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The Catskill Mountain High Elevation

Forest Continuum

Matt Heimel


Geography 399 Independent Study: Advanced Biogeography
SUNY Oneonta Summer 2011
Wittenberg Mountain, the Central Escarpment, and the Blackhead Range beyond, as seen from Cornell Mountain.
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1. Introduction
Earth and the life that inhabit it have evolved together. Continents are continuously being
reshaped, and the form a landmass may take has reaching effects into the future. Regional ecosystems
throughout natural history must sustain themselves and adapt to the changing physical restrictions within
their habitat, and the genetic links that survive are the key to what adaptations and life forms will be
characteristic to the future forms of the planet. These statements may be applied to the Catskill
Mountains; these relatively small mountains are the depository remnants of a once towering mountain
range. Ecology of the ancient delta, evidence of which is recorded in the bedrock strata, reveals
alternating patters of shallow seas, floods, and riparian systems altered by evolving vegetation. Lifes
adaptations to survive in the ancient environment are links to modern life forms for parallel present
environments. Continental glaciations had a major role in further eroding the mountains and adding final
touches. The progressive return of vegetation after glaciations shows a primary succession pattern
following a denuded landscape, leading to a summit forest community displaying characteristics of the
boreal forest ecosystem, presently undergoing a gradual transition to a hardwood community.
The vast, ancient depositional delta has eroded away through glaciations and flowing water.
Highest elevations in the Catskill Mountains are what remains of lesser-transported alluvial deposits. That
zone is the habitat for the spruce-fir assemblage of the Catskill forest community, a part of the eastern
forest biome. The purpose of this study is to present a comprehensive biogeographical natural history and
analysis of the dominant forest ecosystem factors for the highest elevations in the Catskill Mountains. The
five highest mountaintops were chosen as sample plots for discussion because they represent an ultimate,
limiting parameter to vegetational development within the entire Catskill ecosystem. Dominant over,
middle, and under stratum species are determined and their adaptations and factors for distribution are
discussed. Environmental factors at these elevations are unique from the lower elevations because of the
regions topography and history, and dominant species of vegetation respond to these differences.

2. Geographic Location & Environment
The Catskill Mountain ecological sub region discussed in this paper is the northeastern inclusion
of the geomorphological Appalachian Plateau province. Structurally it is a maturely dissected plateau,
with flat mountaintops separated by weathered drainage channels and basins [figure 2-1]. On its eastern
front, there is a 2,000 to 3,000 steep escarpment into the Hudson Valley [figure 2-2]. On the western
front, there is a gentle merge into a hilly landscape typical of the western Catskill region and the
Allegheny Plateau [figure 2-3]. Within the Allegheny and northeast Appalachian Plateau regions, the
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Catskill Mountains have the highest elevations, ranging
from ~900 to 4,182 feet. Most of the terrain is steep, with a
typical steppe like pattern comprised of sandstone ledges
with intermittent slopes containing boulders and glacial till
under organic soils.
The forests on the five highest mountains were used
as sample plots representing the climax spruce-fir
ecosystem. These relatively small areas of forest cover
were chosen because they represent the highest reaches of
elevation on individual mountains. Although the area is
structurally one dissected plateau, each slope has
undergone distinct vegetational migrations to the top
following a post-glacial primary succession pattern. There
are mountains below the five highest elevations that share
the same spruce-fir communities, and some dominated by
hardwoods. The theory presented here is that these
communities are distributed independently of elevation,
precipitation, and temperature, and the presence of spruce-
fir forests is attributed to the soil conditions. The five
highest elevations will provide an analysis of what species
dominate those mountains, and the physiographic site
characteristics can be inferred by what species are present.

