods, especially regarding basal area, are

red oak and red maple. These four species account for

more than 80 percent of the basal area. Large trees in this

parcel are red oak and white pine as indicated by the

relatively small number of trees (12.6 percent of total)

compared to high timber volume (62.6 percent of total).

Without disturbance, red maple and particularly

hemlock will eventually represent a greater proportion of

basal area and volume in this parcel but white pine and

red oak are well represented in all diameter classes (Table

4 and Fig. 10).

Parcel N3 — Most of the area sampled (91 acres) is in

white and red pine planted after the 1947 fire. The area

sampled has 550 live trees/acre, basal area of 168.2 ft2

/

acre, and volume of 4,073 ft3

/acre (Table 5). This parcel

contains 94 percent softwood species by basal area.

Important softwoods include planted white pine and red

pine, in terms of number of trees and basal area. The

small component of hardwoods (< 6 percent of basal

area) consists of pioneer species such as gray birch. More

than 90 percent of the basal area is pine. Ninety-five

percent of the total volume is white and red pine with

white pine comprising most. The diameter distribution

of the parcel is characteristic of an even-aged planting

(Fig. 11).

Parcel N4 — Much of the area sampled (35 acres)

burned in the 1947 fire and was planted with pine. The

plantation east of Jellerson Road survived but the

plantation west of the road failed. This resulted in a mix

of softwood plantation and pioneer hardwood species.

Parcel N4 has 2,059 live trees/acre, basal area of 171.5

ft2

/acre, and volume of 2,872 ft3/acre (Table 6). This

parcel is slightly more than half softwood species by

basal area. Important softwoods include white pine

(mostly planted) and hemlock, in terms of number of

trees and basal area, and important hardwoods include

red maple and paper birch. These four species account

for 80 percent of the basal area, of which white pine

accounts for nearly half. More than half the volume in

the parcel is white pine, while red maple, red pine, paper

birch, and red oak account for most of the remainder.

The mix of softwood plantations and natural, mixed

wood stands has an effect on the diameter distribution of

the parcel in that the larger trees (≥ 5 inch-diameter

Figure 9.—Number of trees per acre by diameter classes and major tree

species in Parcel N1, northern unit, Massabesic Experimental Forest.
19

class) on the parcel are mostly planted white pine while

hemlock and red maple are dominant in smaller

diameter classes (< 5 inches) (Fig. 12).

Parcel S1 — The area sampled (37 acres) has 3,220 live

trees/acre, basal area of 178.9 ft2

/acre, and volume of

2,629 ft3/acre (Table 7). This parcel contains about twothirds white pine and hemlock, by basal area. Important

hardwoods are red maple and red oak, in terms of

number of trees and basal area. These four species

account for more than 86 percent of the basal area.

Nearly half of the volume in the parcel is white pine,

while red oak, red maple, and hemlock account for most

of the rest. Parcel S1 is former agricultural land that

reverted to forest. Most of the hemlock trees are small.

This species accounts for 74 percent of the total trees but

only 37 percent of the basal area. Hemlock less than 6

inches dbh is especially well represented (Fig. 13).

Parcel S2 — This parcel burned extensively in 1947.

There were a few small plantings of pine following the

fire, but most of the area was left to regenerate naturally.

Table 4.—Number of trees and basal area/acre for all live trees ≥ 0.5 inch dbh and gross

volume/acre for all live trees ≥ 4.5 inch dbh by species and species group in Parcel N2,

northern unit, Massabesic Experimental Forest.

Species Number of Basal area Percent Volume

trees basal area

ft2

/acre ft3/acre

Softwoods

Eastern hemlock 881 31.3 20.1 380

Eastern white pine 170 52.6 33.9 1,338

Balsam fir 32 1.5 1.0 20

Red spruce 7 0.1 0.1 2

Red pine 2 0.9 0.5 22

Pitch pine <1 0.1 0.1 3

Subtotal 1,092 86.5 55.7 1,735

Hardwoods

Red maple 281 21.4 13.8 421

American beech 103 5.2 3.3 92

Sweet birch 78 1.9 1.2 9

Paper birch 75 6.8 4.4 82

Yellow birch 74 2.9 1.9 31

Northern red oak 62 21.8 14.0 695

Gray birch 34 1.6 1.0 6

White oak 14 3.9 2.5 112

Black cherry 7 0.3 0.2 3

White ash 4 1.0 0.6 25

Quaking aspen 4 0.7 0.4 13

Moose maple 4 0.2 0.1 —

Sugar maple 2 0.4 0.6 13

Black oak 2 0.2 0.1 3

Black gum 2 0.1 0.1 3

Hophornbeam 2 <0.1 <0.1 —

American elm 1 <0.1 <0.1 —

Serviceberry 1 <0.1 <0.1 —

Bigtooth aspen <1 0.1 0.1 3

Subtotal 751 68.7 44.3 1,511

Total 1,843 155.2 100.0 3,246
20

Figure 10.—Number of trees per acre by diameter classes and major tree species

in Parcel N2, northern unit, Massabesic Experimental Forest.

Table 5.—Number of trees and basal area/acre for all live trees ≥ 0.5 inch dbh and gross

volume/acre for all live trees ≥ 4.5 inch dbh by species and species group in Parcel N3,

northern unit, Massabesic Experimental Forest.

Species Number of Basal area Percent Volume

trees basal area

ft2

/acre ft3/acre

Softwoods

Eastern white pine 306 116.1 69.0 2,960

Red pine 83 37.6 22.4 904

Red spruce 23 4.1 2.4 82

Black spruce 5 <0.1 <0.1 —

Tamarack <1 0.4 0.2 11

Subtotal 417 158.2 94.1 3,957

Hardwoods

Gray birch 69 3.7 2.2 18

Red maple 43 2.7 1.6 28

Quaking aspen 14 2.3 1.4 38

Black cherry 6 0.9 0.5 19

Black gum 1 0.4 0.2 13

Subtotal 133 10.0 5.9 116

Total 550 168.2 100.0 4,073
21

Figure 11.—Number of trees per acre by diameter classes and major tree species

in Parcel N3, northern unit, Massabesic Experimental Forest.

Table 6.—Number of trees and basal/per acre for all live trees ≥ 0.5 inch dbh and gross

volume/acre for all live trees ≥ 4.5 inch dbh by species and species group in Parcel N4,

northern unit, Massabesic Experimental Forest.

Species Number of Basal area Percent Volume

trees basal area

ft2

/acre ft3/acre

Softwoods

Eastern hemlock 654 16.9 9.9 78

Eastern white pine 241 66.2 38.6 1,636

Balsam fir 155 5.2 3.0 —

Red pine 11 5.5 3.2 133

Red spruce 9 0.3 0.2 —

Subtotal 1,070 94.1 54.9 1,847

Hardwoods

Red maple 529 36.2 21.1 563

Paper birch 189 18.0 10.5 133

Yellow birch 82 2.6 1.5 —

American beech 76 5.0 2.9 33

Gray birch 56 5.3 3.1 74

Northern red oak 46 6.7 3.9 126

Quaking aspen 9 2.7 1.6 70

Black oak 2 0.9 0.5 26

Subtotal 989 77.4 45.1 1,025

Total 2,059 171.5 100.0 2,872
22

In the area sampled (107 acres) there are 915 live trees/

acre, basal area of 97.8 ft2

/acre, and volume of 1,887 ft3/

acre (Table 8). The parcel contains slightly more than a

quarter softwood species by basal area. White pine is the

major softwood species, in terms of number of trees and

basal area, and important hardwoods include red maple,

red oak, and paper birch. These four species account for

more than 85 percent of the basal area. Sixty-two percent

of the volume is red oak, red maple, and paper birch.

There is a mix of hardwoods and pine across the

diameter distribution (Fig. 14).

Parcel S3 — This is the largest parcel (1,363 acres) in the

MEF but only 593 acres were suitable for sampling

because of the presence of extensive wetlands. Parcel S3

is diverse in terms of topography, agricultural history,

wetlands, fire history, and effects of hurricanes. It has

had experimental plantings of softwoods and

hardwoods, softwood plantings to replace stands lost to

fire, and management interventions, such as thinning

and herbicide applications, to encourage natural pine.

Parcel S3 has 1,338 live trees/acre, basal area of 189.4

ft2

/acre, and volume of 4,610 ft3/acre (Table 9). Almost

three-fourths of the basal area is softwood species.

Important softwoods are white pine and hemlock, in

terms of number of trees and basal area, and important

hardwoods include red maple and red oak. These four

species account for more than 92 percent of the basal

area. More than half of the volume is white pine;

hemlock, red oak, and red maple account for most of the

remainder. The diameter distribution shows that most

species categories are represented over the range of

classes. Both red oak and white pine are present in the

1 - 4 inch diameter classes (Fig. 15).

Parcel S4 — This parcel was a natural pine forest that

partially burned in the 1947 fire. Although there were a

few small plantings of pine, the area was largely left to

regenerate naturally. In the area sampled (54 acres) there

are 1,220 live trees/acre, basal area of 130.1 ft2

/acre, and

volume of 2,666 ft3

/acre (Table 10). The parcel is half

softwoods and half hardwoods by basal area. White pine

and hemlock are the major softwood species, in terms of

number of trees and basal area, and important

hardwoods include red oak, red maple, and white oak.

These five species comprise nearly 90 percent of the

basal area and 88 percent of the volume. The diameter

distribution indicates that most species are represented

Figure 12.—Number of trees per acre by diameter classes and major tree

species in Parcel N4, northern unit, Massabesic Experimental Forest.
23

Figure 13.—Number of trees per acre by diameter classes and major tree

species in Parcel S1, southern unit, Massabesic Experimental Forest.

Table 7.—Number of trees and basal area/acre for all live trees ≥ 0.5 inch dbh and gross

volume/acre for all live trees ≥ 4.5 inch dbh by species and species group in Parcel S1,

southern unit, Massabesic Experimental Forest.

Species Number of Basal area Percent Volume

trees basal area

ft2

/acre ft3/acre

Softwoods

Eastern hemlock 2,388 66.6 37.2 228

Eastern white pine 115 45.5 25.4 1,270

Subtotal 2,503 112.1 62.6 1,498

Hardwoods

Red maple 350 24.7 13.8 417

Paper birch 174 15.9 8.9 140

Yellow birch 67 3.3 1.8 12

Northern red oak 50 18.5 10.3 512

American beech 38 1.0 0.6 —

Sweet birch 15 0.5 0.3 —

Gray birch 15 1.0 0.6 —

White oak 8 1.9 1.1 50

Subtotal 717 66.8 37.4 1,131

Total 3,220 178.9 100.0 2,629
24

Figure 14.—Number of trees per acre by diameter classes and major tree

species in Parcel S2, southern unit, Massabesic Experimental Forest.

Table 8.—Number of trees and basal area/acre for all live trees ≥ 0.5 inch dbh and gross

volume/acre for all live trees ≥ 4.5 inch dbh by species and species group in Parcel S2

southern unit, Massabesic Experimental Forest.

Species Number of Basal area Percent Volume

trees basal area

ft2

/acre ft3/acre

Softwoods

Eastern white pine 126 25.9 26.5 591

Red pine 11 0.5 0.5 9

Subtotal 137 26.4 27.0 600

Hardwoods

Red maple 311 31.9 32.6 551

Sweet birch 157 3.9 4.0 9

Gray birch 93 4.2 4.3 38

Paper birch 88 12.0 12.3 206

Yellow birch 64 2.4 2.5 —

Northern red oak 42 13.6 13.9 413

White oak 9 1.8 1.8 34

Quaking aspen 5 0.6 0.6 13

Black cherry 5 <0.1 <0.1 —

American elm 3 0.5 0.5 8

Bigtooth aspen 1 0.5 0.5 14

Subtotal 778 71.4 73.0 1,287

Total 915 97.8 100.0 1,887
25

over the range of diameter classes. Both red oak and

white pine are present in the 1 - 4 inch diameter classes

and red oak accounts for a large proportion of trees

greater than 5 inches (Fig. 16).

Parcel S5 — Most of Parcel S5 burned in the 1947 fire

and many of the surviving trees blew down in hurricanes

in 1950 and 1954. Some white pine and red pine were

planted, and hardwood competition was controlled by

mechanical and chemical treatments. Most of the parcel

was left to regenerate naturally. In the area sampled (25

acres) there are 2,856 live trees/acre, basal area of 169.0

ft2

/acre, and volume of 2,495 ft3/acre (Table 11). This

parcel contains more softwood (58 percent of basal area)

than hardwood (42 percent). Hemlock and white pine

are the major softwood species, in terms of number of

trees and basal area, and important hardwoods include

red maple, white oak, and red oak. These five species

account for nearly 92 percent of the basal area. Almost

93 percent of the volume is from white pine, red oak,

white oak, hemlock, and paper birch. The diameter

distribution shows that hemlock in diameter classes

smaller than 6 inches is very abundant (Fig. 17).

Parcel S6 — This parcel burned in 1947 and much of it

was planted following the fire, mostly with white pine,

red pine, and some jack pine. In the area sampled (36

acres) there are 699 live trees/acre, basal area of 160.5 ft2

/

Table 9.—Number of trees and basal area/acre for all live trees ≥ 0.5 inch dbh and gross

volume/acre for all live trees ≥ 4.5 inch dbh by species and species group in Parcel S3,

southern unit, Massabesic Experimental Forest.

Species Number of Basal area Percent Volume

trees basal area

ft2

/acre ft3/acre

Softwoods

Eastern hemlock 675 50.9 26.9 912

Eastern white pine 147 84.7 44.7 2,485

Balsam fir 16 0.4 <1 1

Red pine 3 3.4 1.8 97

Pitch pine <1 0.1 <1 3

Subtotal 841 50.2 73.7 3,498

Hardwoods

Red maple 282 21.4 11.3 438

Northern red oak 87 17.5 9.2 454

Yellow birch 41 1.3 0.7 13

Paper birch 32 4.0 2.1 65

White oak 11 1.9 1.0 53

Sugar maple 10 0.7 0.4 10

White ash 9 0.7 0.4 13

American beech 8 1.1 0.6 34

Moose maple 6 0.2 0.1 —

Sweet birch 3 0.6 0.3 18

Bigtooth aspen 2 <0.1 <0.1 —

Gray birch 2 <0.1 <0.1 —

Black gum 2 <0.1 <0.1 —

Black cherry 1 0.1 <0.1 3

Quaking aspen <1 0.3 0.1 8

Black ash <1 0.1 <0.1 3

Subtotal 497 49.9 26.3 1,112

Total 1,338 189.4 100.0 4,610
26

acre, and volume of 3,566 ft3

/acre (Table 12). More than

80 percent of the basal area is softwood. Important

softwoods include white pine and red pine (mostly

planted) and pitch pine (natural), in terms of number of

trees and basal area. The hardwood component (< 20

percent of basal area) is gray birch, red oak, and white

oak. More than 58 percent of the basal area and nearly

60 percent of the total volume is white pine. The

diameter distribution of the parcel is characteristic of an

even-aged planting (Fig. 18).

