Holland et al. – Vegetation changes in temperate forest
Vegetation changes in temperate forested island communities
in Lake Winnipesaukee, New Hampshire, USA
MARJORIE M. HOLLAND
Department of Biology, University of Mississippi, University, MS 38677
e-mail: mholland@olemiss.edu
WILLIAM M. CLAPHAM
USDA-ARS, Appalachian Farming Systems Research Center, Beaver, WV 24957
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Holland et al. – Vegetation changes in temperate forest
ABSTRACT. We document changes in overstory species composition and foliar cover
during a twenty-three year sampling period, comparing woody species on three small islands in
Lake Winnipesaukee, New Hampshire, and determining changes in dominant plant species over
time. Floristic surveys began on these islands in 1901, and provide valuable information about
earlier vascular plant composition. The three islands, named Three Mile Island, Hawk’s Nest
Island, and Blueberry Island, were first sampled quantitatively in 1978, again in 1991, and most
recently in 2001. Data are compiled here for woody vascular plant species found in long-term
monitoring plots on the islands owned and/or managed by the Appalachian Mountain Club. On
Hawk’s Nest Pinus resinosa, Pinus strobus and Tsuga canadensis were the three most dominant
species. On Blueberry Island Acer rubrum, Betula populifolia and Tsuga canadensis were the
three most dominant woody species when sampling began. However, by 2001 Ilex verticillata,
Myrica gale, and Vaccinium corymbosum were the three most dominant. On Three Mile Island
Pinus strobus and Quercus rubra were the two most dominant species in 1978, but by 2001 Acer
pensylvanicum and Hamamelis virginiana were the dominants. Overall species richness within
the permanent plots increased, suggesting that natural succession has followed similar patterns on
all three islands.
Key words: Acer pensylvanicum, Hamamelis virginiana, Quercus rubra, Pinus strobus, Tsuga
canadensis,
island
plant
diversity,
vegetation
composition,
bootstrapping, forest succession, freshwater island vegetation
2
disturbance,
Holland et al. – Vegetation changes in temperate forest
INTRODUCTION
An understanding of human disturbance is increasingly recognized as critical for the
interpretation of modern landscapes, the evaluation of conservation objectives, and the
development of management approaches for natural areas (Egan and Howell 2001; Foster 2000;
Foster and Motzkin 1998; Hall et al. 2002; Marcucci 2000). Despite widespread alteration of the
New England landscape since European settlement, there have been few major shifts in species
distributions, but rather several notable changes in the relative abundance of tree taxa (Hall et al.
2002). Several comparisons of early settlement and modern data from across the northeastern
United States have documented decreases in Fagus spp., Tsuga spp., Picea spp., Ulmus spp.,
Carya spp., Pinus spp., and Castanea dentata (Glitzenstein et al. 1990; Nowacki and Abrams
1992; Russell 1981; Siccama 1971). Taxa that have increased in relative abundance across the
region include: Acer spp., Betula spp., Populus spp., Abies balsamea Mill., Quercus spp., Prunus
spp., and Fraxinus spp. (McIntosh 1972; Siccama 1971; Whitney 1990). The most dramatic
change has involved Acer spp. (primarily Acer rubrum), which increased from 7 to 27% in
Massachusetts from colonial to modern times (Hall et al. 2002). An examination of pre-settlement
tree surveys for all six New England states indicates that in the 47 towns surveyed in New
Hampshire, Fagus grandifolia represented 25.5%, Tsuga canadensis represented 14.7%, and
Pinus spp. represented 12.3% of tree species across the state (Cogbill et al. 2002).
In this study, we review the influence of environmental variation and land-use history on
forest composition and distribution in islands of central New Hampshire. We focused on the
vegetation of three of the 253 islands in Lake Winnipesaukee, New Hampshire: Blueberry,
Hawk’s Nest, and Three Mile Islands. These three are either owned or managed by the
Appalachian Mountain Club (AMC), which allowed the authors, both of whom are members of
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Holland et al. – Vegetation changes in temperate forest
the Club, easy access for sampling. All three islands are located in the northwestern arm of the
lake, and the last of these three received its name because it is located three miles from the town of
Center Harbor, NH.
Plot-based ecological studies build on the floristic surveys summarized in Holland and
Sorrie (1989). Based on earlier observations of the three islands (Maciejowski et al. 1981), we
predicted species richness would be highest on the largest island (Three Mile), lowest on the
smallest (Blueberry), and that species richness would increase over time (Briggs et al. 2008).
Secondly, we did not expect to see a great change in the importance of Tsuga canadensis and
Fagus grandifolia, which are currently threatened in other places by the non-native invasive
species, the hemlock woolly adelgid beetle (Orwig and Foster 2002; USDA 2005) and beech bark
disease (Gilman and Watson 1993; Houston 1997). Thirdly, we present results from plot based
ecological studies which quantify the frequency, distribution, and dominance of woody species
listed in Holland and Sorrie’s (1989) floristic survey of the vegetation of Three Mile Island.
In 1900 the Appalachian Mountain Club (AMC) established a permanent camp on Three
Mile Island (Rogovin 1983). By the early 1970’s the Camp had grown, and managers realized that
certain uses needed to be restricted, leading to the implementation of a land use plan (Clapham et
al. 2010). Adapted from Odum (1969)’s four types of environments needed by humans, the Camp
was divided into Protective, Productive, Urban, and Compromise zones (Holland and Sorrie
1989). Protective and Productive zones see relatively little human use, while Urban and
Compromise zones support the majority of human activity. By the 1970s five islands were
affiliated with the Camp: Blueberry, Hawk’s Nest, Nabby, Rock, and Three Mile. Since Three
Mile is the largest island, experiencing the greatest human impact and supporting structures which
require regular maintenance of electrical and water fixtures, vegetation sampling was begun there.
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Holland et al. – Vegetation changes in temperate forest
In July 1978, a pilot environmental assessment study at Three Mile Island Camp commenced with
three purposes: (1) to establish a qualitative and quantitative vegetative baseline to allow future
measurement of the floristic change and rate of change; (2) to document the recent history of the
ecosystem and the role of disturbance (logging, fire, windthrow, and hiking) in its maintenance
and development; and (3) to determine whether or not the land use plan “was working” by
providing managers with vegetation data to establish a foundation for critical natural resource
decisions (Maciejowski et al. 1981). During the winter of 1978-1979, it was agreed to sample
Hawk’s Nest and Blueberry Islands during summer 1979, however, Nabby and Rock Islands
would not be sampled due to the small size of these islands.