2a. Climate
The Catskills are within a severe mid-latitude
climate. Temperatures change along a wide spectrum between warm, humid summers and severely cold
winters. Precipitation decreases during the winter months, but overall it is evenly distributed throughout
the year [figure 2-4]. The prevailing westerlies, from the southwest, are the main wind belt for
atmospheric systems in the area. However, seasonal storms such as hurricanes or tropical storms
occasionally reach the Catskills and bring heavy outbursts of precipitation, contributing to flooding and
erosion of riparian areas (Thaler, 1996).
Figure21

Figure22

Figure23

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While the prevailing winds and
atmospheric circulatory patterns bring
the overall climatological classification
to the Catskill region, the primary
differences in localized climate between
mountains are caused by local variations
in weather patterns. Occurrences of
upsloping, downsloping, and the
orographic effect are patterns that
dictate temperature and precipitation differences. The
central Catskill Mountains containing the major
escarpments, and the southwestern mountains with
their intense drainage basins, receive greater rainfall
than the lowlands, as air is forced to rise over the
mountains, cools, and is unable to hold its moisture (Thaler, 1996). Annual
precipitation averages for the valleys ranges from 40 to 48 inches, while it can exceed 60 inches on the
higher mountains, contributing to the extremely lush vegetation and the abundant streams [figure 2-5].
Growing seasons can last between 120 to 160 days, decreasing as elevation increases, which is
evident in the apparent greening of mountain slopes during spring (USGS, 2003). Snow and ice can
remain on the higher mountains through April while plant growth has begun in the lower valleys. Ice and
high winds are a significant disturbance regime in the Catskill forest, commonly causing localized tree
mortalities but stand blowdowns do occur either associated with the heavy buildup of ice or soil and root
saturation (Johnson, 2007).

2b. Soils and Vegetation
Edaphic characteristics of the higher elevation forests are the result of their geological, glacial, and
vegetational histories. The parent material is primarily conglomerate sandstone in the higher elevations.
Over the course of thousands of years, freeze-thaw action and deposition by glaciers has created a mineral
layer or bedrock fractures that vegetation may grow upon once sufficient organics accumulate.
There is no well-developed soil profile present in the higher forests and soils are generally a thin
layer of organics overlying substrate. A mixture of inceptisols and spodosols are present, where the
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Slide Mountain Weather Station, 2647' AMSL
Precipitation, in. Temperature, deg.F
2-4: Climograph for higher elevation areas.
Source: NCDC TD 9641 Clim 81 1961-1990
Normals

2-5: Annual
Precipitation map
for Catskill
region. Higher,
southwestern
mountains receive
above 60 inches.
Source: NRCS
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recently glaciated landscape has been denuded of organics and the organic accumulation is still underway
(Kudish, 1979).
The quantity of glacial till is a primary deciding factor in what tree species has been able to
dominate an area (Kudish, 1979). In the high elevation forests, balsam fir, red spruce, and occasional
paper birch are a community preceding hardwood establishment. These species have a lower required
glacial till thickness, and the tops of the Catskill mountains have the thinnest till in the region. The
summits typically have less than 8 inches of till overlying bedrock, if any (Mcintosh, 1962). As the
organics accumulate over hundreds of years to build up humus layers, hardwood species are able to
tolerate the new deeper thickness and establish root systems. The pH of glacial till is a nill factor, while it
is mildly acidic throughout the mountains and does not affect tree species distribution (Kudish, 1979).
High elevation forests are principally growing mineral soils derived from conglomerate
sandstones. The higher amounts of precipitation in these areas creates greater amounts of leaching,
allowing better-developed podzols with an organic mat 2 to 8 inches thick (Kudish, 1971). Water
retention in these soils varies and is attributed with different forest covers. Generally, red spruce
dominance is in an area with high water retention while balsam fir is in areas with lower water retention.
Overall, the water retention in high elevation forests capacity is high (Mcintosh, 1962). The organic soils
covered by a dense mat of sphagnum moss retain the water, and some boreal bog-like conditions are
sustained through the intact manner of the spruce-fir high elevation ecosystem (Larson, 1980).