Large live trees

Trees at least 18 inches dbh (large trees) are of particular

interest because they often define the ecological character

of a forest. Their importance to wildlife is well

documented (DeGraaf and Yamasaki 2001, DeGraaf et

al. 1992, Morrison et al. 1992). Because of their size and

shape, large trees appeal to people aesthetically and

spiritually. The southern unit contains the largest

Atlantic white-cedar recorded in Maine. In 1999, it was

21.6 inches dbh, had a crown diameter of 19 ft, and was

63 ft tall.

We estimate that there are 14 live trees/acre that are 18

inches dbh or larger, or almost 27,000 on the area

sampled. Of those measured, 66 percent are white pine,

16 percent hemlock, 11 percent red oak, 4 percent red

maple, and 2 percent white oak (Table 13). The

remaining 1 percent includes a few each of red pine,

yellow birch, American beech, sugar maple, and paper

birch. Parcel S3 has more large trees than average for the

MEF, with 19.8 large, live white pines/acre. This

abundance could be due to light harvest activity over the

past century compared to the rest of the property, and

because the parcel includes a relatively large area that did

not burn in 1947. Northern red oak in this size class is

most abundant in Parcel N1, with 4.4 large, live trees/

acre.

Figure 15.—Number of trees per acre by diameter classes and major tree

species in Parcel S3, southern unit, Massabesic Experimental Forest.
27

Figure 16.—Number of trees per acre by diameter classes and major tree

species in Parcel S4, southern unit, Massabesic Experimental Forest.

Table 10.—Number of trees and basal area/acre for all live trees ≥ 0.5 inch dbh and gross

volume/acre for all live trees ≥ 4.5 inch dbh by species and species group in Parcel S4,

southern unit, Massabesic Experimental Forest.

Species Number of Basal area Percent Volume

trees basal area

ft2

/acre ft3/acre

Softwoods

Eastern hemlock 431 24.8 19.1 205

Eastern white pine 206 31.1 23.9 701

Pitch pine 10 6.5 5.0 187

Red pine 7 2.7 2.1 72

Subtotal 654 65.1 50.0 1,165

Hardwoods

Red maple 321 14.0 10.8 225

Northern red oak 124 34.0 26.1 902

White oak 50 12.5 9.6 326

Yellow birch 31 2.1 1.6 18

Paper birch 12 0.6 0.5 —

White ash 12 0.2 0.2 —

Gray birch 6 0.4 0.3 —

Black cherry 6 <0.1 <0.1 —

Quaking aspen 4 1.2 0.9 30

Subtotal 566 65.0 50.0 1,501

Total 1,220 130.1 100.0 2,666
28

Figure 17.—Number of trees per acre by diameter classes and major tree

species in Parcel S5, southern unit, Massabesic Experimental Forest.

Table 11.—Number of trees and basal area/acre for all live trees ≥ 0.5 inch dbh and gross

volume/acre for all live trees ≥ 4.5 inch dbh by species and species group in Parcel S5,

southern unit, Massabesic Experimental Forest.

Species Number of Basal area Percent Volume

trees basal area

ft2

/acre ft3/acre

Softwoods

Eastern hemlock 2,083 64.1 37.9 360

Eastern white pine 148 32.8 19.4 728

Balsam fir 25 0.7 0.4 —

Subtotal 2,256 97.6 57.7 1,088

Hardwoods

Red maple 272 14.0 8.3 159

White oak 131 21.8 12.9 419

Northern red oak 73 22.5 13.3 630

Paper birch 51 9.8 5.8 179

Yellow birch 50 1.6 1.0 —

Gray birch 13 0.4 0.2 —

Sweet birch 10 1.3 0.8 20

Subtotal 600 71.4 42.3 1,407

Total 2,856 169.0 100.0 2,495
29

Figure 18.—Number of trees per acre by diameter classes and major tree

species in Parcel S6, southern unit, Massabesic Experimental Forest.

Table 12.—Number of trees and basal area/acre for all live trees ≥ 0.5 inch dbh and gross

volume/acre for all live trees ≥ 4.5 inch dbh by species and species group in Parcel S6,

southern unit, Massabesic Experimental Forest.

Species Number of Basal area Percent Volume

trees basal area

ft2

/acre ft3/acre

Softwoods

Eastern white pine 403 93.9 58.6 2,124

Pitch pine 63 18.7 11.6 450

Red pine 50 16.4 10.2 351

Subtotal 516 129.0 80.4 2,925

Hardwoods

Gray birch 101 10.0 6.2 81

White oak 34 9.9 6.2 256

Northern red oak 31 10.7 6.7 290

Black cherry 10 0.7 <1 14

Red maple 7 0.2 <1 —

Subtotal 183 31.5 19.6 641

Total 699 160.5 100.0 3,566
30

Dead trees

Dead trees are important as snags, which are used by

wildlife, and as future downed coarse woody

material that will remain on the forest floor for many

years. They provide habitat, carbon and moisture

storage, and a site for regeneration of tree seedlings.

Inventory data for standing dead trees includes trees

4.5 inches dbh or greater, not just large trees. We

summarized these data by parcel, unit, species group,

and size class (Table 14). Parcels N3 and S6 were

largely replanted with softwood species after the

1947 fire, possibly explaining why we found no dead

hardwoods 4.5 inches dbh or larger. There are no

dead softwood trees 11.5 inches dbh or larger

recorded for S6 but N3 has dead softwood trees in

all size classes. The two largest compartments, N2

and S3, have dead trees in all three size-classes for

both softwoods and hardwoods. Both the northern

unit and southern unit have similar numbers of dead

trees/acre in all three size-classes for both softwood

and hardwood species-groups. The northern unit

averages a total of 15.3 dead trees/acre and the

southern unit 22.8 dead trees/acre.

Tree mortality, especially among smaller diameter

classes, can provide clues to likely future

composition of the overstory. For example, at a point

in time one species in a stand may have more stems/

acre than another, suggesting that without

disturbance it will eventually dominate the overstory.

However, if trees of the more abundant species die at

a faster rate, the less abundant species may eventually

dominate. These inventory results represent only one

point in time so the mortality rates among species

cannot yet be determined. Nevertheless, comparing

the percentage of dead trees by diameter class

provides insight into likely changes in species

composition across the forest. Compared to other

species, eastern hemlock has a relatively low

percentage of dead trees in the 1 - 8 inch diameter

classes (Table 15), suggesting that the proportion of

hemlock will increase in the MEF, if it is not infested

by hemlock woolly adelgid (Adelges tsugae Annand.).

White pine has the greatest percentage of dead trees

in both units, and in most diameter classes.

Consequently, without disturbance or management

that focuses on white pine, it could decline in

importance. Overall, the percentage of dead trees is

similar among species in the two units except for red

oak, which has about 10 percent more dead trees in

the southern unit than in the northern unit. Thus,

like white pine, the prevalence of red oak might also

decline, especially in the southern unit.

Table 13.—Number of live trees/acre ≥ 18 inches dbh by species, parcel, unit, and total forest.

Eastern Eastern Red Northern Red White Yellow Sugar American Paper white hemlock pine red oak maple oak birch maple beech birch

Parcel pineN1 4.6 0.5 0.0 4.4 2.2 0.0 0.0 0.0 0.0 0.0N2 5.1 1.0 0.1 2.4 0.5 0.4 0.1 0.1 0.1 <0.1N3 2.8 0.0 0.7 0.0 0.0 0.0 0.0 0.0 0.0 0.0N4 6.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0Northern unit 4.9 0.9 0.1 2.1 0.5 0.3 0.1 <0.1 <0.1 <0.1S1 9.9 0.3 0.0 0.3 0.0 0.0 0.0 0.0 0.0 0.0S2 1.4 0.0 0.0 0.5 0.0 0.0 0.0 0.0 0.0 0.0S3 19.8 6.0 0.2 0.9 0.7 0.2 0.0 <0.1 0.0 0.0S4 3.2 0.0 0.6 0.7 0.0 0.6 0.0 0.0 0.0 0.0S5 1.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0S6 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0Southern unit 14.6 4.2 0.2 0.8 0.5 0.2 0.0 <0.1 0.0 0.0Forest 9.2 2.3 0.1 1.5 0.5 0.3 <0.1 <0.1 <0.1 <0.1
31

Tree Seedlings

Regeneration is the basis of the future forest. Tree

seedlings greater than 1 ft tall and with dbh less than 0.5

inches are present on most but not all 0.01-acre plots.

Among parcels, S3 has the greatest number of seedlings/

acre, followed by S4, N1, and N2. Hemlock is the most

abundant seedling, especially in S3, with an average of

1,091 seedlings/acre (Table 16). This species is also

abundant in N2, S5, and N1. White pine seedlings are

most abundant in N1, followed by S3 and N2. Red

maple seedlings are most abundant in S4, followed by S2

and N1. Oaks, including red and white, are most

abundant in S6, followed by S5, S2, S1, N1, and S4.

Seedlings found in a high percentage of plots throughout

the MEF include red maple, paper birch, white pine,

trembling aspen (Populus tremuloides), black cherry,

white oak, red oak, and hemlock.

Table 14.—Number of dead trees/acre by parcel, unit, species group, and size class in the

Massabesic Experimental Forest.

Softwood Hardwood

Area

Parcel (acres) Smalla Mediumb Largec Smalla Mediumb Largec Total

N1 36 2.3 1.0 0.2 7.8 1.7 0.3 13.3

N2 881 8.1 0.5 0.3 5.2 1.6 0.2 15.9

N3 91 2.3 0.4 0.3 0.0 0.0 0.0 3.0

N4 35 19.1 0.0 0.5 12.9 0.0 0.0 32.5

Northern unit 1043 7.8 0.5 0.3 5.1 1.4 0.2 15.3

S1 37 0.0 2.5 0.4 15.7 1.3 0.0 19.9

S2 107 3.7 0.0 0.0 15.8 0.4 0.1 20.0

S3 593 12.2 0.9 0.5 6.6 0.5 0.1 20.8

S4 54 9.3 0.0 0.0 31.2 0.0 0.0 40.5

S5 25 13.7 0.0 0.0 62.0 0.0 0.0 75.7

S6 36 4.9 0.0 0.0 0.0 0.0 0.0 4.9

Southern unit 852 10.1 0.8 0.4 11.1 0.4 0.1 22.8

a

Small: 4.5 - 11.4 inches dbh

b

Medium: 11.5 - 17.4 inches dbh

c

Large: 17.5 inches dbh or larger

Table 15.—Trees in the sapling and pole size classes in the Massabesic Experimental Forest that were

dead when sampled, by unit and species.

Unit Species Diameter class (inches)

1 2 3 4 5 6 7 8 Overall

Percentage dead trees

North White pine 44.6 37.7 30.4 16.9 15.7 6.8 8.5 2.0 24.3

Red maple 29.5 10.6 5.9 0 0 0 0 3.4 15.6

Red oak 17.6 10.4 6.4 17.3 0 12.7 8.2 7.9 1.3

Hemlock 9.6 3.1 0.9 1.8 0 2.4 0 0 6.6

South White pine 23.8 26.4 27.0 19.0 20.5 8.4 13.0 6.2 21.8

Red oak 30.6 25.8 27.2 19.3 26.9 10.3 0 9.2 21.2

Red maple 27.1 23.2 11.5 4.4 3.3 0 0 0 18.8

Hemlock 16.5 5.8 1.7 2.4 4.4 2.6 5.7 18.3 10.8
32 Table 16.—Seedlings/acre, by parcel in the Massabesic Experimental Forest. Seedlings are defined as trees > 1 

Scientific name Common name Parcel

S1 S2 S3 S4 S5 S6 N1 

Abies balsamea balsam fir 130.6 5.9 12.5 Acer pensylvanicum striped maple 3.6 27.3 Acer rubrum red maple 133.3 235.3 121.6 641.2 75.0 93.3 236.4 Acer saccharum sugar maple 3.6 9.1Betula alleghaniensis yellow birch 29.4 12.6 11.8 Betula lenta sweet birch 17.6 0.9 Betula papyrifera paper birch 5.9 8.1 18.2 Betula populifolia gray birch 8.3 23.5 7.2 37.5 73.3 9.1 Betula sp. birch species 18.2Carpinus caroliniana American hornbeam 0.9Crataegus sp. hawthorn species 1.8 9.1 Fagus grandifolia American beech 17.1 127.3 Fraxinus americana white ash 13.5 70.6 9.1 Juniperus communis common juniper 0.9 

var. depressa

Nyssa sylvatica black tupelo 3.6 

Ostrya virginiana eastern hophornbeam Picea rubens red spruce 0.9 Pinus rigida pitch pine 0.9Pinus strobus eastern white pine 50.0 64.7 255.9 164.7 12.5 60.0 418.2 16Populus tremuloides quaking aspen 17.6 5.4 11.8 Prunus serotina black cherry 100.0 72.1 111.8 206.7 3Quercus alba white oak 183.3 23.5 36.9 317.6 12.5 186.7 81.8 6Quercus ilicifolia bear oak Quercus rubra northern red oak 158.3 182.4 97.3 147.1 212.5 293.3 154.5 8Tsuga canadensis eastern hemlock 91.7 1091.0 58.8 225.0 218.2 34Ulmus americana American elm Ulmus sp. elm species 17.6 Unknown dead unknown dead 1.8

hardwood hardwood

Total 625.0 717.6 1888.3 1541.2 587.5 913.3 1336.4 101
33

Understory Plants—Shrubs and Herbs

Density (stems/acre) of shrubs taller than 1 ft in 0.01-

acre plots is greatest in Parcel N3, followed by S1 and

N2 (Table 17). The most abundant shrub species in the

MEF is beaked hazelnut (Corylus cornuta), especially in

N3 and N2. The genus Viburnum, which includes here

witherod (V. nudum var. cassinoides), arrowwood (V.

dentatum), and maple-leaved viburnum (V. acerifolium),

is common and abundant across the forest, especially in

N3. Highbush blueberry (Vaccinium corymbosum) is

most abundant in S1 and S6. Winterberry (Ilex

verticillata), a facultative wetland species (Appendix II),

is abundant in N1 and S1. Other shrubs that average

more than 100 stems/acre on a single parcel include

witch-hazel (Hamamelis virginiana), sheep laurel (Kalmia

angustifolia), and speckled alder (Alnus incana ssp.

rugosa), an obligate wetland species (Appendix II). Parcel

N4 has no plots with shrubs taller than 1 ft, although

shrub species shorter than 1 ft were present and shrubs

were evident in the meander transects. We did not

collect shrub data for parcels S2 and S5.