SITE DESCRIPTION
The shore of Lake Winnipesaukee is 154 meters above sea level in the foothills of the
White Mountains. The lake has a water surface area of 187 square kilometers, and the distance
around the lake is 463 kilometers (New Hampshire GRANIT 2009). The lake is fed by a number
of assorted springs, and has a forested shoreline and crystal clear water. The three islands studied
are erosional remnants left by the last glacier in the final retreat of the ice sheet between 10,000
and 15,000 years ago (Likens et al. 1967). The islands range in size from Blueberry (0.27 ha) to
Hawk’s Nest (0.41 ha) to Three Mile Island (17.4 ha).
The geology of the islands in Lake Winnipesaukee in the reach between Moultonboro
Neck and Meredith Neck (Bear, Pine, Black Cat, Three, Five, and Six Mile Islands) is quite
uniform. These islands are all comprised of outcrops of the Winnipesaukee Quartz Diorite, which
is generally described as a medium grained, gray quartz diorite (New Hampshire Highway
Department 1940). In the places in which it is found it is often seen in contact or in gradational
sections with granite, granodiorite and diorite. The bedrock core of Three Mile Island is covered
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Holland et al. – Vegetation changes in temperate forest
with a surface mantle of broken blocks of bedrock from the local area produced as a result of the
intense frost-prying action which came to bear on this area immediately following the departure of
the ice. The mantle also contains unevenly distributed pockets of glacial till, which in this case is
comprised largely of silt, sand and small stones.
Three Mile Island. Records from the 1890’s indicate that Three Mile Island (TMI) was
then largely covered with a second-growth forest of Betula spp. and Populus spp., with an
abundant understory of evergreen conifers (Holland and Sorrie 1989; Rogovin 1983). At the turn
of the 20th century the island was described as “picturesquely rocky,” supporting a heavy stand of
mixed timber with a richly diversified undergrowth of herbs and shrubs (Atkins 1972; Holland and
Sorrie 1989). In May 1901, as a member of the Three Mile Island advisory board, H. P. Kelsey
introduced 481 plants representing 60 predominantly woody species on Three Mile Island,
contributed from his plant nursery (Kelsey 1902). A list was prepared in August of 1903 of the
more important species of plants found on Three Mile Island (Kelsey 1903). By 1911 when Pease
published his list of the 265 plant species at Three Mile Island, 51 of Kelsey’s 60 introductions
persisted (Pease 1911; Table 1). By 1978, the island’s forests were dominated by Fagus
grandifolia, Quercus rubra, and Pinus strobus with Acer rubrum, Betula papyrifera, and Tsuga
canadensis also prominent (Maciejowski et al. 1981; Table 3). In a floristic survey in the 1980s,
Holland and Sorrie (1989) reported 243 plant species on the island (Table 1).
Hawks’s Nest Island. Hawk’s Nest Island (HNI) is 100 m east of TMI and currently has
no structures on it. HNI was privately owned by a family of AMC members through the mid1960s, but their house fell into disrepair, and when HNI management was assumed by the AMC in
the 1960s, the old house was dismantled, and management was assigned to the TMI Camp.
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Holland et al. – Vegetation changes in temperate forest
Currently HNI is managed as part of the Protective zone, so permission for campfires is limited by
the TMI Camp management. No botanical sampling had been conducted at HNI prior to 1979.
Blueberry Island. Blueberry (BI; also known as Minister Island) is 3.2 km northeast of
TMI and is privately owned by AMC members who live outside of New England, so BI is
managed by TMI staff. Blueberry Island is primarily used for picnicking, although the owners
sometimes spend time in a primitive cabin there during the summers. William I. Haven, along
with two college friends, purchased Blueberry on 22 December 1887 for $25.00 (Albert Gray, coowner, pers. comm. 1980). An etching of the BI cabin dated 1899 shows an open area
surrounding the structure with a few small conifers close by. Today Haven’s descendants in the
North, Anderson, and Gray families share ownership for BI. The northeast and southeast ends of
BI are managed by TMI staff as part of the Protective zone, while the central area near the 1899
cabin is managed as part of the Compromise zone. No botanical sampling had been conducted at
BI prior to 1979.
MATERIALS AND METHODS
Plots were established and quantitative sampling of the canopy, shrub, and herb strata of
the islands’ mixed deciduous-coniferous forest were initiated at TMI in the summer of 1978, and
extended to the two smaller islands in 1979 (Holland et al. 1983; Maciejowski et al. 1981). A
circle, often used as a sample unit in forestry studies (Mueller-Dombois and Ellenberg 1974) was
chosen for two reasons: it is easy to mark and relocate using only two coordinates after a period
of time (Maciejowski et al. 1981) and it minimizes edge effect (Chapman 1976). To ensure
random distribution a grid was drawn over a map of the three islands (Phillips 1959), each
intersection on the grid was numbered consecutively, and numbered slips of paper were randomly
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Holland et al. – Vegetation changes in temperate forest
drawn from a hat until 25 plot-centers had been identified. All 25 plot-centers have been
permanently marked with plastic stakes, and further studies during 1991 and 2001 utilized the
same plots. Each circle used to sample the overstory has a diameter of 34 m; 19 plots were
established on TMI (the largest island), and three each were located on BI and HNI (the two
smaller islands) in order to sample 12.55% of the overall islands’ area. Nomenclature followed
USDA, NRCS (2011). Taxonomic authorities are provided in Tables 3, 4, and 5.
Within each circle, woody species (both trees and shrubs) composition, numbers of
individuals of each species, and dominance were recorded. Total frequency of each species was
recorded as the number of plots in which a species was present out of the total number of plots on
each island (19 on TMI, 3 on HNI and 3 on BI). Importance values were calculated using the sum
of the relative frequency (frequency of species A / total frequency) × 100), relative dominance
(dominance of species A / total dominance) × 100, where dominance = rating of species A / sum
of ratings for all species), and relative density (density of species A / total density) × 100, where
total density = sum of the number of individuals of species A from all plots) of each species. All
of the trees in each circle were identified and rated as to whether they were Dominant,
Codominant, Intermediate, or Overtopped in relation to each other (Maciejowski et al. 1981;
Smith 1962, 1986). Dominant trees have crowns extending above the general level of crown
cover and receiving full light from above and partly from the sides (thus, received the highest rank
of #4). Co-dominant trees (ranked #3) had medium-sized crowns and received little light from the
sides. Intermediate trees (ranked #2) were shorter than those in the two preceding classes;
receiving a little direct light from above but none from the sides. Overtopped (formerly
“Suppressed” in Smith 1962) trees [ranked # 1; e.g. short trees, saplings, and tall shrubs] had
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Holland et al. – Vegetation changes in temperate forest
crowns entirely below the general level of the crown cover (Smith 1986). Species richness was
recorded as the number of species per plot sampled on each island.