3. Catskill Paleogeography
Ecological processes of the Catskill forest take place upon depository remnants of ancestral
continental landforms. The Acadian mountain range orogeny is the main geological event that reshaped
the northern Appalachians of which the Catskills are a part. This occurred during the late Devonian period
beginning ~375 million years ago and continuing and for another 50 million years, and later another uplift
in the Late Paleozoic pushed the Catskill basin higher to its current position, weathering away ever since.
During the Devonian period, the earths crust was folded and deformed to form the Acadian
mountains. This occurred through lateral plate compression from the ancestral tectonic Avalonia
and Laurentia plates, where Avalonia was pushed over and accreted to Laurentia. The long series
of uplifts and volcanic centers throughout the extremely large mountain range would have released clastic
rocks, slowly breaking down away the Acadians (USGS, 2003). While the tectonic collision proceeded,
the subsidence pressure created by the increased mass of rock along the mountain chain created an impact
formation known as the Catskill basin to east and north of the Acadian mountains (Murphy, 2006). Today,
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6 3-2: Alternating layers of sandstones, conglomerates, and red sandstone conglomerates
indicate diverse periods of stream flow intensities to carry different sized deposits.
the basin would encompass parts of northern New Jersey,
southeastern New York, and eastern Pennsylvania. The
Catskill basin, because of its proximity to the Acadian
highlands, would have received the bulk of clastic rock
from the mountain weathering, while farther to the east an
epicontinental sea would have ultimately received the
basins meandering streams and extremely fine grained
sediments [figure 3-1].
Precipitation drainage from the continuously
eroding Acadian mountains was the principal source for
the accumulation of the Catskill delta (Titus, 2004). The
new land shelf widened over time, and a medium grade of
deposition has been noted in the Alleghany and northern
Appalachian regions of which this formation comprises (USGS, 2003). Closer to the rough boundary of
the Hudson River to the former Acadian Highlands is where the greater Catskill Mountains formed. Their
larger size than the rest of the region is attributed to their composition of primarily sandstones and
conglomerate on the higher mountains, which is much more resistant to weathering than the farther and
more distributed clays, siltstones, and shale that make up valley bottoms and the western Catskill and
Alleghany regions (USGS, 2003). The sedimentary basin eventually reached an elevation of ~7,000 feet.
The depositional landmass along the eastern edge, where the
higher mountains are located, had the thickest accumulation of
sediment with exceptionally high concentrations of
conglomerate and sandstones when compared to the rest of the
deltaic clastic-wedge region (Titus, 2004).
The deposition of Catskill delta material was a complex
series of fluctuating shorelines and flood environments. The
landmass progressively moved north, originating in an area
south of the equator. Its shifting position through the equator
climatological zone caused it to experience shifting weather
patterns (Titus, 2004). This is recorded in alternate layers of
deposited sandstone and shale strata, where there are distinct
boundaries between times of deep flood sequences and average
3-1: The Acadian source area eroded, and its
sediment distribution is traced with a relationship
between distance carried and alluvium size.
Source: USGS
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stream flow deposition [figure 3-2]. High seasonal rainfall in the Acadian highlands would have formed
braided, meandering streams throughout the delta. Before vegetation was able to stabilize the region,
broad, shallow, interconnected streams traversed the alluvial plane. These channels were very unstable,
consisting solely of fine sediments and coarser sands that could be constantly shifted in response to
changing stream currents. This created a challenging aquatic and riparian habitat, as the continually
shifting sediments would hamper root development and organic matter accumulation (Murphy, 2006).
Depositional patterns from the later Devonian period indicate a stream flow pattern more akin to
meandering channels (Murphy, 2006). In such a system, the water is confined to a single channel that
slowly erodes its banks. These banks are usually kept stable by streamside vegetation, and the presence of
these streams would suggest that the vegetation during this era had adapted to the alluvial plane
environment. The Devonian era is also attributed to the emergence of deep-rooted trees, which would
have aided in the stabilization process to create meandering stream channels (Titus, 2004).
As terrestrial vegetation became able to establish itself along the waters edge, some of the first
descendants of modern forests began to appear. The genus Archaeopteris has been found in fossil records
in most Devonian landmasses, including a Catskill site located along the Schoharie Creek in Gilboa
(Titus, 2004). Mature stands of this tree would have created a leaf canopy, moderating the temperature
and moisture regimes and creating a shelter for any invertebrates and microbes present. Morphologically,
Archaeopteris would have resembled a modern fern, only much larger, growing
up to 90 feet and living for 40-50 years [figure 3-4]. The new forest possessed
adaptations allowing it to survive in and stabilize its environment, and leaps in
evolution created a forest ecosystem that can be characteristically linked to
modern forests (Murphy, 2006).
Many of the late Devonian plants that grew in the new forests are
regarded as the ancestors of seed plants. Reproductive and vegetative growth
features are shared, such as the branching clusters common to vascular plant
anatomy (Titus, 2004). It is not completely clear which species creates the link
to modern forest and Devonian, but the ancient forest ecosystem is a picture of
one moment in the evolutionary progression towards the modern forests.