A common shrub, low sweet blueberry (Vaccinium

angustifolium), is not represented in the stem count data

(Table 17) because of its growth habit, which is usually

shorter than 1 ft, yet it has a higher frequency than most

other understory species (Table 18; Figs. 19 and 20). We

found this species in 55 percent of the 211 plots in the

northern unit and in 63 percent of those in the southern

unit. The frequency of low sweet blueberry could be due

to sandy soils, fire history, and seed longevity in the soil.

Other common shrubs found in more than 10 percent

of plots for one or both units are beaked hazelnut,

maple-leaved viburnum, shadbush (Amelanchier sp.),

Northern Unit

10 20 30 40 50 60 70

  	

 

 

   	  	

 
	

  

   

   

  	 


 

  

   

 
	 

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# 

 	

 	

  

 $ 

Taxa

Percentage of 211 plots in which species are found

Figure 19.—Common understory plants in the northern unit, Massabesic Experimental Forest.
34

Table 17.—Shrub stems/acre found on fixed plots in eight parcels and by unit in the Massabesic Experimental are not available.Scientific Name Common Name N1 N2 N3 N4 NU S1 

Total 

Alnus incana ssp. rugosa speckled alder 7 5 

Alnus sp. alder species 18 0 1 

Amelanchier sp. shadbush species 18 13 27 14 

Carpinus caroliniana American hornbeam 0 0 

Cornus alternifolia alternate-leaved dogwood <1 <1 Corylus cornuta beaked hazlenut 360 568 344 

Crataegus sp. hawthorn species 36 16 14 

Diervilla lonicera bush-honeysuckle 9 0 <1 Gaylussacia baccata black huckleberry 0 0 

Hamamelis virginiana witch-hazel 132 105 Ilex verticillata winterberry 318 4 20 169 Juniperus communis v. depressa common juniper <1 <1 Kalmia angustifolia sheep laurel 0 0 185 

Lyonia ligustrina maleberry 2 2 169 

Nemopanthus mucronatus mountain holly <1 <1 77 Photinia melanocarpa black chokeberry 1 1 8 

Quercus ilicifolia bear oak 5 4 

Rhododendron canadense rhodora 12 9 

Spiraea latifolia meadowsweet 0 0

Vaccinium corymbosum highbush blueberry 23 18 354 Viburnum acerifolium maple-leaved viburnum 42 33 Viburnum lentago unknown seedling 1 1 

Viburnum nudum v. cassinoides witherod 2 477 52 

Viburnum dentatum arrowwood 1 1 

Total 622 1073 625 962 
35

witch-hazel, highbush blueberry, mountain holly

(Nemopanthus mucronatus), black huckleberry

(Gaylussacia baccata), winterberry, sheep laurel, and

witherod (Figs. 19 and 20).

One way to examine the frequency of understory plants

is by the number of parcels in which they are found. In

addition to low sweet blueberry, all 10 parcels have

highbush blueberry, wintergreen (Gaultheria

procumbens), mountain rice (Oryzopsis asperifolia),

bracken fern (Pteridium aquilinum) and club moss

(Lycopodium obscurum). Nine of ten parcels have wild

sarsaparilla (Aralia nudicaulis), partridgeberry (Mitchella

repens), swamp dewberry (Rubus hispidus), pink lady’s

slipper (Cypripedium acaule), Canada mayflower

(Maianthemum canadense), and cow-wheat (Melampyrum

lineare) (Figs. 19 and 20).

Another way to consider frequency is in the number of

plots where a species was found (Table 18). The three

most frequent understory species in each unit are the

same and in the same order—starflower (Trientalis

Southern Unit 

10 20 30 40 50 60 70

 




 


 

 



 


 


 



 

 



   

 
 

! 


" 

# 

 


$ 


#
% 


 


 


 


&
 

 
 

' 


  


(
 



 


 

 

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Taxa

Percentage of 188 plots in which species are found

Figure 20.—Common understory plants in the southern unit, Massabesic Experimental Forest.
36 Table 18.—Percentage of plots in which given species was found, by parcel.

No number indicates species was not found during survey.Number of plots/parcel are shown in parentheses beside parcel name.

Northern unit Southern 

N1 (11) N2 (167) N3 (22) N4 (11) Total (211) S1 (13) S2 (22) S3 (113) S4 (17) 

Coniferous treesAbies balsamea 18.2 27.5 4.5 63.6 26.5 16.8 17.6 Larix laracina 4.5 0.5Picea rubens 1.8 9.1 9.1 2.8 1.8 Pinus resinosa 1.2 36.4 9.1 5.2 4.5 3.5 11.8 Pinus rigida 0.6 0.5 5.9 Pinus strobus 81.8 80.2 95.5 90.9 82.5 61.5 72.7 92.9 100.0 Pinus sylvestris 0.6 0.5Tsuga canadensis 72.7 63.5 13.6 72.7 59.2 76.9 4.5 83.2 58.8 Deciduous treesAcer pensylvanicum 9.1 7.8 6.6 4.5 0.9 Acer rubrum 100.0 96.4 72.7 100.0 94.3 92.3 90.9 95.6 100.0 Acer saccharum 18.2 1.8 2.4 7.7 2.7 Betula alleghaniensis 63.6 24.0 9.1 22.7 38.5 22.7 20.4 5.9 Betula lenta 9.1 7.8 6.6 7.7 31.8 6.2 5.9 Betula papyrifera 18.2 34.1 9.1 45.5 31.3 46.2 68.2 25.7 11.8 Betula populifolia 9.1 17.4 36.4 36.4 19.9 7.7 22.7 1.8 5.9 Betula sp. 9.1 2.4 2.4 9.1 Carpinus caroliniana 0.0 0.9 Crataegus cf. macrosperma 9.1 0.5

Crataegus sp. 9.1 9.0 7.6 1.8 11.8 Fagus grandifolia 72.7 42.5 9.1 37.9 23.1 15.9 Fraxinus americana 27.3 7.8 7.6 8.0 5.9 Fraxinus nigra 0.6 0.5Fraxinus sp. 

Malus sylvestris 4.5 0.5Nyssa sylvatica 1.2 4.5 1.4 4.5 0.9 Ostrya virginiana 1.8 1.4Populus grandidentata 1.8 9.1 1.9 0.9 5.9 Populus tremuloides 4.2 22.7 18.2 6.6 9.1 1.8 5.9 Prunus serotina 28.7 63.6 45.5 31.8 50.0 29.2 29.4 Prunus virginiana 4.5 0.5 4.5 0.9 Prunus sp. 0.6 4.5 0.9
37

Northern unit Southern 

N1 (11) N2 (167) N3 (22) N4 (11) Total (211) S1 (13) S2 (22) S3 (113) S4 (17) 

Quercus alba 36.4 38.9 13.6 27.3 35.5 30.8 40.9 30.1 64.7 Quercus ilicifolia 1.2 0.9Quercus rubra 100.0 86.2 40.9 90.9 82.5 76.9 72.7 87.6 100.0 Quercus sp. 0.9 Quercus velutina 0.6 0.5Sorbus americana 1.2 0.9Ulmus americana 0.6 0.5Ulmus sp. 0.6 0.5 4.5 ShrubsAlnus incana ssp. rugosa 0.6 0.5 4.5 0.9 5.9 

Alnus cf. incana ssp. rugosa 4.5 

Alnus sp. 9.1 0.5Amelanchier sp. 18.2 22.8 36.4 22.7 22.7 10.6 52.9 Comptonia peregrina 4.8 22.7 6.2 13.6 0.9 Cornus alternifolia 1.2 0.9Cornus cf. racemosa 4.5 

Cornus sp. 1.2 0.9 Corylus cornuta 54.5 33.5 18.2 18.2 32.2 22.7 13.3 11.8 Diervilla lonicera 3.0 9.1 3.3 4.5 1.8 Gaylussacia baccata 1.2 4.5 1.4 7.7 4.5 9.7 41.2 Gaylussacia cf. baccata 4.5 

Hamamelis virginiana 81.8 26.3 9.1 25.6 4.5 33.6 29.4 Ilex verticillata 9.1 3.0 9.1 3.8 15.4 22.7 9.7 17.6 Juniperus communis var. depressa 0.6 0.5 0.9 5.9 

Kalmia angustifolia 9.1 6.0 4.5 5.7 30.8 27.3 38.1 64.7 Lonicera canadensis 1.2 4.5 1.4Lonicera sp. 0.9 

Lyonia ligustrina 2.4 1.9 15.4 9.1 8.0 11.8 Nemopanthus mucronatus 2.4 13.6 9.1 3.8 15.4 9.1 18.6 5.9 Photinia melanocarpa 1.8 18.2 3.3 7.7 4.4 17.6 Rhododendron canadense 0.6 0.5Rubus allegheniensis 9.1 0.9Rubus cf. canadensis 4.5 Rubus idaeus 0.6 0.5Rubus occidentalis 4.5 0.5 0.9 
38

Rubus sp. 1.2 18.2 2.8 2.7 Spiraea latifolia 9.1 2.4 18.2 0.0 4.3 36.4 5.3 Taxus canadensis 0.6 0.5Vaccinium angustifolium 81.8 51.5 72.7 54.5 55.5 38.5 77.3 56.6 88.2 Vaccinium corymbosum 18.2 2.4 4.5 18.2 4.3 23.1 45.5 19.5 17.6 Vaccinium myrtilliodes 9.1 1.4Viburnum acerifolium 27.3 29.9 25.1 4.5 10.6 Viburnum dentatum 9.1 2.4 2.4 4.5 0.9 5.9 Viburnum lantanoides 9.1 2.4 2.4Viburnum lentago 0.6 0.5Viburnum cf. lentago 1.2 0.9

Viburnum nudum var. cassinoides 54.5 14.4 13.6 15.6 15.4 27.3 15.0 29.4 Viburnum sp. 0.6 0.5SubshrubsAralia nudicaulis 63.6 56.9 4.5 36.4 50.7 38.5 45.5 45.1 17.6 Cornus canadensis 18.2 8.4 18.2 9.1 10.0 7.7 4.5 17.7 17.6 Epigaea repens 36.4 6.6 9.1 7.6 13.6 21.2 29.4 Gaultheria procumbens 81.8 38.3 18.2 18.2 37.4 23.1 45.5 50.4 70.6 Mitchella repens 63.6 18.6 13.6 9.1 19.9 7.7 13.6 44.2 17.6 Rubus hispidus 27.3 6.0 54.5 11.8 31.8 23.0 11.8 Rubus pubescens 4.8 3.8 13.6 4.4 5.9 HerbsAchillea millefolium 9.1 0.5Actaea pachypoda 0.6 0.5Anaphalis margaritacea 0.6 0.5 4.5 Anemone quinquefolia 0.6 4.5 0.9 5.9 Apocynum androsaemifolium 4.8 9.1 4.7 22.7 Apocynum sp. 0.6 18.2 2.4Arisaema triphyllum 3.6 2.8 13.6 0.9 5.9 Aster macrophyllus 1.8 9.1 1.9Aster sp. 9.1 2.4 13.6 9.1 4.3 22.7 1.8 Asteraceae sp. 7.2 13.6 7.1 0.9 5.9 Brassica sp. 4.5 

Circaea alpina 0.6 0.5 4.5 0.9 

Northern unit Southern 

N1 (11) N2 (167) N3 (22) N4 (11) Total (211) S1 (13) S2 (22) S3 (113) S4 (17) 

Table 18.—Continued.
39

Circaea cf. alpina 0.6 0.5

Clintonia borealis 9.1 7.2 9.1 6.6 7.7 5.3 Coptis trifolia 18.2 8.4 4.5 18.2 9.0 7.7 13.6 21.2 5.9 Cypripedium acaule 2.4 36.4 3.8 7.7 27.3 12.4 23.5 Desmodium sp. 0.6 0.5Eupatorium maculatum 4.5 

Fallopia cilinodis 4.5 Fragaria virginiana 0.6 0.5 4.5 Fragaria cf. vesca 4.5 0.5Galium asprellum 0.6 0.5Galium sp. 0.6 0.5 4.5 Goodyera tesselata 9.0 9.1 8.1 8.0 Hieracium sp. 1.8 1.4Houstonia caerulea 0.6 0.5 4.5 Hydrocotyle americana 1.2 0.9 4.5 Hypericum sp. 4.5 

Impatiens capensis 4.5 Iris versicolor 9.1 0.6 0.9 9.1 Lactuca sp. 0.6 0.5Lilium philadelphicum 4.5 Lycopus uniflorus 9.1 2.4 2.4 18.2 0.9 Lycopus sp. 4.5 

Lysimachia quadrifolia 10.2 4.5 8.5 50.0 3.5 11.8 Lysimachia terrestris 1.2 0.9 18.2 0.9 Maianthemum canadense 36.4 46.7 86.4 72.7 51.7 46.2 77.3 65.5 58.8 Maianthemum racemosum 9.1 1.8 1.9 0.9 11.8 Medeola virginiana 63.6 31.1 9.1 18.2 29.9 7.7 22.7 44.2 23.5 Melampyrum lineare 3.0 27.3 18.2 6.2 7.7 13.6 0.9 17.6 Mentha sp. 0.6 0.5Mitella nuda 0.6 0.5Moneses uniflora 0.9 5.9 Monotropa hypopithys 4.5 9.1 0.9 1.8 Monotropa uniflora 9.1 11.4 4.5 0.0 10.0 23.1 9.1 13.3 Oclemena acuminata 9.1 3.0 9.1 3.3 0.9 5.9 Oxalis montana 1.2 0.9

Northern unit Southern 

N1 (11) N2 (167) N3 (22) N4 (11) Total (211) S1 (13) S2 (22) S3 (113) S4 (17) 
40

Oxalis stricta 5.9 Oxalis cf. stricta 4.5 

Panax trifolius 9.1 0.5Persicaria arifolia 9.1 Persicaria sp. 4.5 

Platanthera clavellata 9.1 Polygala paucifolia 10.8 18.2 45.5 12.8 13.6 4.4 Polygonatum pubescens 2.4 18.2 2.8Polygonatum sp. 0.6 0.5Polygonum sp. 0.6 0.5Potentilla canadensis 0.9 Potentilla norvegica 4.5 Potentilla simplex 9.1 0.5 9.1 0.9 Potentilla cf. simplex 4.5 Prenanthes altissima 1.8 1.4Prenanthes sp. 18.2 6.0 4.5 18.2 7.1 9.1 Pyrola americana 2.4 4.5 9.1 2.8 4.5 3.5 Pyrola elliptica 0.6 4.5 0.9Pyrola sp. 0.6 0.5 0.9 Rorippa sp. 0.6 0.5Rubus dalibarda 18.2 4.8 4.7 8.8 Rumex acetosella 4.5 Scutellaria galericulata 0.6 0.5 9.1 Scutellaria sp. 0.6 0.5Solidago cf. puberula 0.6 0.5