The number of overstory species was summarized for each of the four land use zones each
year of measurement using bootstrap resampling (sampling with replacement) (Efron and
Tibshirani 1998). The resampling procedure involved pooling the data for each year and zone, for
example one data pool contained data from the protective plots assessed in 1978. Then N samples
were randomly selected with replacement from a given data pool where N equals the number of
circular plots within a given land use zone. The mean of the N samples was then calculated and
recorded and the procedure repeated for each measurement year and land use zone combination
through 10,000 iterations using @Risk software (Palisade, Inc., Ithaca, NY), within Excel
(Microsoft Inc., Redmond, WA). The frequency distributions of the 10,000 generated means are
presented as probability density plots and cumulative frequency curves. Changes in mean species
number between sample years were assessed using a non-parametric two-sample bootstrap
hypothesis testing procedure described by Efron and Tibshirani (1993). In this procedure the null
hypothesis, H0, is that the two distributions of interest (i.e. the mean number of species for years
1978 and 1991 for the Urban Zone) are equal. Output of the test is the probability that H0 is true.
RESULTS
Distributions of overstory species were modeled by bootstrapping data for each sampling
period. Figure 1 (a-d) displays frequency distributions and cumulative probabilities for the total
numbers of overstory species per plot for the four land uses for the three sampling periods. The
increase in total overstory species in the protective and compromise zones between 1978 and 1991
was significant (Table 2). No other significant differences in the total overstory species in any of
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Holland et al. – Vegetation changes in temperate forest
the other land-use zones over the periods 1978-1991 and 1991-2001 were significant (Clapham et
al. 2009).
The number of species of woody plants increased in the permanent plots between 1978 and
2001 (Tables 3, 4, and 5; Figure 1). For the twenty-three year sampling period, overstory species
richness increased on TMI and BI (Tables 3 and 5). Due to the marked increase in species richness
on Three Mile Island from 1978 to 1991, which was an increase of 18 species, we predicted that in
the 2001 sampling year the species richness would have increased again. However, species
richness on Three Mile Island decreased slightly between 1991 and 2001 (Table 3). In the late
1970’s, the forests on Three Mile Island were dominated by Fagus grandifolia, Quercus rubra,
and Pinus strobus (Maciejowski et al. 1981). Other prominent species included Acer rubrum, and
Tsuga canadensis (Table 3) and these five have continued to be important into the 21st century.
As of 2001, there was no evidence of any hemlock wooly adelgid outbreaks on the three
islands sampled. Other species of interest (Hawboldt 1944; Houston 1994; Siccama 1971)
included Fagus grandifolia, Betula alleghaniensis and Betula papyrifera (Tables 3, 4, and 5), and
Vaccinium corymbosum (Tables 4 and 5). While evidence of beech bark disease was observed on
Three Mile Island by 1981 (Holland photo collection), beech tree density had increased over the
sampling period (Table 3). As the island’s name suggests, blueberries (Vaccinium corymbosum)
were a prominent component of the Blueberry Island flora throughout the sampling period (Table
5). While Betula alleghaniensis has not been sampled in the permanent plots on Blueberry Island
(Table 5), it disappeared on Hawk’s Nest (Table 4), and on Three Mile Islands (Table 3). Betula
papyrifera thrived on all three islands in 2001 (Tables 3, 4, and 5).
On Three Mile Island, Betula papyrifera, Fagus grandifolia, Tsuga canadensis, Acer
rubrum, Pinus strobus, and Quercus rubra all decreased in importance values from the sampling
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Holland et al. – Vegetation changes in temperate forest
in 1978 to the sampling in 2001 (Table 3). In 1978 Pinus strobus and Quercus rubra exhibited
highest relative frequency (Table 3). Based on 1991 data, we expected Pinus strobus and Acer
rubrum to be the most frequent species in 2001, but A. rubrum, A. pensylvanicum, and Quercus
rubra were each found in all 19 plots, so were the three most frequent species in 2001 (Table 3).
Species with the highest dominance values in 2001 on Hawk’s Nest Island (Table 4) were
Pinus resinosa (3.33), Betula papyrifera (2.75), and Pinus strobus (2.72). On HNI, Tsuga
canadensis maintained an importance value over 15 throughout the sampling period, while Pinus
resinosa, Pinus strobus, and Quercus rubra all decreased in importance values from 1978 to 2001
(Table 4). In 2001 Amelanchier laevis, Pinus resinosa, Pinus strobus, Quercus rubra, and Tsuga
canadensis shared the highest relative frequency on HNI, while in 1978 Pinus strobus and
Quercus rubra exhibited highest relative frequency.
Throughout the sampling period on Blueberry, Hawk’s Nest, and Three Mile Islands, the
overall density recorded changed for individual species (Tables 3, 4, and 5). On Blueberry Island,
Vaccinium corymbosum, Ilex mucronata, and Ilex verticillata increased in density (Table 5). On
Hawk’s Nest, Tsuga canadensis showed the largest increase in density followed by Alnus incana
spp. rugosa, which showed the second largest increase in density (Table 4). On Three Mile Island,
Hamamelis virginiana and Acer pensylvanicum had the largest change in density over the three
sampling years (Table 3). On Blueberry Island, three dominant species in 1979 (Acer rubrum,
Pinus strobus, and Tsuga canadensis) all decreased in importance values by 2001, when Ilex
mucronata, Ilex verticillata, and Vaccinium corymbosum exhibited the highest importance values
(Table 5).
The greatest change in species richness is seen on Three Mile Island, which is most likely
due to movement into the plots of species previously established in other locations on the island.
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Holland et al. – Vegetation changes in temperate forest
Based on sampling in the 1970s, we expected to see Acer pensylvanicum, Acer rubrum, Fagus
grandifolia, Pinus strobus, Quercus rubra, and Tsuga canadensis to have high importance which
they did, but Hamamelis virginiana had the highest 2001 importance value (Table 3). Hamamelis
virginiana, Acer rubrum, and Acer pensylvanicum were expected to have the highest density in
2001, but Fagus grandifolia had more individuals than Acer rubrum (Table 3). Quercus rubra,
Quercus alba, and Betula alleghaniensis were expected to have the highest dominance (mean
rating) for Three Mile Island in 2001, but in 2001 the four most dominant species were Betula
lenta (3.5), Populus grandidentata (3.42), Quercus alba (4.00) and Quercus rubra (3.23)
according to the mean dominance ratings (Table 3). Hamamelis virginiana (972) and Acer
pensylvanicum (776) had the highest densities in 2001 on Three Mile Island, while Fagus
grandifolia (531) also had a high density (Table 3).
DISCUSSION
Overall, overstory species numbers per plot and their standard deviations increased from
the earliest sampling to the more recent on all three islands during the three sampling periods.