4. Present Day Structure of the Catskill Mountains
After millions of years of deposition, uplift, and erosion, the Catskill Mountains are now in their
modern form. Sedimentary rocks, formed by the river deposits in the ancient Catskill delta, lay stratified
3-4: Archaeopteris. Source:
Devonian Times
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in a single plateau, that has been dissected through
complex phases in glacial advances, glacial melt water
surges, and mass wasting through stream flooding.
Where ancient streambeds distributed their deposits,
depending on weight, is outlined by the weathering
resistant ledges surrounded by eroded valleys. Higher
mountains consist of conglomerate and sandstone,
which would have been deposited in river channels
as coarse sand and pebbles [figure 4-1]. In valley
bottoms around the larger mountains where
significant erosion has taken place, beds of soft red
shale are often found [figure 4-2]. These are
depository remains of ancient soils, paleosols, and are
what is left of the Devonian forests (USGS, 2003).
Wherever an escarpment is exposed for a
relatively long part of its length, it is evident that there
is one large mass of sandstone that has been subjected
to great stress. This is indicated by the presence of intersecting layers of sandstone and joint fractures
[figure 4-3]. During the Alleghanian Orogeny 330 to 225 million years ago, the Catskill delta was uplifted
to become its current plateau (USGS, 2003). In the many years
since the rock has been eroding away, following these fractures.
Multiple channels became streams as precipitation traveled down
the plateaus developing drainage basins.
This has been the erosional pattern for the Catskill
Mountains, where there is one relative plateau elevation across the
greater Catskill region consisting of dense conglomerate
sandstones, broken down into central drainage basins. Three main
escarpments comprise the mountains- Slide Mountain and its
surrounding ridge, Hunter Mountain and the surrounding central
escarpment mountains, and the Blackhead range. Each range is
4-1: Sandstone outcropping on the summit of Black Head.
4-2: Valley bottom soft red shale, eroded by a nearby
stream, under a large sandstone outcrop.
4-3: Massive sandstone wall on Hunter
Mountain over Stony Clove (figure 5-2)
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separated by large valleys that act as major
watersheds for all of the stream water.
Each Catskill Mountain has a common form.
The summits are plateaus, surrounded on all sides by
steep slopes to the valley bottoms [figure 4-4]. There
are gentler spur ridges coming off of the summits,
reaching down to the valleys [figure 4-5]. Between
these ridges are the steeper slopes, with channels and
streams in the centers. It is this common characteristic
that makes the Catskills recognizable as a single
landform unit that has been carved into individual
sections by water and ice. It takes a stretch of the
imagination to conceptualize what forms the mountains
would take without the vegetation, soil, and glacial till
cover.