Solidago cf. rugosa 4.5 

Solidago sp. 18.2 3.6 22.7 9.1 6.6 22.7 5.9 Streptopus sp. 0.6 0.5Symphyotrichum lateriflorum 9.1 0.5Symphyotrichum sp. 0.6 0.5Symplocarpus foetidus 3.0 2.4Thalictrum pubescens 0.9 Thalictrum sp. 1.8 1.4Triadenum virginicum 0.6 0.5 9.1 Trientalis borealis 63.6 58.1 72.7 72.7 60.7 38.5 90.9 68.1 70.6 

Northern unit Southern 

N1 (11) N2 (167) N3 (22) N4 (11) Total (211) S1 (13) S2 (22) S3 (113) S4 (17) 

Table 18.—Continued.
41

Northern unit Southern 

N1 (11) N2 (167) N3 (22) N4 (11) Total (211) S1 (13) S2 (22) S3 (113) S4 (17) 

Trillium sp. 0.9 Trillium undulatum 18.2 4.8 18.2 5.7 5.3 Uvularia sessilifolia 72.7 46.7 22.7 9.1 43.6 15.4 27.3 38.9 29.4 Veratrum viride 0.6 0.5Verbena hastata 9.1 Viola hastata 0.6 0.5Viola cf. adunca 4.5 0.5

Viola sp. 9.1 5.4 13.6 9.1 6.6 13.6 0.9 5.9 Unknown herb 4.5 0.0 Ferns and alliesAthyrium filix-femina 2.4 1.9 4.5 Botrychium virginianum 0.6 0.5Dennstaedtia punctilobula 18.2 2.4 2.8 9.1 9.7 Diphasiastrum complanatum 9.1 0.6 4.5 1.4Diphasiastrum cf. complanatum 4.5 

Diphasiastrum digitatum 1.2 4.5 1.4 7.7 9.7 Diphasiastrum sp. 0.9 

Dryopteris campyloptera 0.6 0.5Dryopteris carthusiana 0.6 0.5 4.5 4.4 Dryopteris cf. carthusiana 4.5 

Dryopteris cristata 1.2 0.9 9.1 Dryopteris intermedia 7.2 9.1 6.6 7.1 5.9 Dryopteris marginalis 4.5 Dryopteris sp. 9.1 1.2 4.5 1.9 4.5 0.9 Equisetum sylvaticum 0.6 0.5Equisetum sp. 0.6 0.5 4.5 0.9 Gymnocarpium dryopteris 0.9 Huperzia lucidula 2.4 1.9 0.9 Lycopodium annotinum 0.6 4.5 0.9Lycopodium clavatum 3.6 13.6 27.3 5.7 1.8 Lycopodium cf. dendroideum 9.1 

Lycopodium obscurum 9.1 17.4 22.7 36.4 18.5 30.8 9.1 35.4 52.9 Onoclea sensibilis 9.1 3.0 9.1 3.3 0.0 18.2 3.5 5.9 Osmunda cinnamomea 27.3 9.0 9.1 18.2 10.4 23.1 27.3 24.8 5.9 Osmunda claytoniana 9.1 7.2 6.2 7.7 0.0 7.1 
42

Osmunda regalis 18.2 2.4 4.5 3.3 7.7 13.6 2.7 Polypodium appalachiana X P. virginianum 4.5 Polystichum acrostichoides 0.6 0.5Pteridium aquilinum 54.5 44.9 81.8 18.2 47.9 23.1 63.6 38.9 76.5 Thelypteris novaboracensis 3.0 18.2 3.3 9.7 5.9 Thelypteris palustris 0.6 0.5 4.4 5.9 Thelypteris sp. 0.6 0.5Unknown fern 2.4 1.9 1.8 GrassesAgrostis gigantea 0.9 Agrostis perennans 4.5 1.8 Agrostis scabra 0.9 Agrostis cf. capillaris 0.6 0.5

Agrostis sp. 9.1 0.5Anthoxanthum odoratum 5.9 

Brachyelytrum septentrionale 18.2 4.8 4.5 9.1 5.7 1.8 Calamagrostis canadensis 0.6 0.5 4.5 Cinna latifolia 0.6 0.5Danthonia compressa 0.9 Danthonia spicata 9.1 2.4 4.5 2.8 13.6 1.8 Danthonia cf. spicata 4.5 0.5

Danthonia sp. 4.5 0.5Glyceria canadensis 4.5 

Glyceria striata 1.8 1.4 4.5 Glyceria cf. striata 0.6 0.5

Glyceria sp. 4.5 

Oryzopsis asperifolia 54.5 47.9 68.2 18.2 48.8 7.7 45.5 29.2 29.4 Panicum cf. oligosanthes var. scribnerianum 0.6 0.5

Panicum sp. 9.1 1.2 1.4 0.9 Unknown grass 1.2 4.5 9.1 1.9 13.6 1.8 5.9 SedgesCarex albicans 9.1 0.9Carex arctata 0.6 0.5 4.5 Carex brachyglossa 0.9 

Northern unit Southern 

N1 (11) N2 (167) N3 (22) N4 (11) Total (211) S1 (13) S2 (22) S3 (113) S4 (17) 

Table 18.—Continued.
43

Northern unit Southern 

N1 (11) N2 (167) N3 (22) N4 (11) Total (211) S1 (13) S2 (22) S3 (113) S4 (17) 

Carex brunnescens 0.6 0.5 9.1 Carex canescens 9.1 Carex communis 4.5 0.5 4.5 Carex cf. communis 9.1 0.5

Carex crawfordii 0.6 0.5 4.5 

Carex debilis 9.1 1.2 13.6 9.1 3.3 4.5 1.8 5.9 Carex cf. echinata 9.1 

Carex foenea 0.9 Carex folliculata 0.6 0.5 4.5 5.9 Carex gynandra 0.6 0.5 4.5 Carex cf. gynandra 0.6 0.5

Carex intumescens 1.8 1.4 5.9 Carex leptalea 4.5 

Carex cf. lucorum 45.5 25.7 31.8 18.2 48.2 23.1 77.3 35.4 76.5 

Carex cf. stricta 0.6 0.5 7.7 

Carex tenera 0.9 

Carex tonsa 4.5 0.5Carex cf. tonsa 4.5 0.5

Carex trisperma 0.9 

Carex umbellata 4.5 0.5Carex cf. utriculata 4.5 

Carex sp. 9.1 3.0 4.5 9.1 3.8 4.5 1.8 5.9 Juncus brevicaudatus 0.9 Juncus sp. 9.1 0.6 4.5 1.4VinesAmphicarpa bracteata 2.4 1.9 0.9 Parthenocissus quinquefolia 1.2 0.9 0.9 Smilax herbacea 0.6 0.5Smilax sp. 9.1 0.5Toxicodendron radicans 2.4 18.2 2.8 4.5 3.5 5.9 OtherUnknown seedling 9.1 0.6 0.9 4.5 5.9 

sp. means there are potentially numerous species that could not be conclusively determined.cf. means “compare to.”
44

borealis), low sweet blueberry, and Canada mayflower.

Across both units, common herbs are starflower in 63

percent of plots, Canada mayflower in 56 percent,

bracken fern in 48 percent, wild sarsaparilla in 45

percent, wintergreen in 43 percent, wild oats (Uvularia

sessifolia) in 40 percent, and mountain rice in 40 percent.

In the northern unit, 18 understory species are found in

more than 10 percent of the 211 plots and eight of these

were in at least 40 percent (Fig. 19). In the southern

unit, 29 understory species are found in at least 10

percent of the 188 plots, and six were in at least 40

percent (Fig. 20).

Possible reasons for the frequent occurrence of species

that are found in more than 10 percent of plots per unit

include: broad ecological niche; successful reproductive

strategy; long history in the area; pollinators and

dispersers are common; vigorous response to

disturbance; or some combination of these. None of the

common species are exclusive to one or the other unit.

There are, however, two common species in the northern

unit that are not among the most common in the

southern unit: maple-leaved viburnum and bird on the

wing (Polygala paucifolia). Nine of the most common

species in the southern unit are not among the most

common in the northern unit: sheep laurel, witch-hazel,

highbush blueberry, mountain holly, pink lady’s slipper,

goldthread (Coptis trifolia ssp. groenlandica), bunchberry

(Cornus canadensis), black huckleberry, Indian pipe

(Monotropa uniflora), and whorled loosestrife (Lysimachia

quadrifolia). These differences between the northern and

southern units, and between parcels within units, could

be due to: historic distribution of plant species; response

to disturbance, which is patchy and heterogeneous, and

includes agriculture, fire, harvest, and windthrow in an

array of intensities; varying soil moisture and topography;

dispersal capability; or some combination of these.

As expected, patchiness of herb species within the parcels

varies. Our sampling effort was similar across parcels of

differing sizes, yet some parcels, such as N4 and S6, have

individual herb species that occur in at least 50 percent

of plots, while in most of the other parcels these same

herb species occur in few plots (Table 18). The two

largest parcels, N2 and S3, have no species that occur in

more than 10 percent of all plots. Patchiness among the

herb communities in these parcels could have to do with

the heterogeneous distribution of a wide range of soil

types, or the abundance of small wetland inclusions

interspersed among uplands, or various disturbances and

land use histories, or some combination.

In addition to the common occurrence of pink lady’s

slipper, other orchids occur in the MEF in some

unexpected habitats. For example, we were surprised to

find that orchids in the genus Goodyera, the rattlesnake

plantains, are much more abundant in pine plantations

than in natural stands. We found two species (G.

pubescens and G. tesselata) and a possible hybrid. Neither

is considered particularly rare in Maine. These herbs

produce numerous dust-like, wind-dispersed seeds. The

association between Goodyera and plantations suggests

that these species are capable of persisting in or

colonizing after intense disturbance. They appear to

benefit from a relatively open understory that lacks dense

vegetation. A population of about 90 individuals of an

orchid that grows in sphagnum bogs, swamp pink

(Arethusa bulbosa), was found in a red maple swamp in

the southern unit.

Rare plants

We found numerous uncommon and rare plants, some

of which are listed by the state or federal governments

for protection (Table 19). There are two small

populations of a federally threatened and stateendangered orchid, small whorled pogonia (Isotria

medeoloides). One population is in mature pine-oak and

the other in mixed woods on a former agricultural field;

both are on gradual slopes. The species appears to be

associated with braided streams; i.e., an interlacing

network of channels where leaf litter is shifted during

heavy rains. Other rare plants in the MEF are state

recognized but not federally listed (Maine Department

of Conservation 1999). A single individual of the stateendangered spotted wintergreen (Chimaphila maculata)

was discovered in 1999 by a local botanist. It was found

under hemlock, an uncharacteristic habitat for the

species, and its flowers did not open the year it was

found. The MEF also has two small populations of a

state-endangered plant, upright bindweed (Calystegia

spithamaea). One occurs in a pine plantation, and a

much larger population of at least 100 stems is in an area

of rocky woods. No flowers or fruits were seen. A local

botanist found swamp saxifrage (Saxifraga pensylvanica),

a state-threatened plant. Atlantic white-cedar is listed as

“Special Concern” in Maine and is near the northern

limit of its range. The Maine Natural Areas Program

classifies the Atlantic white-cedar swamp-community as

state “Imperiled” and globally “Vulnerable” (Gawler and

Cutko in press). We found four additional species

designated as state “Special Concern” in the inventory:

Wiegand’s sedge (Carex wiegandii), a tall sedge of

wetlands; Bailey’s sedge (Carex baileyi), a sedge of

forested wetlands and swamps; long-beaked sedge (Carex

sprengelii), found at a cellar hole in mixed woods; and

smooth winterberry holly (Ilex laevigata), a shrub in the

Atlantic white-cedar swamp.
45

Plants found only on meander transects

Because our focus was on uplands, plants associated with

wetter soils were not as well represented in the inventory.

However, meander transects led observers through many

small wetlands and larger swamps and greatly enhanced

preparation of a more complete flora. Among plants

tallied on transects but not on plots, 94 are obligate

wetland species (Appendix II). Plants found only on

transects include three deciduous tree species, four

conifers, 20 shrubs, five subshrubs, 11 ferns or fern allies,

11 grasses, 37 sedges (28 in the genus Carex), and 77 other

herbs, including seven orchid species (Appendix I).

Applying GIS

Variable-radius plot, fixed plot, and transect data can be

related to landscape features such as soils, wetlands, and

topography. There are many potential applications for

GIS in research and management in the MEF. Maps can

be produced that show clusters of plots with particular

characteristics; e.g., high regeneration rate of selected

tree species, percentage sawtimber, inoperable areas, and

distance from a road, wetland, or stream. For example,

the distribution of plots with trees 18 inches dbh or

greater (Fig. 21) shows that in the northern unit larger

trees are in stands south of Cooks Brook and west of CC

Road, and also on the hill near the Thirteen Cellar

Holes. In the southern unit, large trees are mostly west

of Ida Jim Road. Examining the association of forest

cover types with soil types is another example. In Figure

22 the area of the MEF covered by pine and pine-oak

types where more than 50 percent of the trees are

classified as sawtimber-size (shown in orange) was

selected in the GIS. The proportion of soil types

underlying this area is shown on the pie chart (Fig. 22).

Soils supporting pine and pine-oak types with a high

proportion of sawtimber-size trees in the MEF are

mostly very stony, fine sandy loams.

Table 19.—State- and federal-listed rare plants found in the Massabesic Experimental Forest.