Frequency distributions (Figure 1) provide an excellent means to display information about
populations, mean variability and ‘shape.’ Since the data were bootstrapped, most distributions
show a well-developed normal distribution. The insets showing the cumulative probability show
clearly the differences among the distributions - their position relative to the x-axis, and the degree
of variability – the slope of the probability function can be observed directly. The increase in
overstory species richness may be attributed to a combination of natural succession (including
blowdowns and lightning strikes) and the implementation of a land-use management plan on all
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Holland et al. – Vegetation changes in temperate forest
three islands. Use of a similar land-use management plan for other islands could be helpful in
protecting rare species and habitats.
In a study of Timber Island, Bradley and Crow (2010) compared the flora of Three Mile
Island to that of three other Lake Winnipesaukee islands (Bear, Rattlesnake, and Timber). They
speculated that an overall higher than anticipated species richness on Three Mile Island (compared
to other island flora surveyed) may be related to the fact that the floristic study of TMI involved
multiple-year sampling. We believe that increased protection of the island through a land use
management plan which restricted heavy use of TMI to the southern end of the island (Clapham et
al. 2010) may have allowed species with varied requirements to survive in different habitats
spread throughout the island. Perhaps multiple-year sampling and protection have both played a
significant role.
Regardless of what factors may have influenced the high species diversity on Three Mile
Island, we can compare relative dominance and relative frequency of woody plants between
Timber and Three Mile Islands. At Timber Island, Acer rubrum was present in all plots found in
the Acer rubrum - Dulichium arundinaceum cover type (Bradley and Crow 2010), while Acer
rubrum was present in all 19 plots on Three Mile Island (Table 3). Tsuga canadensis was present
in 18 plots (95%) on Three Mile Island (Table 3), while it was found in 100% of the Tsuga
canadensis cover type plots on Timber Island (Bradley and Crow 2010). Pinus strobus also
occurred in 18 plots (95%) on Three Mile Island, while it was found in 94% of the Pinus strobus
cover type plots on Timber Island (Bradley and Crow 2010). Quercus rubra occurred in 19 plots
(100 %) on Three Mile Island, while it was found in 90% of the Pinus resinosa cover type plots on
Timber Island (Bradley and Crow 2010).
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Holland et al. – Vegetation changes in temperate forest
Application of the Simple Matching Index to the island floras of Three Mile Island and
Timber Island produced a value of 65.59% (Bradley and Crow 2010). This high similarity value
is striking given the difference in human use of the two islands. Timber Island (135 acres) is the
largest undeveloped island in Lake Winnipesaukee, while Three Mile Island (43 acres)
experiences regular human disturbance from June through September when an AMC Camp is in
session. It is possible, however, that this heavy human use has protected Three Mile Island from
deer grazing in recent years. In fact, one moose was sighted on TMI about a decade ago, but only
at the beginning of the Camp’s season. Deer have not been reported at TMI since the start of the
vegetation sampling in 1978, so impacts from large grazing herbivores have not been noted.
Five out of the 25 plots sampled on BI, HNI, and TMI are shoreline plots, bordered by the
Lake. It is in these plots that woody shrubs have proliferated, and the preponderance of
Gaylussacia baccata, Ilex mucronata, Ilex verticillata, and Vaccinium corymbosum have been
documented. Undoubtedly the easy access to sunlight has allowed these shrubs to dominate the
shorelines of the three islands. By the mid-1980s, the number of non-indigenous species
introduced to TMI by Kelsey had dropped to 10 species (Holland and Sorrie 1989), and most of
these are growing in the island’s center in “Rhododendron Swamp,” part of the Protective Zone.
Cogbill et al. (2002) reviewed pre-settlement tree surveys to determine the natural patterns
of vegetation throughout historical New England. The pre-settlement northern hardwood forests
dominated by Fagus grandifolia were distinct from the Quercus spp. forests in southern New
England (Cogbill et al. 2002). Since the earliest records for BI, HNI, and TMI date from the
1890s, we assume that the trees noted in pre-settlement New Hampshire may have provided the
seed source for these islands. Seeds of Acer spp., Betula spp., and conifers are wind dispersed,
thus with the mainland less than 0.5 miles away from the islands, the mainland could have
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Holland et al. – Vegetation changes in temperate forest
provided the seed source. Specific changes on the three Lake Winnipesauke Islands sampled can
be attributed to natural phenomena such as lighting strikes and other storm events, along with
human uses associated with summer recreation (primarily in the Compromise and Urban zones). .
Whereas records from the 1890’s (Rogovin 1983) indicate that Three Mile Island was covered by
Betula spp. and Populus spp., by the late 1900’s the most frequently encountered species were
Acer rubrum, Fagus grandifolia, Pinus strobus, Quercus rubra, and Tsuga canadensis.
The sampling of the three islands on Lake Winnipesaukee is part of ongoing floristic and
ecological studies (Clapham et al. 2010; Hartmann 1941; Holland and Sorrie 1989; Kelsey 1902,
1903; Maciejowski et al. 1981; Pease 1911). As of this writing, healthy populations of Tsuga
Canadensis were found on all three islands. In spite of the infestation of beech bark disease on
Three Mile Island, density of Fagus grandifolia has increased over the study period; beech has not
been found on Hawk’s Nest, but the one lone beech on Blueberry Island continues to survive.
Numbers of “overtopped” woody specimens (shrubs and short trees) grew throughout the plots on
all three islands. On Blueberry Island the density of Nemopanthus mucronatus, Vaccinium
corymbosum, and Ilex verticillata increased each sampling year. On Hawk’s Nest Island, the
species richness remained relatively constant, but density of Alnus incana ssp. rugosa, Lyonia
ligustrina, and Viburnum cassinoides increased over the twenty-three year sampling period. On
Three Mile Island, the density of Acer pensylvanicum and Hamamelis virginiana increased
consistently from one sampling period to the next. We predict when the plots are next sampled the
shorter woody species (both trees and shrubs) currently exhibiting high importance values on the
islands’ edges will continue their importance there.
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Holland et al. – Vegetation changes in temperate forest
ACKNOWLEDGMENTS.
The authors thank Marin Andrus, Raymond W. Prach Sr.,
Hannah Holland, and Raymond W. Prach Jr. for assistance with 2001 field work; James Fedders
for help with statistical analyses; and C. John Burk, Elizabeth Farnsworth, Louis Palmer, Jr., Jay
Maciejowski, Bruce Sorrie, Jason Hoeksema and two anonymous reviewers for reviews of various
drafts of the manuscript. Logistical and moral support is acknowledged from Three Mile Island
Camp managers, committee members, crew, and campers. Financial support from the Three Mile
Island Camp Committee, the College of New Rochelle, and the University of Mississippi is
gratefully acknowledged. Help from Forrest Briggs, Dustin Culver, Tara Davis, Shannon Moore,
and Cherrelle Williams with manuscript preparation is greatly appreciated.