Species Common State Global State Federal

name ranka rank status status

Agrostis hiemalis ticklegrass (rare*)

Calystegia spithamaea Upright bindweed S1 G4G5 Threatened

Carex baileyi Bailey’s sedge S1? G4 Special Concern

Carex sprengelii Long-beaked sedge S2 G5? Special Concern

Carex wiegandii Weigand’s sedge S2 G3 Special Concern

Chamaecyparis thyoides Atlantic white-cedar S2 G4 Special Concern

Chimaphila maculata Spotted wintergreen S1 G5 Endangered

Ilex laevigata Smooth winterberry holly S2 G5 Special Concern

Isotria medeoloides Small whorled pogonia S2 G2G3 Endangered Threatened

Saxifraga pennsylvania Swamp saxifrage S2 G5 Threatened

a

Explanation of codes:

S1 = critically imperiled in Maine because of extreme rarity (< 5 occurrences or very few remaining individuals or

acres) or because some aspect of its biology makes it especially vulnerable to extirpation from the State of Maine

S1? = assumed S1 but more information needed

S2 = imperiled in Maine because of rarity (6-20 occurrences or few remaining individuals or acres) or because of

other factors making it vulnerable to further decline

G2= imperiled (typically 6-20 occurrences or 1000-3000 individuals)

G3 = vulnerable (rare; typically 21—100 occurrences or 3000-10,000 individuals)

G4 = apparently secure (uncommon but not rare; some cause for long-term concern; usually > 100 occurrences and

10,000 individuals)

G5 = secure (common, widespread and abundant)

G5? = thought to be secure but in need of verification

* = rare and proposed for official state status (Arthur D. Haines, 2002, personal communication.)
46







 

 

































































































































 





























 











  







































  





 

 









 



























 









 









 





































     





















































 































 













 

 







 



 

  

	











      



Figure 21.—Distribution of large trees (diameter breast height ≥ 18 inches) by permanent sample

points, Massabesic Experimental Forest.
47

Figure 22.—Locations of stands in the Massabesic Experimental Forest with > 50 percent pine

or pine—oak sawtimber, are shown in orange. The proportion of soil types underlying these

areas are depicted in the pie chart.

012 0.5 Miles

Northern Unit, areas with 

more than 50% sawtimber 

in pine and pine - oak type

Southern Unit , areas with 

more than 50% sawtimber 

in pine and pine - oak type



Ratio of soils supporting more than 

50% sawtimber in pine and pine - oak type

very stony, fine sandy loam

fine sandy loam

extremely stony, fine sandy loam

gravelly-loamy sand humic material

loamy sand

peat

unweathered bedrock

sand

very fine sandy loam
48

Summary and Conclusions

We inventoried the flora on approximately one-half of

the 3,700-acre MEF using a combination of VRP and

fixed-area plot sampling, and meander transects. We

concentrated on upland portions of the forest preparatory

to establishing manipulative experiments in those areas.

Half of the forest was not sampled and is made up of

wetlands and a 52-acre administrative site. We

established and sampled a grid of 399 permanent sample

points. The points determined where VRP sampling was

conducted and also defined centers of 0.01-acre circular

plots. We inventoried trees 4.5 inches dbh or greater

with VRP sampling, and quantified smaller trees and all

other vegetation in the fixed-area plots. Because of the

emphasis on uplands, and because a grid is not always

the most efficient way to sample plant populations that

may be clumped, meander transects between plots were

used to ensure a more complete listing of plants.

We found six forest-cover types in the uplands,

representing more than 2,500 acres. The most common

of these is the white pine type with more than 900 acres,

followed by pine-oak with more than 800 acres, and

pine-hemlock with slightly more than 700 acres. A type

that is rare in Maine, Atlantic white-cedar swamp, is

relatively abundant in the southern unit with a total of

160 acres.

We found an unexpected high density of trees 18 inches

dbh or greater in some parcels - up to 19.8 trees/acre for

white pine, perhaps due to minimal harvest and low

intensity or no fire in the recent past. Tree mortality

suggests that without natural disturbances or

management intervention, there will be a decrease in

white pine and red oak, and an increase in hemlock over

time. Hemlock is the most abundant tree seedling (taller

than 1 ft and less than 0.5 inches dbh), followed by

maples (especially red maple), oaks (northern red and

white), and white pine. Parcel S3 has the highest seedling

density, followed by Parcels S4 and N1. Density of shrub

stems taller than 1 ft is greatest for beaked hazelnut,

followed by viburnums, highbush blueberry, and

winterberry. Shrub stems/acre is highest in N3, followed

by S1 and N2. The most common understory plant is

starflower, found in more than 60 percent of plots in

each unit. Other common herbs and shrubs include low

sweet blueberry, Canada mayflower, bracken fern,

witherod, shadbush, pink lady’s slipper, and

partridgeberry. Listed rare plants include a federally

threatened orchid, small whorled pogonia, and eight

additional state-listed species.

The MEF ecosystems appear to be intact, given recent

disturbances, and are mostly of high ecological integrity

as evidenced by a noteworthy absence of invasive, exotic

plants. Mature and over-mature stands provide

important wildlife habitat in a matrix of successional

post-fire regeneration. Together the database of flora and

stand attributes, and the GIS with physical descriptions

and data summaries, provide the information needed to

devise, plan, and establish forest management

experiments in the MEF. The data also provide a baseline

against which to compare changes in forest structure and

composition resulting from those experiments or from

natural or anthropogenic disturbances.

Acknowledgments

This research and resulting publication is the product of

many people. We are grateful to Research Work Unit

NE-4155, especially Clayton Carl, Jim Sweeney, Bill

Leak, and Heather Smith. David Funk, Peter Garrett,

and Charles Swett (retired assistant station director,

project leader, and forestry technician, respectively)

provided background about the MEF.

We also thank our intrepid volunteers who helped us

establish the grid of permanent sample points, identified

and measured plants, and helped enter data. Volunteers

included: Lawrence Carl, Al Carlson, Michael Cormier,

Melissa Dow, Ray Drown, George and Ann Dugovic,

Barbara Grunden, Charles Kokernak, Richard and Judy

Logan, Vera Matthews, Nancy McReel, Joe Morrill,

Betsy Newcomer, Jack Parker, Patricia Smith, and Gerald

Sparkowich. Students from l’Ecole Nationale

d’Ingénieurs des Travaux Agricoles of Bordeaux, France,

came to us as volunteers as part of their educational

experience. They included: Magali Boyce, Valerie

Cussigh, Florence Delpy, Christine Fortin, Vincent

Herbretau, Thomas Larrieu, Caroline Lucas, Laure

Mirande, and David Pergaud.

We appreciate guidance from Jeff Heppinstall at the

University of Maine and Bill Frament of the USDA

Forest Service, Northeastern Area, State and Private

Forestry, who helped develop the GIS. We gratefully

acknowledge the Maine Forest Service, especially Dennis

Brennan, Claudette Désetelle, Steven Harding, and

Bruce Small, for providing logistic support. We also are

grateful to Alfred Conservation Commission members

William Bullard, Al and Barbara Carlson, Ray Drown,

Patricia Smith, Jack and Pat Parker, and Judy Vasalle for

their support, and the Small Woodland Owners

Association of Maine (SWOAM), especially Everett

Towle and Gordon Stuart.

We are grateful to members of the New England

Botanical Club (especially Dr. Lisa Standley), the Maine

Mycological Association (especially Dorothy Spaulding,
49

Dr. Samuel Ristich, and Gary Marshall), the Maine

Entomological Society (especially Dick Dearborn and

Dan Jennings), and the 27th Annual Andrews Foray

(mosses, liverworts and lichens) for conducting field

trips in the MEF in 1999-2002, for increasing our

knowledge of the biota in the MEF, and for

contributions to our understanding of inventory results.

Dr. James Hinds and Patricia Hinds provided a survey of

the lichens in various habitats, while Bruce Allen

identified numerous moss specimens from the MEF.

Arthur D. Haines provided confirmation on some of our

plant identifications and determinations for some

specimens. Thanks also to Bill Leak, Nancy Sferra, and

Gary Wade, who reviewed earlier drafts of this report.

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52 Appendix I. Checklist of vascular plants found during the inventory by parcel and by sampling technique.

“Other” includes parcel S7 and reconnaissance surveys.

Abies balsamea (L.) P. Mill. balsam fir X X X X X X X X X X X X X X X X X X X X Acer negundo L. box elder X Acer pensylvanicum L. striped maple X X X X X X X X X X X X X X Acer rubrum L. red maple X X X X X X X X X X X X X X X X X X X X X X Acer saccharum Marsh. sugar maple X X X X X X X X X X X X X X Achillea millefolium L.* common yarrow X X X X X X X Actaea pachypoda Ell. white baneberry X X X X X XActaea sp. baneberry species X X X Agrostis canina L. velvet bentgrass X 

Agrostis capillaris L.* Rhode Island bentgrass X X X X X X Agrostis gigantea Roth* redtop XX Agrostis hyemalis (Walt.) B.S.P. ticklegrass X 

Agrostis perennans (Walt.) Tuckerman autumn bentgrass X X X X X X X Agrostis scabra Willd. ticklegrass X Agrostis sp. bentgrass species X X X X X X X X X Alnus incana (L.) Moench ssp. speckled alder X X X X X X X X X X X X X 

rugosa (DuRoi) Clause

Alnus serrulata (Ait.) Willd. smooth alder 

Alnus sp. alder species X X X X X Ambrosia artemisiifolia L.* common ragweed X X Ambrosia sp.* ragweed species X XAmelanchier cf. obovalis shadbush X X X 

Amelanchier cf. stolonifera Wieg. thicket shadbush 

Amelanchier sp. shadbush species X X X X X X X X X X X X X X X X X X X X X X X X X Amphicarpaea bracteata (L.) Fern. hog-peanut X X X X X X X X Anaphalis margaritacea (L.) pearly everlasting X X X X X X X 

Benth. & Hook. f.*

Andromeda polifolia L. var. bog rosemary X 

glaucophylla (Link) DC.

Andropogon cf. gerardii Vitman big bluestem X X

Anemone canadensis L. Canada anemone X XAnemone quinquefolia L. wood anemone X X X X X X X X X X Antennaria howellii ssp. neodioica pussytoes 

Greene

Anthoxanthum odoratum L.* sweet vernal grass X

Apios americana Medik. common groundnut X X XApocynum androsaemifolium L. spreading dogbane X X X X X X X X X X X X X X Apocynum sp. dogbane species X X X X X X X X 

N1 N2 N3 N4 Total S1 S2 S3 S4 

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Aralia nudicaulis L. wild sarsaparilla X X X X X X X X X X X X X X X X X X X X X X X X X X Arethusa bulbosa L. dragon's mouth X 

Arisaema triphyllum (L.) Schott jack-in-the-pulpit X X X X X X X X X X X X X X X X X Asclepias exaltata L. tall milkweed 

Asclepias incarnata L. swamp milkweed X XAsclepias sp. milkweed species X X X X Aster macrophyllus L. big-leaved aster X X X X X X X X X X Aster sp. aster species X X X X X X X X X X X X X X X X X X X X Asteraceae sp. aster family X X X X X X X X X X X X X X X X X Athyrium filix-femina (L.) Roth ex lady fern X X X X X X X X X X X X 

Mertens var. angustifolium(Willd.) Lawson

Berberis thunbergii DC.* Japanese barberry X X X X X Betula alleghaniensis Britt. yellow birch X X X X X X X X X X X X X X X X X X X X X X Betula lenta L. sweet birch X X X X X X X X X X X X X X Betula papyrifera Marsh. paper birch X X X X X X X X X X X X X X X X X X X X X X X X X Betula populifolia Marsh. gray birch X X X X X X X X X X X X X X X X X X X X X X X X X Betula sp. birch species X X X X X X X X X Bidens connata Muhl. ex Willd. purple stem beggar X 

 ticks

Bidens frondosa L. devil's beggar ticks X X X Bidens vulgata Greene tall beggar ticks X 

Bidens sp. beggar ticks species X X X Botrychium virginianum (L.) Sw. rattlesnake fern X X X Botrychium matricariifolium (Doll) Mingan moonwort X X X

A. Braun ex Koch

Brachyelytrum septentrionale grass X X X X X X X X X X X X X X X X X X 

(Babel) G. Tucker

Brassica sp. mustard species X Bromus erectus Huds.* bromegrass XBromus sp. bromegrass X XBulbostylis capillaris (L.) sedge 

Kunthe ex C. B. Clarke

Calamagrostis canadensis (Michx.) bluejoint grass X X X X X X X X X X X X X X X X 

Beauv.

Calopogon tuberosus (L.) B. S. P. grass-pink 

Calystegia spithamaea (L.) Pursh upright bindweed X XCardamine pensylvanica Muhl. Pennsylvania X X X X 

ex Willd. bitter-cress

Cardamine sp. bitter cress, toothwort X XCarex albicans Willd. ex Spreng. white-tinged sedge X X X X X X 

N1 N2 N3 N4 Total S1 S2 S3 S4 

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Carex albicans var. emmonsii white-tinged sedge X X

(Dewey ex Torr.) J. Rettig

Carex aquatilis Whalenb. water sedge 

Carex arctata Boott northern clustered X X X X X X X X X X X X 

 sedge

Carex argyrantha Tuckerman sedge X X Carex atlantica Bailey eastern sedge X X X X Carex atlantica ssp. capillacea Bailey eastern sedge 

Carex baileyi Britt. Bailey's sedge 

Carex blanda Dewey woodland sedge XCarex brachyglossa Mackenzie yellow-fruited sedge X X X X X X X Carex brunnescens (Pers.) Poir. brownish sedge X X X X X X X X X X X X X X X Carex bullata Schkuhr ex Willd. button sedge X

Carex canescens L. silvery sedge X X X X X X X X X Carex communis Bailey fibrous-rooted sedge X X X X X X X X Carex cf. communis Bailey fibrous-rooted sedge X X

Carex comosa Boott bristly sedge X 

Carex crawfordii Fern. sedge X X X X X X X X X X Carex crinita Lam. drooping sedge 

Carex cumulata (Bailey) Fern. sedge X Carex debilis Michx. white-edged sedge X X X X X X X X X X X X X X X X X X X Carex deflexa Horem. northern sedge X Carex cf. disperma Dewey two-seeded sedge X Carex echinata Murr. star sedge X X X X 

Carex foenea Willd. sedge X Carex folliculata L. long sedge X X X X X X X X X X X X X X X X Carex gracillima Schwein. graceful sedge X 

Carex gynandra Schwein. nodding sedge X X X X X X X X X Carex houghtoniana Torr. ex Dewey Houghton's sedge 

Carex intumescens Rudge sedge X X X X X X X X X X X Carex lacustris Willd. lake bank sedge X X X X Carex laxiflora Lam. loose-flowered sedge XCarex leptalea Wahlenb. bristly-stalked sedge X X X X X X Carex leptonervia (Fern.) Fern. two-edged sedge X X X Carex lucorum Willd. ex Link sedge X X X 

Carex cf. lucorum Willd. ex Link sedge X X X X X X X X X X X X X X X X X X X X X X X X 

Carex lurida Wahlenb. sallow sedge X X X Carex magellanica ssp. irrigua bog sedge X X X X 

(Wahlenb.) Hulten

Carex merritt-fernaldii Mackenzie Fernald's sedge 

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Carex novae-angliae Schwein. New England sedge