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and land-use history on upland forests of the Cary Arboretum, Hudson Valley, New York.
Bulletin of the Torrey Botanical Club 117: 106–122.
Gray, A.L. 1980, Personal Communication.
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Hall, B., G. Motzkin, D. Foster, M. Syfert, and S. J. Burk. 2002. Three Hundred Years of Forest
and Land-Use Change in Massachusetts, USA. Journal of Biogeography 29: 1319-1335.
Hartmann, E. 1941. The flora of Three Mile Island. Appalachia 23: 560-563.
Hawboldt, L.S. 1944. History of spread of beech scale, Cryptococcus fagi (Baerensprung), an
insect introduced to the Maritime provinces. Acadian Naturalist 1: 137-146.
Holland, M.M., W.M. Clapham, and J.D. Maciejowski. 1983. Volunteer success in natural
resource management at an Appalachian Mountain Club Camp in New Hampshire, pp.162170. In: Proceedings of Volunteers in the Backcountry Conference, U.S. Forest Service,
Durham, NH.
Holland, M.M., and B.A. Sorrie. 1989. Floristic dynamics of a small island complex in Lake
Winnipesaukee, New Hampshire. Rhodora 91: 315-338.
Houston, D.R. 1994. Major new tree epidemics: beech bark disease. Ann. Rev. Phytopathol. 32,
75-87.
Houston, D.R. 1997. Beech Bark Disease. Exotic pests of eastern forests, Conference Proceedings.
USDA Forest Services website. www.invasive.org/symposium/houston.html. Accessed 13
Nov. 2011.
Kelsey, H.P. 1902. Reports of the Councillors for the autumn of 1901: natural history.
Appalachia 10: 75-59.
______. 1903. Reports of the Councillors for the autumn of 1902: natural history. Appalachia
10: 202-203.
Lake Winnipesaukee website. Available at www.winnipesaukee.com. Accessed 15 Sept. 2011.
Likens, G.E., F.H. Bormann, N.M. Johnson, and R.S. Pierce. 1967. The calcium, magnesium,
potassium and sodium budgets for a small Forested ecosystem. Ecology 48(5): 772-785.
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Holland et al. – Vegetation changes in temperate forest
Maciejowski, J., W. Clapham, and M.M. Holland. 1981. Environmental assessment begins at
Three Mile Island Camp. Appalachia 43:137-142.
Marcucci, D.J. 2000. Landscape history as a planning tool. Landscape and Urban Planning, 49,
67–81.
McIntosh, R.P. 1972. Forests of the Catskill Mountains, NewYork. Ecological Monographs, 42,
143–161.
Mueller-Dombois, D. and H. Ellenberg. 1974. Aims and methods of vegetation ecology. John
Wiley and JSons. New York. 547 pp.
New Hampshire GRANIT. 2008, University of New Hampshire, Durham, NH. Website
(http://www.granit.unh.edu). Accessed September 2011.
New Hampshire Highway Dept. 1940. Geologic map and structure sections of the Winnipesaukee
Quadrangle, New Hampshire. Concord, NH.
Nowacki, G.J. & Abrams, M.D. 1992. Community, edaphic, and historical analysis of mixed oak
forests of the Ridge and Valley Province in central Pennsylvania. Canadian Journal of
Forest Research, 22, 790–800.
Odum, E.P. 1969. Strategy of ecosystem development. Science 164: 262-270.
Orwig, D.A. and Foster, D. R. 2002. Stand, landscape, and ecosystem analyses of Hemlock
Woolly Adelgid outbreaks in southern New England: an overview. Journal of
Biogeography. Vol. 10.11:1475-1487.
Pease, A.S. 1911. List of plants on Three Mile Island: Pteridophyta and Spermatophyta.
Appalachia 12: 266-276.
Phillips, E.A. 1959. Methods of Vegetation Study. Henry Holt and Company, Inc. 107 pp.
19
Holland et al. – Vegetation changes in temperate forest
Rogovin, L. S. 1983. A History of Three Mile Island Camp. Published by Three Mile Island
Camp, Meredith, NH.
Russell, E.W.B. 1981. Vegetation of northern New Jersey before European settlement. American
Midland Naturalist, 105, 1–12.
Siccama, T.G. 1971. Presettlement and present forest vegetation in Northern Vermont with special
reference to Chittenden County. American Midland Naturalist, 85, 153–172.
Smith, D. 1962. The Practice of Silviculture. John Wiley and Sons, Inc., Chapter 2.
______. 1986. The Practice of silviculture. Eighth edition. John Wiley and Sons, Inc. Ch. 2.
USDA Forest Service. 2005. An Exotic Pest Threat to Eastern Hemlock: An Initiative for
Management of Hemlock Woolly Adelgid.
www.na.fs.fed.us/fhp/hwa/pub/hwa_mgt_plan.pdf Accessed 6 October 2011
United States Department of Agriculture Forest Service. August 2005. Hemlock Woolly Adelgid.
(www.na.fs.fed.us/spfo/pubs/pest_al/hemlock/hwa05.htm). Accessed 13 Nov. 2011.
USDA, NRCS. 2011. The PLANTS Database. National Plant Data Center, Baton Rouge, LA,
Website (http://plants.usda.gov). Accessed September 2011.
Whitney, G.G. 1990. The history and status of the hemlock hardwood forests of the Allegheny
Plateau. Journal of Ecology, 78, 443–458.
20
Holland et al. – Vegetation changes in temperate forest
Table 1. Total reported, new taxa, and persistence from previous collections of plant
species at Three Mile Island, New Hampshire, USA (from Holland and Sorrie (1989). * = Nine
species planted by Kelsey were indigenous to the islands; ** = Ten species first observed on the
islands by Hartmann persisted in 1989.
Number of Species
Persistence of
NonIndigenous
Total
Species
Persistence of
Species
Introduced by
Indigenous
Reporte
New to
Kelsey in
Species Reported
d
TMI
1901
by Pease (1911)
60
-
-
-
Pease (1911)
265
-
51
-
Hartmann (1941)**
193
31
9
153
MMH/BAS (1989 study)
243
66
10
157
Collector
Kelsey (1902)*
21
Holland et al. – Vegetation changes in temperate forest
Table 2. Significance of differences of the mean number of overstory species per circular plot
among the years indicated (from Clapham, Holland and Fedders 2009).
1978 to 1991
1991 to 2001
Land use zone
Protective
0.031
0.344
Productive
0.068
0.471
Compromise
0.05
0.182
Urban
0.284
0.793
22
23
TOTALS
Viburnum acerifolium L.
Viburnum nudum L. var. cassinoides (L.)
Torr. & A. Gray
Tilia americana L.
Tsuga canadensis (L.) Carrière
Vaccinium corymbosum L.
Robinia hispida L.
Rhus typhina L.
Sambucus nigra L. ssp. canadensis (L.) R.
Bolli
Nyssa sylvatica Marsh.
Ostrya virginiana (Mill.) K. Koch
Pinus resinosa Aiton
Pinus strobus L.
Populus grandidentata Michx.
Populus tremuloides Michx.
Prunus pensylvanica L. f.
Prunus serotina Ehrh.
Quercus alba L.
Quercus rubra L.
Ilex mucronata (L.) Powell, Savolainen &
Andrews
Ilex verticillata (L.) A. Gray
Lyonia ligustrina (L.) DC.
Hamamelis virginiana L.
Fraxinus americana L.
Gaylussacia baccata (Wangenh.) K. Koch
Halesia carolina L.
Fagus grandifolia Ehrh.
Betula populifolia Marsh.
Cephalanthus occidentalis L.
Betula lenta L.
Betula papyrifera Marsh.
Amelanchier laevis Wiegand
Betula alleghaniensis Britton var.
alleghaniensis
Alnus incana (L.) Moench ssp. rugosa (Du
Roi) R.T. Clausen
Species
Acer pensylvanicum L.
Acer rubrum L.
Acer saccharum Marsh.
Year
1
0
1
4
5
8
19
4
1
1
3
2
18
1
1
1
1
14
5
7
3
204
0
1
16
0
0
0
146
3
5
9
0
1
16
3
1
1
16
0
3
16
0
0
17
0
0
0
13
18
19
6
3
3
Freq
0
0
17
14
9
1991
0
0
0
2
3
9
19
0
3
0
2
0
19
0
0
Freq
1978
0
0
0
100
0
0
2 0.68
18 10.96
5
0
7
0
1
0
2
0
2
0
3 1.37
4 2.05
9 6.16
18 13.01
3
0
1 2.05
0
0
4 1.37
1
0
19 13.01
1
0
0
0
0
0
1 2.05
12 10.96
2
0
0
0
18 11.64
2
0
2
0
0
0
16
8.9
3
1
Table 3. Calculations of Importance Values : Three Mile Island (TMI)
0
0.97
8.7
2.4
3.4
0.49
0.97
0.97
1.5
1.9
4.4
8.7
1.5
0.49
0
1.9
0.49
9.2
0.49
0
0
0.49
5.8
0.97
0
8.7
0.97
0.97
0
7.7
1.5
0.49
9.2
9.2
3.8
1.47
1.5
100 99.76
0.49
0.49
6.86
2.45
3.43
0.49
0
0.49
1.96
2.45
3.92
9.31
1.96
0.49
0.49
1.47
0.98
8.82
0.49
0.49
1.47
2.45
4.41
0
0.49
7.84
1.47
0.49
0.49
7.84
1.47
1.47
8.82
9.31
2.94
0
1968
0
3
236
0
0
0
0
0
3
64
49
345
0
11
0
2
0
311
0
0
0
48
113
0
0
293
0
0
0
190
0
0
264
209
66
13
3759
1
10
160
26
35
92
0
2
27
142
36
321
12
2
3
8
5
275
94
7
16
16
57
0
1
293
4
8
1
811
116
20
562
556
27
29
4537
0
4
259
90
30
109
5
25
17
188
50
362
12
2
0
14
1
326
9
0
0
2
97
8
0
531
5
16
0
972
104
15
776
410
69
0
100
0
0.15
11.99
0
0
0
0
0
0.15
3.25
2.49
17.53
0
0.56
0
0.1
0
15.8
0
0
0
2.44
5.74
0
0
14.89
0
0
0
9.65
0
0
13.41
10.62
3.35
0.35
100
0.03
0.27
4.26
0.69
0.93
2.45
0
0.05
0.72
3.78
0.96
8.54
0.32
0.05
0.08
0.21
0.13
7.32
2.5
0.19
0.43
0.43
1.52
0
0.03
7.79
0.11
0.21
0.03
21.57
3.09
0.53
14.95
14.79
0.72
0.64
99.96
0
0.09
5.7
1.98
0.66
2.4
0.11
0.55
0.37
4.14
1.1
7.98
0.29
0.04
0
0.31
0.02
7.19
0.2
0
0
0.04
2.14
0.19
0
11.7
0.11
0.35
0
21.42
2.29
0.33
17.1
9.04
1.52
0
4255
0
6
416
0
0
0
0
0
8
76
105
713
0
34
0
0
3
795
0
0
0
94
304
0
0
537
0
0
0
245
0
0
386
408
125
13
5683
1
16
345
26
35
92
0
2
35
180
63
581
30
5
3
9
13
799
94
7
37
41
119
0
1
455
9
8
2
824
120
20
674
965
59
29
6578
0
9
462
90
30
109
5
25
25
252
100
614
41
4
0
16
4
899
9
0
0
7
182
9
0
744
11
16
0
945
110
15
899
783
134
0
28.29
0
2
1.76
0
0
0
0
0
2.67
1.19
2.14
2.07
0
3.09
0
1.5
0
2.56
0
0
0
1.96
2.69
0
0
1.83
0
0
0
1.29
0
0
1.46
1.95
1.89
1
1.6
2.16
1
1
1
0
1
1.3
1.27
1.75
1.81
2.5
2.5
1
1.13
2.6
2.91
1
1
2
1.02
1
53.45
1
2.31
2.56
2.09
0
1
1.55
2.25
1.03
1
1.2
1.74
2.19
1
52.38
0
2.25
2.05
1
1
1
1
1
1.47
1.34
2.27
2.06
3.42
2
0
1.14
4
3.23
1
0
0
3.5
1.9
1.13
0
1.54
2.5
1
0
1.08
1.06
1
1.35
2.15
1.94
0
100
0
7.07
6.23
0
0
0
0
0
9.43
4.2
7.57
7.3
0
10.93
0
5.3
0
9.04
0
0
0
6.48
0
0
0
4.56
0
6.92
9.51
0
0
5.17
6.9
6.69
1.87
100
1.87
2.99
4.03
1.87
1.87
1.87
0
1.87
2.43
2.37
3.27
3.39
4.68
4.68
1.87
2.1
4.86
5.44
1.87
1.87
1.87
2.91
4.21
1.87
3.74
1.9
4.33
4.79
3.91
1.94
1.87
2.24
3.25
4.09
0 2.16 1.93
0 1.29 0.91
0
0
0
3.65
3.17
5.41
12.62
0
4.51
0
2.26
0
12.62
0
0
1.6
0
0.8
1.7
2.87
2.72
7.08
2.32
1.74
0.81
1.26
1.99
7.19
1.62
0.85
1.6
0.99
1.14
1.56
2.86
3.28
6.87
2.75
1.45
0
1.46
2.7
7.52
0.87
0
1.91
99.84
0 1.23 1.35
100 100 99.85
0
0 0.8
0
4.29 2.64 1.25 1.78
3.91 9.73 5.05 6.1
1.91
0 1.67 2.1
1.91
0 2.08 1.99
1.9
1.9
1.9
2.81
2.55
4.33
3.93
6.5
3.81
0
2.17
7.6
6.16
1.91
0
0
0 2.4
0
6.7 3.81 2.23 2.41
3.6 8.74 3.28 3.85
2.2
1.12
0
0 0.8
0
2.94
11 6.18 7.78
4.77
0 1.93 1.95
1.9
0 0.86 1.07
0
0 1.42
0
2.06 7.71 10.44 10.39
2
1.9
2.57 10.08 8.67 9.62
4.1 9.04 9.12 7.45
3.7 5.4 2.58
3
1978 1991 2001
1978
1991
2001 1978 1991 2001
1978
1991
2001
1978
1991
2001
1978
1991
2001 1978 1991 2001
Rel.