Carex ormostachya Wieg. sedge X X XCarex pallescens L. sedge X XX X

Carex pensylvanica Lam. Pennsylvania sedge X X

Carex projecta Mackenzie necklace sedge X X X X X X Carex pseudocyperus L. cyperus-like sedge X X Carex scabrata Schwein. rough sedge X XCarex scoparia Schkuhr ex Willd. pointed broom sedge X X X X X X X 

Carex cf. scoparia Schkuhr ex Willd. pointed broom sedge X XCarex sprengelii Dewey ex Spreng. long-beaked sedge

Carex stipata Muhl. ex Willd. awl-fruited sedge X X X X X 

Carex stricta Lam. tussock sedge

Carex cf. stricta Lam. tussock sedge X X X X X X X X X X X 

Carex swanii (Fern.) Mackenzie sedge X X X XCarex tenera Dewey sedge XXXX X 

Carex tonsa (Fern.) Bickn. deep green sedge X X X X X Carex tonsa var. cf. rugosperma deep green sedge X X

(Mackenzie) W.J. Crins

Carex trisperma Dewey three-seeded sedge X X X X X X Carex trisperma (Dewey) var. trisperma sedge X Carex trisperma var. billingsii Knight three-seeded sedge X 

Carex umbellata Schkuhr ex Willd. umbel-like sedge X X

Carex utriculata Boot beaked sedge X 

Carex vesicaria L. inflated sedge X X X Carex virescens Muhl. ex Willd. ribbed sedge X X Carex wiegandii Mackenzie Wiegand's sedge X XCarex sp. (includes many sedge species X X X X X X X X X X X X X X X X X X X X X X 

possible species)

Carpinus caroliniana Walt. ssp. American hornbeam X X X X 

virginiana (Marsh.) Furlow

Carya ovata (P. Mill.) K. Koch shagbark hickory X X X X

Cephalanthus occidentalis L. button bush X X X X X X Celastrus orbiculata Thunb.* Asian bittersweet

Chamaecyparis thyoides (L.) B. S. P. Atlantic white-cedar X Chamaedaphne calyculata (L.) Moench leatherleaf X X X X X X Chelone glabra L. white turtlehead X X X X X Chimaphila maculata (L.) Pursh spotted wintergreen

Chimaphila umbellata (L.) W. Bart. ssp. pipsissewa X X X X X X X 

cisatlantica (Blake) Hulten

Chrysosplenium americanum Schwein. water carpet X X X X X X X 

ex Hook.

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Cicuta maculata L. water hemlock X X X Cinna cf. arundinacea L. common woodreed X X

Cinna latifolia (Trev. ex Goepp.) Griseb. drooping woodreed X X X X X Circaea alpina L. alpine enchanter's- X X X X X X X X X X 

 nightshade

Circaea lutetiana L. ssp. canadensis (L.) common enchanter's- X

Aschers. & Magnus nightshade

Circaea sp. enchanter's-nightshade X X X X X X X X X Clematis virginiana L. virgin's bower X X XClematis sp. clematis species X XClintonia borealis (Ait.) Raf. blue-bead lily X X X X X X X X X X X X X X X Comandra umbellata (L.) Nutt. bastard toadflaxComptonia peregrina (L.) Coult. sweetfern X X X X X X X X X X X X X X X X X X X X Coptis trifolia (L.) Salisb. goldthread X X X X X X X X X X X X X X X X X X X X X X X X Cornus alternifolia L. f. alternate-leaved X X X X X X X 

 dogwood

Cornus amomum P. Mill. ssp. amomum silky dogwood X X XCornus canadensis L. bunchberry X X X X X X X X X X X X X X X X X X X X X X X X X Cornus racemosa Lam. X

Cornus cf. racemosa Lam. northern swamp X X X X 

 dogwood

Cornus sp. dogwood species X X X X X X X Corylus cornuta Marsh. beaked hazlenut X X X X X X X X X X X X X X X X X X X X X X X X Crategus macrosperma Ashe hawthorn X X XCrategus cf. macrosperma Ashe hawthorn X X XCyperus lupulinus (Spreng.) Marcks sedge 

ssp. macilentus (Fern.) Marcks

Cypripedium acaule Ait. pink lady's slipper X X X X X X X X X X X X X X X X X X X X X X Danthonia compressa Austin ex Peck wild oatgrass X X X X Danthonia spicata (L.) Beauv. poverty oatgrass X X X X X X X X X X X X X X X 

ex Roemer & J. A.

Danthonia sp. oatgrass species X X X X X X X X X X X X X Daucus carota L.* Queen Anne's lace X 

Decodon verticillatus (L.) Ell. water-willow XDennstaedtia punctilobula (Michx.) hay-scented fern X X X X X X X X X X X X X X X X X X X 

T. Moore

Deparia acrostichoides (Sw.) M. Kato silvery spleenwort X X X X X 

Desmodium glutinosum cluster-leaf tick-trefoil X

(Muhl. ex Willd.) Wood

Desmodium sp. tick-trefoil species X X X X X XDiervilla lonicera P. Mill. bush-honeysuckle X X X X X X X X X X X X X X X 

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Digitaria ischaemum (Schreb.) Muhl.* smooth crabgrass X Digitaria sanguinalis (L.) Scop.* northern crabgrass 

Digitaria sp.* crabgrass species X XDiphasiastrum complanatum (L.) northern running-pine X X X X X X X X X X 

Holub

Diphasiastrum digitatum southern running-pine X X X X X X X X X X X X X X X 

(Dill. ex A. Braun) Holub

Diphasiastrum tristachyum blue ground-cedar X 

(Pursh) Holub

Diphasiastrum sp. running-pine species X X X X X X X X X X X Dirca palustris L. leatherwood X XDoellingeria umbellata (P. Mill.) Nees flat-topped white aster X X X Drosera intermedia Hayne spatulate-leaved X

 sundew

Drosera rotundifolia L. round-leaved sundew X Dryopteris campyloptera (Kunze) mountain wood fern X X X X

Clarkson

Dryopteris carthusiana (Vill.) spinulose wood fern X X X X X X X X X X X X X 

H. P. Fuchs

Dryopteris cristata (L.) Gray crested wood fern X X X X X X X X X X X X X X X X Dryopteris intermedia evergreen wood fern X X X X X X X X X X X X X X X 

(Muhl. ex Willd.) Gray

Dryopteris marginalis (L.) Gray marginal wood fern X X X X X X X X X Dryopteris sp. wood fern X X X X X X X X X X X X X X X X X X X Dulichium arundinaceum (L.) Britt. three-way sedge X X X X X X X X Echinochloa muricata (Beauv.) Fern. barnyard grass 

Elaeagnus angustifolia L.* Russian-olive Elymus repens (L.) Gould* witch grass X Elymus cf. trachycaulus (Link) Gould slender wheatgrass X 

ex Shinners

Epifagus virginiana (L.) W. Bart. beechdrops X X X X X Epigaea repens L. mayflower X X X X X X X X X X X X X X X X X X X X X X Epilobium ciliatum Raf. American willow-herb X X X Epilobium cf. coloratum Biehler eastern willow-herb X X X 

Epilobium leptophyllum Raf. American marsh 

 willow-herb

Epilobium sp. willow-herb X X X X Equisetum fluviatile L. river horsetail XEquisetum pratense Ehrh. meadow horsetail X XEquisetum sylvaticum L. woodland horsetail X X X X X X X X X X X Equisetum sp. horsetail species X X X X X X X X X 

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Erechtites hieraciifolia (L.) Raf. ex D.C. fireweed X

Erigeron strigosus Muhl. ex Willd. rough fleabane 

Erigeron sp. fleabane species Eriophorum virginicum L. tawny cotton-grass X 

Eriophorum sp. cotton-grass species X 

Eupatorium maculatum L. spotted joe-pye weed X X X X Eupatorium sp. joe-pye weed species X XEuthamia graminifolia (L.) Nutt. common flat-topped X X 

 goldenrod

Euthamia sp. goldenrod species X XFagus grandifolia Ehrh. American beech X X X X X X X X X X X X X X X X X X X X X Fallopia cilinodis (Michx.) Holub fringed bindweed X X Fallopia japonica (Houtt.) Decraene* Japanese bindweed XX 

Festuca rubra L.* red fescue X Festuca sp. sheep fescue species X XFragaria vesca L. woodland strawberry X XFragaria virginiana Duchesne wild strawberry X X X X X X X Fragaria sp. strawberry species X X X X X Frangula alnus P. Mill.* alder-leaved buckthorn XFraxinus americana L. white ash X X X X X X X X X X X X X X X Fraxinus nigra Marsh. black ash X X X X X X X X X X X X Galium aparine L. spring cleavers 

Galium asprellum Michx. rough bedstraw X X X X X X X Galium lanceolatum Torr. wild-licorice XGalium palustre L. marsh bedstraw X 

Galium tinctorium (L.) Scop. southern three-lobed X X 

 bedstraw

Galium trifidum L. northern three-lobed X X X X 

bedstraw

Galium triflorum Michx. sweet-scented bedstraw X X X X X X Galium sp. bedstraw/cleavers X X X X X X X X X X X 

 species

Gaultheria hispidula (L.) Muhl. creeping snowberry X X X 

ex Bigelow

Gaultheria procumbens L. wintergreen X X X X X X X X X X X X X X X X X X X X X X X X X X Gaylussacia baccata (Wangenh.) black huckleberry X X X X X X X X X X X X X X X X X X X X X X X X 

L. Koch

Gaylussacia dumosa (Andr.) TorFern. dwarf huckleberry X 

Gentiana linearis Froel. closed gentian XGlyceria borealis (Nash) Batchelder northern mannagrass X X X Glyceria canadensis (Michx.) Trin rattlesnake mannagrass X X 

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Glyceria grandis S. Wats. American mannagrass X Glyceria melicaria (Michx.) northeastern X

F. T. Hubbard mannagrass

Glyceria obtusa (Muhl.) Trin. coastal mannagrass X X Glyceria striata (Lam.) A. S. Hitchc. fowl mannagrass X X X X X X X X X X X Glyceria sp. mannagrass species X X X X X X X X X X Gnaphalium sp. cudweed species X 

Goodyera pubescens (Willd.) R. Br. . downy rattlesnake X X X X X X X X 

ex Ait. f plantain

Goodyera cf. repens (L.) R. Br. ex Ait. f. dwarf rattlesnake X X

 plantain

Goodyera tesselata Lodd. checkered rattlesnake X X X X X X X X X X X X X X 

 plantain

Goodyera tesselata x repens var. rattlesnake plantain X X

ophioides

Goodyera sp. rattlesnake plantain X X

 species

Gymnocarpium dryopteris (L.) Newman oak fern X X X Hamamelis virginiana L. witch-hazel X X X X X X X X X X X X X X X X X X X X X X X Hemerocallis sp.* day lily species X 

Hieracium caespitosum Dumort.* yellow king devil X 

Hieracium cf. piloselloides Vill.* glaucous king devil X X

Hieracium sp. hawkweed species X X X X X X X X Houstonia caerulea L. bluets X X X X X X X X X X X Huperzia lucidula (Michx.) Trevisan shining clubmoss X X X X X X X X X Huperzia/Lycopodium sp. clubmoss species X X X X X Hydrocotyle americana L. marsh pennyroyal X X X X X X X X X X X Hypericum boreale (Britt.) Bickn. St. Johnswort X 

Hypericum cf. boreale (Britt.) Bickn. St. Johnswort X X

Hypericum canadense L. Canada St. Johnswort X Hypericum ellipticum Hook. St. Johnswort XHypericum gentianoides (L.) B. S. P. orange-grass XHypericum perforatum L.* common St. Johnswort X 

Hypericum punctatum Lam. spotted St. Johnswort XX 

Hypericum sp. St. Johnswort species X X X X X XX 

Ilex laevigata (Pursh) Gray smooth winterberry X Ilex verticillata (L.) Gray winterberry X X X X X X X X X X X X X X X X X X X X X X Impatiens capensis Meerb. orange touch-me-not X X X X X X X X X Ionactis linariifolius (L.) Greene still aster X X X X X

Iris versicolor L. northern blue flag X X X X X X X X X X X X X X X X X Isotria medeoloides (Pursh) Raf. small whorled-pogonia X X X XJuncus brevicaudatus (Engelm.) Fern. short-tailed rush X X 

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Juncus effusus L. soft rush X XX 

Juncus filiformis L. thread rush X X Juncus greenei Oakes & Tuckerman Green's rush X X 

Juncus tenuis Willd. path rush X X X Juncus sp. rush species X X X X X X X X X Juniperus communis L. var. depressa common juniper X X X X X X X X X X X X X X X 

Pursh

Kalmia angustifolia L. sheep laurel X X X X X X X X X X X X X X X X X X X X X X X X X Kalmia polifolia Wangenh. bog laurel X 

Lactuca biennis (Moench) Fern. tall blue lettuce Lactuca canadensis L. tall lettuce 

Lactuca sp. lettuce species X X X X X X X X Lamiaceae sp. mint family X XLarix laricina (Du Roi) K. Koch larch X X X X X X Lechea tenuifolia Michx. slender pinweed X X Lechea sp. pinweed species X X X X Leersia oryzoides (L.) Sw. rice cutgrass X Lilium philadelphicum L. wood lily X X X X X X X X Linnaea borealis L. ssp. longifolia twinflower X X X 

(Torr.) Hulten

Lobelia cardinalis L. cardinal flower XLobelia inflata L. Indian-tobacco X X X X X Lobelia kalmii L. Kalm's lobelia Lonicera canadensis Bartr. ex Marsh. fly honeysuckle X X X X X X X X X X X X 

Lonicera morrowii Gray* Morrow's honeysuckle 

Lonicera sp.* honeysuckle species X X X Ludwigia sp. water purslane X XLuzula multiflora (Ehrh.) Lej. common woodrush X X XLuzula pallidula Kirsch.* woodrush Luzula sp. common woodrush X X X X X X X X X Lycopodium annotinum L. stuff clubmoss X X X X X X X X X X Lycopodium clavatum L. wolfclaw clubmoss X X X X X X X X X X X X X X X X X X Lycopodium dendroideum Michx. prickly tree clubmoss X X Lycopodium obscurum L. princess-pine X X X X X X X X X X X X X X X X X X X X X X X X X X Lycopodium sp. clubmoss species X X X X Lycopus americanus Muhl. ex W. Bart. American water- X X X 

 horehound

Lycopus uniflorus Michx. northern water- X X X X X X X X X X X X X X X X X X 

 horehound

Lycopus virginicus L. Virginia water- X X X 

 horehound

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Lycopus sp. water-horehound X X X X 

 species

Lyonia ligustrina (L.) DC. maleberry X X X X X X X X X X X X X X X X X X X X X X X Lysimachia ciliata L. fringed loosestrife X Lysimachia quadrifolia L. whorled loosestrife X X X X X X X X X X X X X X X X X X Lysimachia terrestris (L.) B.S.P. swamp candles X X X X X X X X X X X X X X X Lysimachia sp. loosestrife species X X X X Maianthemum canadense Desf. Canada mayflower X X X X X X X X X X X X X X X X X X X X X X X X X Maianthemum racemosum (L.) Link false spikenard X X X X X X X X X X X X X X Maianthemum trifolium (L.) Sloboda three-leaved false X X 