Rel.
Rel.
Rating Rating Rating
Imp Imp Imp
Freq Freq Freq Density Density Density Rel. Den Rel. Den Rel. Den summed summed summed Dominance Dominance Dominance Rel. Dom Rel. Dom Rel. Dom Values Values Values
19 11.64
19 9.59
8 6.16
3
206
Freq
2001
Holland et al. – Vegetation changes in temperate forest
24
TOTALS
Viburnum nudum L. var. cassinoides
(L.) Torr. & A. Gray
Viburnum recognitum Fernald
Pinus resinosa Aiton
Pinus strobus L.
Populus tremuloides Michx.
Quercus rubra L.
Tsuga canadensis (L.) Carrière
Vaccinium corymbosum L.
Lyonia ligustrina (L.) DC.
Hamamelis virginiana L.
Betula populifolia Marsh.
Gaylussacia baccata (Wangenh.) K.
Koch
Betula lenta L.
Betula papyrifera Marsh.
Amelanchier laevis Wiegand
Species
Acer pensylvanicum L.
Acer rubrum L.
Acer saccharum Marsh.
Alnus incana (L.) Moench ssp. rugosa
(Du Roi) R.T. Clausen
Year
0
0
20
1
1
30
0
2
1
3
3
1
3
3
0
2
2
0
3
1
1
1
1
1
1
0
1
0
2
3
0
3
2
0
1
1
2
1
2
1
0 0 3.125
5 6.67 6.25
0 3.33 6.25
10 10 9.375
15 10 9.375
0 3.33 0
15 10 9.375
10 10 9.375
0 0 6.25
5 6.67 6.25
5 6.67 9.375
5 0 0
5 10 6.25
5 3.33 3.125
5 3.33 0
10 3.33 6.25
5 6.67 0
56
31
0
6
37
1
5
8
85
27
0
4
67
1.69
0
17.51
26.55
0
10.73
28.25
0
0.56
0.56
2.26
1.69
6.78
0
2.69
13.87
7.87
8.49
0.62
7.87
27.33
0
11.59
6.42
0
1.24
7.66
1
1
5
5
12
1
5
2
56
31
0
15
42
1
13
10
0
1.5
0
4.82
4.82
6.03
8.04
4.82
0
0
100
3.9
3.9
100
0
4.42
7.03
9.61
9.84
2.62
8.62
15.67
0
2.9
4.81
6.18
9.77
9.56
0
8.6
15.1
3.74
7.39 7.76
5.66 5.85
0 0
6.99 6.01
5.14 5.8
4.06 0 2.75 5.67
4.06 0 2.55 2.7
100 100 100 100
0
3.84
0
13.27
18.23
0
12.56
15.68
0
3.46
3.46
4.43
4.91
5.53
0 3.46 2.48 0
6.09 6.1 4.83 5.54
0 5.07 4.4 0
3.9 4.06
3.9 4.06
0
0
9.74 11.17
4.42 4.06
4.82 3.9
6.03 10.13
9.65 4.87
1
0
0 4.06
1 4.82 3.9 4.06
1
0 3.9 4.06
3.33 12.29 10.97 13.54
2.72 13.14 11.02 11.06
0
0 3.9
0
2.69 11.93
8 10.95
2.61 8.78 9.68 10.62
1
0
0 4.06
85 1 1 1
27 1 1 1
0 1.25 0 0
11 1.67 2.5 2.75
67 1 1.14 1
0 1 1
42 1.25 2.6
0 2 1.25
1.52 0 0 10
0
4.12 3 13 27 1 1
8.23 0 67 54 0 1
6.4 79 107 140 2.55 2.82
8.23 128 116 147 2.72 2.83
0 0 3 0 0 1
5.49 47 78 97 2.47 2.05
25.3 91 328 434 1.82 2.48
0.91 0 0 6 0 0
12.96
4.12
0
0.61
10.21
0 0.56 0.21 0
28 2.26 1.04 4.27
0 0.56 1.66 0
0 0 10
3 13 27
0 67 54
31 38 42
47 41 54
0 3 0
19 38 36
50 132 166
0 0 6
1
1
4
3
12
1
4
1
2 0 3.33 6.25 0 5 44 0 1.04 6.71 0 5 44 0 1 1
1 0 3.33 3.125 0 2 6 0 0.41 0.91 0 2 6 0 1 1
32 100 100 100 177 483 656 100 100 100 380 887 1197 20.73 25.67 24.61
1
2
2
3
3
0
3
3
2
2
3
0
2
1
0
2
0
1979 1991 2001 1979 1991 2001 1979 1991 2001 1979 1991 2001 1979 1991 2001 1979 1991 2001 1979 1991 2001 1979 1991 2001
Rating Rating Rating Dominan Dominan Dominan
Imp Imp Imp
Freq Freq Freq Rel.Freq Rel.Freq Rel.Freq Density Density Density Rel. Den Rel. Den Rel. Den summed summed summed ce ce ce Rel. Dom Rel. Dom Rel. Dom Values Values Values
Table 4. Calculations of Importance Values: Hawk's Nest Island
Holland et al. – Vegetation changes in temperate forest
25
TOTALS
Viburnum nudum L. var. cassinoides
(L.) Torr. & A. Gray
Tilia americana L.
Tsuga canadensis (L.) Carrière
Vaccinium corymbosum L.
Spiraea alba Du Roi var. latifolia
(Aiton) Dippel
Pinus resinosa Aiton
Pinus strobus L.
Populus grandidentata Michx.
Populus tremuloides Michx.