 Solomon's seal

Malus sylvestris P. Mill.* apple X X X Medeola virginiana L. indian cucumber root X X X X X X X X X X X X X X X X X X X X X X X X X X Melampyrum lineare Desr. cow wheat X X X X X X X X X X X X X X X X X X X X X X X Menyanthes trifoliata L. buckbean X 

Mimulus ringens L. Allegheny X

 monkeyflower

Mitchella repens L. partridgeberry X X X X X X X X X X X X X X X X X X X X X X X X X X Mitella nuda L. naked miterwort X X X XMoehringia lateriflora (L.) Fenzl grove sandwort XX 

Moneses uniflora (L.) Gray one-flowered shinleaf X XX Monotropa hypopithys L. pinesap X X X X X X X X X X X X X X X X X Monotropa uniflora L. Indian pipe X X X X X X X X X X X X X X X X X X X X X X Muhlenbergia racemosa (Michx.) muhly X

B. S. P.*

Nemopanthus mucronatus (L.) Loes. mountain holly X X X X X X X X X X X X X X X X X X X X X 

Nuphar variegata Dur. yellow water lilly X 

Nymphoides cordata (Ell.) Fern. little floating heart XNyssa sylvatica Marsh. black tupelo X X X X X X X X X X Oclemena acuminata (Michx.) Nesom whorled aster X X X X X X X X X X X X X X X X X Oenothera parviflora L. small-flowered 

 evening-primrose

Oenothera perennis L. little sundrops 

Onoclea sensibilis L. sensitive fern X X X X X X X X X X X X X X X X X X X X Oryzopsis asperifolia Michx. rough-leaved ricegrass X X X X X X X X X X X X X X X X X X X X X X X X Oryzopsis pungens (Torr. ex Spreng.) ricegrass X 

A. S. Hitch

Osmunda cinnamomea L. cinnamon fern X X X X X X X X X X X X X X X X X X X X X X X Osmunda claytoniana L. interrupted fern X X X X X X X X X X X X X X X X X X X X X Osmunda regalis L. var. spectabilis royal fern X X X X X X X X X X X X X X X X X X X X X X 

(Willd.) G

Osmunda sp. fern species X X

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Ostrya virginiana (P. Mill.) K. Koch eastern hophornbeam X X X XOxalis montana Raf. common wood-sorrel X X X X X XOxalis stricta L. common yellow X X X X 

 wood-sorrel

Oxalis sp. wood-sorrel X X X X Panax trifolius L. dwarf ginseng X X X X X X X X X X X X X Panicum acuminatum ssp. fasciculatum woolly panic grass X 

(Torr.) Freckmann & Lelong

Panicum acuminatum ssp. implicatum woolly panic grass X X X

(Schribn.) Freckmann & Lelong

Panicum acuminatum ssp. spretum wooly panic grass X X X X

(J.A. Schultes) Freckmann & Lelong

Panicum clandestinum L. deer tongue grass 

Panicum depauperatum Muhl. starved panic grass X X X Panicum dichotomiflorum L. fall panicum Panicum sp. panic grass species X X X X X X X X X X X X Parthenocissus quinquefolia (L.) Planch Virginia creeper X X X X X X X X X X X Parthenocissus cf. vitacea (Knerr) grape woodbine X 

A.S. Hitchc.

Parthenocissus sp. woodbine species X XPeltandra virginica (L.) Schott tuckahoe XPersicaria arifolia (L.) Haroldson halberd-leaved X X X X X X 

 tearthumb

Persicaria hydropiperoides water pepper X X

(Michx.) Small

Persicaria maculosa S. F. Gray* lady's thumb 

Persicaria cf. punctata (Ell.) Small dotted smartweed Persicaria sagittata (L.) H. Gross arrow-leaved X X X X X X 

 tearthumb

Persicaria sp. tearthumb species X X Phegopteris connectilis (Michx.) Watt long beech fern X X 

Phleum pratense L.* timothy 

Photinia x floribunda (Lindl.) black chokeberry X X X X X X X X X X X X X X X X X 

Robertson & Phipps

Photinia sp. chokeberry species X XPicea glauca (Moench) Voss white spruce X XPicea mariana (P. Mill.) B. S. P. black spruce X X X X X Picea rubens Sarg. red spruce X X X X X X X X X X X X X X X X Picea sp. spruce species X XPinus monticola Dougl. ex D. Don western white pine X 

Pinus resinosa Ait. red pine X X X X X X X X X X X X X X X X X X X X 

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Pinus rigida P. Mill. pitch pine X X X X X X X X X X Pinus strobus L. white pine X X X X X X X X X X X X X X X X X X X X X X X Pinus sylvestris L.* Scotch pine X X X X X Pinus sp. pine species X XPlantago major L.* common plantain X XPlatanthera clavellata (Michx.) Luer green woodland orchid X X X X X X X X X Platanthera grandiflora (Bigelow) Lindl. large purple-fringed X X X

 orchid

Platanthera cf. hyperborea (Lindl. var. leafy northern green X X X

huronensis (Nutt.) Luer orchid

Platanthera lacera (Michx.) G. Don ragged orchid X 

Platanthera sp. orchid species X X X X Poa compressa L.* Canada bluegrass X 

Poa palustris L. fowl meadowgrass X Poa pratensis L.* Kentucky bluegrass X X X X X Poa sp. bluegrass species 

Pogonia ophioglossoides (L.) Ker-Gawl. rose pogoniaPolygala paucifolia Willd. bird on the wing X X X X X X X X X X X X X X X X X X X Polygala sanguinea L. bitter milkwort X Polygonatum pubescens (Willd.) Pursh hairy Solomon's seal X X X X X X X X X X Polygonatum sp. Solomon's seal species X X X X X XPolygonella articulata (L.) Meisn. jointweed X Polypodium appalachianum Appalachian polypody X X X 

Haufler & Windham

Polypodium appalachiana x polypody hybrid X 

P. virginianum

Polypodium virginianum L. rock polypody X X X X X X X X X Polystichum acrostichoides Christmas fern X X X X X X X 

(Michx.) Schott

Pontederia cordata L. pickerelweed X Populus grandidentata Michx. big-toothed aspen X X X X X X X X X X X X X Populus tremuloides Michx. quaking aspen X X X X X X X X X X X X X X X X X X X X X Populus sp. aspen species X XPotentilla canadensis L. running cinquefoil X X Potentilla norvegica L. rough cinquefoil X Potentilla cf. recta L.* rough-fruited X 

 cinquefoil

Potentilla simplex Michx. old-field cinquefoil X X X X X X X X X X X Potentilla sp. cinquefoil species X X X X X X Prenanthes alba L. rattlesnake root X XPrenanthes altissima L. tall white-lettuce X X X X X X X X Prenanthes trifoliata (Cass.) Fern. gall of the earth 

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Prenanthes sp. white-lettuce species X X X X X X X X X X X X X X X X X X Prunella vulgaris L.* selfheal Prunus pensylvanica L. f. pin cherry X X X X X X Prunus serotina Ehrh. black cherry X X X X X X X X X X X X X X X X X X X X X X Prunus virginiana L. choke cherry X X X X X X X X X X X X Prunus sp. cherry species X X X X X XPteridium aquilinum (L.) Kuhn ex bracken X X X X X X X X X X X X X X X X X X X X X X X X X X X 

Decken var. latiusculum (Desv.)

Underwood ex Heller

Pyrola americana Sweet round-leaved pyrola X X X X X X X X X X X X Pyrola elliptica Nutt. shinleaf X X X X X X X X X X X X X X Pyrola sp. pyrola species X X X X X X X X X Quercus alba L. white oak X X X X X X X X X X X X X X X X X X X X X X X X X X Quercus ilicifolia Wangenh. bear oak X X X X X X X X Quercus rubra L. northern red oak X X X X X X X X X X X X X X X X X X X X X X X X X Quercus velutina Lam. black oak X X X X X X X XQuercus sp. oak species X X Rhamnus sp. buckthorn species X XRhododendron canadense (L.) Torr. rhodora X X X X X X X X X X X Rhododendron groenlandicum Labrador-tea X 

(Oeder) Kron & Judd

Rhynchospora alba (L.) Vahl. white beak-rush XRhynchospora capitellata (Michx.) Vahl beak-rush X Ribes hirtellum Michx. bristly gooseberry X XRibes sp. gooseberry/current X X

 species

Rorippa sp.* water-cress species X X X XRosa carolina L. pasture rose 

Rosa multiflora Thunb. ex Murr.* multiflora rose X 

Rosa sp. rose species X X X X Rubus allegheniensis Porter common blackberry X X X X X X X X Rubus cf. canadensis L. Canada blackberry X X 

Rubus dalibarda L. dewdrop X X X X X X X X X X X X X X X X Rubus cf. flagellaris Willd. northern dewberry X XRubus hispidus L. swamp dewberry X X X X X X X X X X X X X X X X X X X X X X X X Rubus idaeus L. red raspberry X X X X X X X X X X X Rubus idaeus ssp. strigosus red raspberry 

(Michx.) Focke*

Rubus occidentalis L. black raspberry X X X X X X X Rubus pubescens Raf. dwarf raspberry X X X X X X X X X X X X X X X X Rubus sp. raspberry species X X X X X X X X X X X X X X 

Appendix I.—continued.

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Rumex acetosella L.* sheep sorrel X X X X X X X Sagittaria latifolia Willd. common arrowhead X XSalix discolor Muhl. pussy willow XSalix humilis var. humilis Marsh prairie willow X

Salix sp. willow species X X X XSambucus canadensis L. common elder X X X Sambucus racemosa L. ssp. pubens red-berried elder X X X 

(Michx.) House

Sarracenia purpurea L. pitcher plant X X 

Schizachyrium scoparium (Michx.) Nash little bluestem X X X X X Scirpus cyperinus (L.) Kunth wool-grass 

Scirpus microcarpus J. & K. Presl X 

Scirpus pedicellatus Fern. bulrush X 

Scirpus sp. bulrush species X 

Scutellaria galericulata L. common skullcap X X X X X X X X X X X Scutellaria lateriflora L. mad-dog skullcap X X X X X X X Scutellaria sp. skullcap species X X X X X X Senecio sp. groundsel species X XSium suave Walt. water parsnip X X XSmilax herbacea L. carrion flower X X X X X X X Smilax rotundifolia L. common greenbrier X XSmilax sp. brier species X X X X X X XSolanum dulcamara L.* bittersweet nightshade X X X X Solidago bicolor L. silverrod X X 

Solidago juncea Ait. early goldenrod X X Solidago nemoralis Ait. gray goldenrod X XSolidago puberula Nutt. downy goldenrod X X X X X Solidago rugosa P. Mill. rough-stemmed X X X X X 

 goldenrod

Solidago uliginosa Nutt. northern bog X X 

 goldenrod

Solidago sp. goldenrod species X X X X X X X X X X X X X X X X X X X X Sorbus americana Marsh. American X X X X X 

 mountain-ash

Sorbus sp. mountain-ash species X XSparganium americanum Nutt. lesser bur reed X 

Sparganium sp. bur reed species X X X X X X Spiraea alba Du Roi var. latifolia meadowsweet X X X X X X X X X X X X X X X X X X 

(Ait) Dippel

Spiraea tomentosa L. hardhack X X X X X Streptopus amplexifolius (L.) DC. twisted stalk X XStreptopus lanceolatus (Ait.) Reveal rose twisted stalk X X

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Streptopus sp. twisted stalk species X X X X X Symphotrichum cordifolium common blue heart- X X X X X 

(L.) Nesom leaved aster

Symphyotrichum ericoides (L.) Nesom squarrose white aster 

Symphyotrichum lateriflorum (L.) calico aster X X X X X X 

A. & D. Love

Symphyotrichum novae-angliae (L.) New England aster X 

Nesom

Symphyotrichum cf. racemosum (Ell.) bristly aster X 

NesomSymphyotrichum undulatum (L.) clasping heart- X X X X X X 

Nesom leaved aster

Symphyotrichum sp. aster species X X 

Symplocarpus foetidus (L.) Salisb. skunk cabbage X X X X X X

ex Nutt.

Taxus canadensis Marsh. American yew X X X X X XThalictrum pubescens Pursh tall meadow-rue X X X X X X X X 

Thalictrum sp. meadow-rue X X X X X X X X Thelypteris noveboracensis (L.) Nieuwl. New York fern X X X X X X X X X X X X X X Thelypteris palustris Schott var. marsh ferns X X X X X X X X X X X X X 

pubescens (Lawson)

Thelypteris simulata (Davenport) Massachusetts fern X 

Nieuwl.

Thelypteris sp. fern X X X X X X X Tiarella cordifolia L. foam flower X X X X X Tilia americana L. basswood X XToxicodendron radicans (L.) Kuntze poison-ivy X X X X X X X X X X X X X X X X X Toxicodendron vernix (L.) Kuntze poison-sumac X Triadenum virginicum (L.) Raf. marsh St. Johnswort X X X X X X Triadenum sp. 

Trichostema dichotomum L. blue curls X X XTrientalis borealis Raf. star flower X X X X X X X X X X X X X X X X X X X X X X X X X Trifolium pratense L.* red clover X Trifolium repens L.* white clover X XTrifolium sp.* clover species X 

Trillium cernuum L. nodding trillium 

Trillium erectum L. purple trillium 

Trillium undulatum Willd. painted trillium X X X X X X X X X X X X X X X Trillium sp. trillium species X X X X Tsuga canadensis (L.) Carr. eastern hemlock X X X X X X X X X X X X X X X X X X X X X X X Ulmus americana L. American elm X X X X X X X X X X 

Appendix I.—continued.