Prunus pensylvanica L. f.
Prunus serotina Ehrh.
Quercus rubra L.
Picea rubens Sarg.
Nyssa sylvatica Marsh.
Ilex mucronata (L.) Powell, Savolainen
& Andrews
Ilex verticillata (L.) A. Gray
Lyonia ligustrina (L.) DC.
Myrica gale L.
Hamamelis virginiana L.
Fraxinus americana L.
Gaylussacia baccata (Wangenh.) K.
Koch
Halesia carolina L.
Fagus grandifolia Ehrh.
Betula populifolia Marsh.
Betula papyrifera Marsh.
Amelanchier laevis Wiegand
Species
Acer pensylvanicum L.
Acer rubrum L.
Acer saccharum Marsh.
Alnus incana (L.) Moench ssp. rugosa
(Du Roi) R.T. Clausen
Year
Freq
0
0
1
0
0
2
0
21
0
0
3
1
0
33
0
1
3
2
1
1
1
1
2
2
3
2
0
0
2
0
0
2
2
1
3
0
0
1
0
1
3
1
0
0
0
0
0
2
1
0
1
0
0
0
0
1
0
3
0
Freq
1991
1
2
1
1979
Freq
3
46
1
0
3
3
1
3
2
1
3
2
2
2
1
0
1
1
0
1
0
1
3
3
2
3
1
0
0
3
0
2001
0
100
0
0
14.29
4.76
0
0
0
0
9.52
4.76
0
4.76
0
0
0
0
4.76
0
0
9.52
4.76
0
4.76
14.29
4.76
0
4.76
9.52
4.76
0
100
0
3.03
9.09
6.06
3.03
3.03
3.03
3.03
6.06
6.06
9.09
6.06
0
0
6.06
0
0
0
0
3.03
6.06
6.06
3.03
9.09
0
0
0
9.09
0
1991
6.52
100
2.17
0
6.52
6.52
2.17
6.52
4.35
2.17
6.52
4.35
4.35
4.35
2.17
0
2.17
2.17
0
2.17
0
2.17
6.52
6.52
4.35
6.52
2.17
0
0
6.52
0
2001
0
131
0
0
56
5
0
0
0
0
5
1
0
17
0
0
0
0
5
0
0
12
3
0
4
10
1
0
1
10
1
1979
0
2176
0
29
32
803
352
250
20
409
11
18
30
5
0
0
8
0
0
0
0
6
43
21
4
103
0
0
0
32
0
1991
93
2764
10
0
42
652
746
626
179
19
21
5
14
28
2
0
17
14
0
1
0
5
96
36
2
117
1
0
0
38
0
2001
0
100
0
0
42.75
3.82
0
0
0
0
3.82
0.76
0
12.98
0
0
0
0
3.82
0
0
9.16
2.29
0
3.05
7.63
0.76
0
0.76
7.63
0.76
1979
0
100
0
1.33
1.47
36.9
16.18
11.49
0.92
18.8
0.51
0.83
1.38
0.23
0
0
0.37
0
0
0
0
0.28
1.98
0.97
0.18
4.73
0
0
0
1.47
0
1991
3.36
100
0.36
0
1.52
23.59
26.99
22.65
6.48
0.69
0.76
0.18
0.51
1.01
0.07
0
0.62
0.51
0
0.04
0
0.18
3.47
1.3
0.07
4.23
0.04
0
0
1.37
0
2001
Table 5. Calculations of Importance: Blueberry Island
0
239
0
0
109
5
0
0
0
0
6
3
0
32
0
0
0
0
12
0
0
12
3
0
9
18
1
0
1
26
2
0
2419
0
65
53
803
352
250
20
409
27
48
95
12
0
0
8
0
0
0
0
6
43
22
11
139
0
0
0
56
0
93
3081
10
0
102
683
746
626
179
19
43
13
42
89
3
0
17
21
0
1
0
5
96
47
4
165
2
0
0
75
0
0
24.08
0
0
1.95
1
0
0
0
0
1.2
3
0
1.88
0
0
0
0
2.4
0
0
1
1
0
2.25
1.8
1
0
1
2.6
2
0
29.48
0
2.24
1.66
1
1
1
1
1
2.45
2.67
3.17
2.4
0
0
1
0
0
0
0
1
1
1.05
2.75
1.35
0
0
0
1.75
0
1
35.99
1
0
2.43
1.05
1
1
1
1
2.05
2.6
3
3.18
1.5
0
1
1.5
0
1
0
1
1
1.31
2
1.41
2
0
0
1.97
0
0
100
0
0
8.08
4.15
0
0
0
0
4.98
12.46
0
7.82
0
0
0
0
9.97
0
0
4.15
4.15
0
9.34
7.48
4.15
0
4.15
10.8
8.31
0
100
0
7.6
5.62
3.39
3.39
3.39
3.39
3.39
8.33
9.04
10.74
8.14
0
0
3.39
0
0
0
0
3.39
3.39
3.55
9.33
4.58
0
0
0
5.94
0
2.78
100
2.78
0
6.75
2.91
2.78
2.78
2.78
2.78
5.69
7.22
8.34
8.83
4.17
0
2.78
4.17
0
2.78
0
2.78
2.78
3.63
5.56
3.92
5.56
0
0
5.48
0
0
100
0
0
21.71
4.24
0
0
0
0
6.11
5.99
0
8.52
0
0
0
0
6.18
0
0
7.61
3.74
0
5.72
9.8
3.23
0
3.23
9.32
4.61
0
100
0
3.99
5.39
15.45
7.53
5.97
2.45
8.41
4.96
5.31
7.07
4.81
0
0
3.27
0
0
0
0
2.23
3.81
3.53
4.18
6.13
0
0
0
5.5
0
4.22
100
1.77
0
4.93
11.01
10.65
10.65
4.53
1.88
4.32
3.92
4.4
4.73
2.14
0
1.86
2.28
0
1.66
0
1.71
4.26
3.82
3.33
4.89
2.59
0
0
4.46
0
1979
1991
2001
1979
1991
2001
1979
1991
2001
1979
1991
2001
Rating Rating Rating Dominan Dominan Dominan
Imp
Imp
Imp
Rel.Freq Rel.Freq Rel.Freq Density Density Density Rel. Den Rel. Den Rel. Den summed summed summed ce
ce
ce
Rel. Dom Rel. Dom Rel. Dom Values Values Values
1979
Holland et al. – Vegetation changes in temperate forest
Holland et al. – Vegetation changes in temperate forest
FIGURES
Figure 1: Frequency and cumulative frequency distributions for the number of overstory
species per circular plot in the four ecological zones for the years 1978, 1991 and 2001.
26
Holland et al. – Vegetation changes in temperate forest
Figure 1.
27