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Uvularia sessilifolia L. wild-oats X X X X X X X X X X X X X X X X X X X X X X X X X Vaccinium angustifolium Ait. low sweet blueberry X X X X X X X X X X X X X X X X X X X X X X X X X Vaccinium corymbosum L. highbush blueberry X X X X X X X X X X X X X X X X X X X X X X X X X Vaccinium macrocarpon Ait. large cranberry X X Vaccinium myrtilloides Michx. velvet-leaf blueberry X X X X X X X X X X Vaccinium oxycoccos L. small cranberry X 

Vaccinium sp. blueberry/cranberry X X 

 species

Veratrum viride Ait. false hellebore X X X X X X X X Verbena hastata L. blue vervain X X X X X X Veronica scutellata L. marsh speedwell X 

Viburnum acerifolium L. maple-leaved X X X X X X X X X X X X X X X X X X 

 viburnum

Viburnum dentatum L. var. lucidum Ait. arrowwood X X X X X X X X X X X X X X X X X X Viburnum lantanoides Michx. hobblebush X X X X X X X X X X X X Viburnum lentago L. nannyberry X X XViburnum nudum L. var. cassinoides witherod X X X X X X X X X X X X X X X X X X X X X X X 

(L.) Torr. & Gray

Viburnum opulus L. var. highbush cranberry 

americanum Ait.

Viburnum sp. viburnum species X X X X X X X X Vicia cracca L.* cow vetch X 

Viola blanda Willd. sweet white violet XViola cf. adunca Sm. hook-spurred violet X XViola cf. affinis Le Conte violet X Viola cucullata Ait. blue marsh violet X X X X Viola hastata halberd-leaved violet X XViola macloskeyi Lloyd ssp. pallens wild white violet X

(Banks ex D.C.) M.S. Baker

Viola sagittata Ait. arrowhead violet X X XViola sagittata Ait. var. ovata arrowhead violet 

(Nutt.) Torr. & Gray

Viola sp. violet species X X X X X X X X X X X X X X X X X X X X X Vitis cf. labrusca fox grape X X X X X X X X X X X X X X Woodwardia virginica (L.) Sm. Virginia chain fern X X X X X X Unknown unknown X X X X X X X X X X X X X X X X X X X 

Specimens for most of the less common taxa are stored in Bradley, ME or are deposited at the Maine Herbarium, University of Maine, Orono.* indicates species not native to North America.sp. means there are potentially numerous species that could not be conclusively determined.cf. means "compare to."

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Appendix II.—Wetland species found in the MEF and their status.

Latin Name Common Name Wetland Statusa

Agrostis gigantea Roth redtop FACWAlnus serrulata (Ait.) Willd. smooth alder OBL

Andromeda polifolia L. var. glaucophylla (Link) DC. bog rosemary OBL

Anemone canadensis L. Canada anemone FACW

Apios americana Medik. common groundnut FACW

Arethusa bulbosa L. dragon’s mouth OBL

Arisaema triphyllum (L.) Schott jack-in-the-pulpit FACWAsclepias incarnata L. swamp milkweed OBL

Bidens connata Muhl. ex Willd. purple stem beggar ticks FACW+

Bidens frondosa L. devil’s beggar ticks FACW

Calamagrostis canadensis (Michx.) Beauv. bluejoint grass FACW+

Calopogon tuberosus (L.) B. S. P. grass-pink FACW+

Cardamine pensylvanica Muhl. ex Willd. Pennsylvania bitter-cress OBL

Carex aquatilis Whalenb. water sedge OBL

Carex atlantica Bailey eastern sedge FACW+

Carex atlantica ssp. capillacea Bailey eastern sedge OBL

Carex brunnescens (Pers.) Poir. brownish sedge FACW

Carex bullata Schkuhr ex Willd. button sedge OBL

Carex canescens L. silvery sedge OBL

Carex comosa Boott bristly sedge OBL

Carex crinita Lam. drooping sedge OBL

Carex echinata Murr. star sedge OBL

Carex folliculata L. long sedge OBL

Carex gynandra Schwein. nodding sedge OBL

Carex intumescens Rudge sedge FACW+

Carex lacustris Willd. lake bank sedge OBL

Carex leptalea Wahlenb. bristly-stalked sedge OBL

Carex leptonervia (Fern.) Fern. two-edged sedge FACW

Carex lurida Wahlenb. sallow sedge OBL

Carex magellanica ssp. irrigua (Wahlenb.) Hulten bog sedge OBL

Carex projecta Mackenzie necklace sedge FACW

Carex psuedocyperus L. cyperus-like sedge OBL

Carex scabrata Schwein. rough sedge OBL

Carex scoparia Schkuhr ex Willd. pointed broom sedge FACW

Carex stipata Muhl. ex Willd. awl-fruited sedge OBL

Carex stricta Lam. tussock sedge OBL

Carex trisperma Dewey three-seeded sedge OBL

Carex vesicaria L. inflated sedge OBL

Carex wiegandii Mackenzie Wiegand’s sedge OBL

Cephalanthus occidentalis L. button bush OBL

Chamaecyparis thyoides (L.) B. S. P. Atlantic white-cedar OBL

Chamaedaphne calyculata (L.) Moench leatherleaf OBL

Chelone glabra L. white turtlehead OBL

Chrysosplenium americanum Schwein. ex Hook. water carpet OBL

Cicuta maculata L. water hemlock OBL

Cinna latifolia (Trev. ex Goepp.) Griseb. drooping woodreed FACW

Circaea alpina L. alpine enchanter’s-nightshade FACW

Continued.
69

Coptis trifolia (L.) Salisb. goldthread FACW

Cornus amomum P. Mill. ssp. amomum silky dogwood FACW

Decodon verticillatus (L.) Ell. water-willow OBL

Drosera intermedia Hayne spatulate-leaved sundew OBL

Drosera rotundifolia L. round-leaved sundew OBL

Dryopteris cristata (L.) Gray crested wood fern FACW+

Dulichium arundinaceum (L.) Britt. three-way sedge OBL

Eleocharis acicularis (L.) Roemer & J. A. Schultes needle spikerush OBL

Epilobium leptophyllum Raf. American marsh willow-herb OBL

Equisetum fluviatile L. river horsetail OBL

Equisetum pratense Ehrh. meadow horsetail FACW

Equisetum sylvaticum L. woodland horsetail FACW

Eriophorum virginicum L. tawny cotton-grass OBL

Eupatorium maculatum L. spotted joe-pye weed FACW

Fraxinus nigra Marsh. black ash FACW

Galium asprellum Michx. rough bedstraw OBL

Galium palustre L. marsh bedstraw OBL

Galium tinctorium (L.) Scop. southern three-lobed bedstraw OBL

Galium trifidum L. northern three-lobed bedstraw FACW+

Gaultheria hispidula (L.) Muhl. ex Bigelow creeping snowberry FACW

Glyceria borealis (Nash) Batchelder northern mannagrass OBL

Glyceria canadensis (Michx.) Trin rattlesnake mannagrass OBL

Glyceris melicaria (Michx.) F. T. Hubbard northeastern mannagrass OBL

Glyceria obtusa (Muhl.) Trin. coastal mannagrass OBL

Glyceria striata (Lam.) A. S. Hitchc. fowl mannagrass OBL

Huperzia lucidula (Michx.) Trevisan shining clubmoss FACWHydrocotyle americana L. marsh pennyroyal OBL

Hypericum boreale (Britt.) Bickn. St. Johnswort OBL

Hypericum canadense L. Canada St. Johnswort FACW

Hypericum ellipticum Hook. St. Johnswort OBL

Ilex laevigata (Pursh) Gray smooth winterberry OBL

Ilex verticillata (L.) Gray winterberry FACW+

Impatiens capensis Meerb. orange touch-me-not FACW

Iris versicolor L. northern blue flag OBL

Juncus brevicaudatus (Engelm.) Fern. short-tailed rush OBL

Juncus effusus L. soft rush FACW+

Juncus filiformis L. thread rush FACW

Kalmia polifolia Wangenh. bog laurel OBL

Larix laricina (Du Roi) K. Koch larch FACW

Leersia oryzoides (L.) Sw. rice cutgrass OBL

Lobelia cardinalis L. cardinal flower FACW+

Lobelia kalmii L. Kalm’s lobelia OBL

Lycopus americanus Muhl. ex W. Bart. American water-horehound OBL

Lycopus uniflorus Michx. northern water-horehound OBL

Lycopus virginicus L. Virginia water-horehound OBL

Lyonia ligustrina (L.) DC. maleberry FACW

Lysimachia ciliata L. fringed loosestrife FACW

Latin Name Common Name Wetland Status

Appendix II.—continued.

Continued.
70

Lysimachia terrestris (L.) B.S.P. swamp candles OBL

Maianthemum trifolium (L.) Sloboda three-leaved false Solomon’s seal OBL

Menyanthes trifoliata L. buckbean OBL

Mimulus ringens L. Allegheny monkeyflower OBL

Mitella nuda L. naked miterwort FACWNemopanthus mucronatus (L.) Loes. mountain holly OBL

Nuphar variegata Dur. yellow water lilly OBL

Nymphoides cordata (Ell.) Fern. little floating heart OBL

Onoclea sensibilis L. sensitive fern FACW

Osmunda cinnamomea L. cinnamon fern FACW

Osmunda regalis L. var. spectabilis (Willd.) G royal fern OBL

Panicum dichotomiflorum L. fall panicum FACWPeltandra virginica (L.) Schott tuckahoe OBL

Persicaria arifolia (L.) Haroldson halberd-leaved tearthumb OBL

Persicaria hydropiperiodes (Michx.) Small water pepper OBL

Persicaria sagittata (L.) H. Gross arrow-leaved tearthumb OBL

Picea mariana (P. Mill.) B. S. P. black spruce FACWPlatanthera clavellata (Michx.) Luer green woodland orchid FACW+

Platanthera grandiflora (Bigelow) Lindl. large purple-fringed orchid FACW

Platanthera lacera (Michx.) G. Don ragged orchid FACW

Poa palustris L. fowl meadowgrass FACW

Pogonia ophioglossoides (L.) Ker-Gawl. rose pogonia OBL

Pontederia cordata L. pickerelweed OBL

Rhododendron canadense (L.) Torr. rhodora FACW

Rhododendron groenlandicum (Oeder) Kron & Judd Labrador-tea OBL

Rhynchospora alba (L.) Vahl. white beak-rush OBL

Rhynchospora capitellata (Michx.) Vahl beak-rush OBL

Rubus hispidus L. swamp dewberry FACW

Rubus pubescens Raf. dwarf raspberry FACW

Sagittaria latifolia Willd. common arrowhead OBL

Sambucus canadensis L. common elder FACW

Sarracenia purpurea L. pitcher plant OBL

Scirpus cyperinus (L.) Kunth wool-grass FACW+

Scirpus microcarpus J. & K. Presl bulrush OBL

Scirpus pedicellatus Fern. bulrush OBL

Scutellaria galericulata L. common skullcap OBL

Scutellaria lateriflora L. mad-dog skullcap FACW+

Sium suave Walt. water parsnip OBL

Solidago uliginosa Nutt. northern bog goldenrod OBL

Sparganium americanum Nutt. lesser bur reed OBL

Spiraea alba Du Roi var. latifolia (Ait) Dippel meadowsweet FACW+

Spiraea tomentosa L. hardhack FACWSymphyotrichum lateriflorum (L.) A. & D. Love calico aster FACWSymplocarpus foetidus (L.) Salisb. ex Nutt. skunk cabbage OBL

Thalictrum pubescens Pursh tall meadow-rue FACW+

Thelypteris palustris Schott var. pubescens (Lawson) marsh ferns FACW+

Thelypteris simulata (Davenport) Nieuwl. Massachusetts fern FACW

Latin Name Common Name Wetland Status

Appendix II.—continued.

Continued.
71

Toxicodendron vernix (L.) Kuntze poison-sumac OBL

Triadenum virginicum (L.) Raf. marsh St. Johnswort OBL

Ulmus americana L. American elm FACWVaccinium corymbosum L. highbush blueberry FACWVaccinium macrocarpon Ait. large cranberry OBL

Vaccinium oxycoccos L. small cranberry OBL

Veratrum viride Ait. false hellebore FACW+

Verbena hastata L. blue vervain FACW+

Veronica scutellata L. marsh speedwell OBL

Viburnum nudum L. var. cassinoides (L.) Torr. & Gray witherod FACW

Viburnum opulus L. var. americanum Ait. highbush cranberry FACW

Viola blanda Willd. sweet white violet FACW

Viola cucullata Ait. blue marsh violet FACW+

Viola macloskeyi Lloyd ssp. pallens (Banks ex D.C.) M.S. B wild white violet OBL

Viola sagittata Ait. arrowhead violet FACW

Viola sagittata Ait. var. ovata (Nutt.) Torr. & Gray arrowhead violet FACW

Woodwardia virginica (L.) Sm. Virginia chain fern OBL

a

OBL = obligate wetland, FACW = facultative wetland.

FACW + = more frequently found in wetlands, - = less frequently found in wetlands.

Latin Name Common Name Wetland Status

Appendix II.—continued.
Printed on Recycled Paper

Dibble, Alison C.; Rees, Catherine A.; Sendak, Paul E.; Brissette, John C. 2004.

Vegetation of forested uplands in the Massabesic Experimental Forest. Gen.

Tech. Rep. NE-320. Newtown Square, PA: U.S. Department of Agriculture, Forest

Service, Northeastern Research Station. 71 p.

A summary of an inventory of vascular plants in the 3,700-acre Massabesic

Experimental Forest in York County, Maine. We identified about 500 species and

subspecies. The most common overstory trees were eastern white pine, eastern

hemlock, northern red oak, and red maple. Hemlock was the most abundant tree

seedling. Shrub density was greatest for beaked hazlenut, followed by several

viburnum species, highbush blueberry, and winterberry. The most common herb was

starflower. We found nine listed rare plants, including a federally threatened orchid,

small whorled pogonia. A geographic information system was developed and includes

inventory data, forest type, soils, fire history, topography, roads, and streams.

Keywords: Inventory, sampling, geographic information system, timber volume,

shrubs and herbs, rare plants, regeneration, white pine, northern red oak, eastern

hemlock, red maple, Atlantic white-cedar
Headquarters of the Northeastern Research Station is in Newtown Square,

Pennsylvania. Field laboratories are maintained at:

Amherst, Massachusetts, in cooperation with the University of Massachusetts

Burlington, Vermont, in cooperation with the University of Vermont

Delaware, Ohio

Durham, New Hampshire, in cooperation with the University of New Hampshire

Hamden, Connecticut, in cooperation with Yale University

Morgantown, West Virginia, in cooperation with West Virginia University

Parsons, West Virginia

Princeton, West Virginia

Syracuse, New York, in cooperation with the State University of New York,

College of Environmental Sciences and Forestry at Syracuse University

Warren, Pennsylvania

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programs.) Persons with disabilities who require alternative means for communication of

program information (Braille, large print, audiotape, etc.) should contact the USDA's TARGET

Center at (202)720-2600 (voice and TDD).

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Whitten Building, 14th and Independence Avenue SW, Washington, DC 20250-9410, or call

(202)720-5964 (voice and TDD). USDA is an equal opportunity provider and employer.

“Caring for the Land and Serving People Thr ving People Through Research”