5. The Effects of Glaciers
During the Pleistocene Epoch of the past 1.6 million years, there have been cyclic ice ages leaving
dramatically altered landscapes as they retreat. The exact forms that the Catskill Mountains took after
each continental ice sheet is unknown, and recent glacition alters the evidence of past glaciations. The last
ice sheet to have had a great effect on the Catskill Mountains was the Wisconian Stage and later scattered
alpine glaciations. ~21,000 years ago this ice sheet would have reached its maximum thickness of over
one mile, completely inundating the Catskills with a sea of ice, and its retreat from the region would be
well underway by ~12,000 years ago (Titus, 2003). This relatively recent event has set conditions for
revegetation patterns and large-scale flood erosion in the modern world.
During the glaciations, the massive slabs of Catskill sedimentary bedrock were abraded and
quarried. Erosion was greatly accelerated by the sheer volume of ice, and the mountains were broken
down into a mixture of rocky fragments varying in size. The advance of glaciers through the valleys and
eventually over mountaintops has left characteristic U-shaped valleys [figure 5-1].Along the eastern
escarpment wall over the Hudson valley, there are two deep cuts into the mountainsides where the
4-4: Cornell & Wittenberg mountain with the characteristic
plateau and steep slopes of every Catskill mountain.
4-5: Spur ridge off the summit of Black Dome
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glaciers are thought to have made
an advance into the mountains
(Titus, 2003). These areas are
Platte and Kaaterskill cloves,
characterized by steep rushing
streams with headwaters at 2,000
feet above the lower valley
floors. Because these two valleys
were deeply cut by advancing
glaciers and eroded into a main
route or gully, they received
much of the melt water once
glaciers retreated and are two
major drainage routes for the
eastern escarpment.
As the climate began to
warm, the Pleistocene ended, and
ice sheets covering the Catskills
were melting and eventually full
deglaciation occurred. Some
lobes remained for a slightly
longer duration in higher
mountain elevations as alpine
glaciers, creating complex
obstacles for melt water and
revegetation (Rich, 1906). Melt
water from the glaciers became a
principal agent of erosion and
deepened many of the central
drainage basins that are prevalent today. The melting process was an array of ice dammed lakes and
raging melt water torrents. Remaining lobes of ice in the mountain valleys blocked the central drainage
basins, and water levels reached a critical point where they could pour over and cut through a valley
Copyrightinfo2.MapsetwascreatedusingMapSetToolKit1.50.gps_mapper
5-1: U-shaped Spruceton Valley. Along the north of the central escarpment, this
map shows the hollowed nature of the valley from a glacier tat once pushed
through it.
Copyrightinfo2.MapsetwascreatedusingMapSetToolKit1.50.gps_mapper
5-2: Topographic view of the deep notch between Hunter and Plateau mountains,
created by a surge of meltwater that was impounded by a glacial lake.
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5-3: One of the many gorges in the southwestern
Catskills, established through glacial melt torrents
and fed by abundant orographic precipitation.
(Titus, 2003). This process created localized, deep-cut notches
between the higher mountains [figure 5-2]. The localized lakes would
have reached elevations between 1300 and 3200 feet, inundating most
of the Catskill region (DEP, 2009). The same lobes of remaining ice
that created lakes were blocking the central drainage areas. This
diverted water through the southeastern valleys near Slide Mountain,
and many gorges were deepened in the southeastern most extension of
the Catskills (DEP, 2009). The drainage channels between ridges in
this area, characteristic of every mountain discussed earlier, are
exceptionally weathered, with deep ravines showing signs of intense
water erosion as the high amounts of melt water was sent through this one region [figure 5-3].
Periods of glaciations in the Catskill Mountains have worked to accelerate erosion of the
sedimentary bedrock, and deposited glacial till throughout the mountains to reorganize the soil regolith
regimes. The Catskill forest has been recovering from this recent significant period that has greatly altered
the landscape. The forest communities that have migrated and established mature climax forests in the
mountains follow a pattern of facultative seral succession, where the physical limitations on life are
altered as the ecosystem develops. In the following sections, the high elevation forests and their species
are evaluated to determine their ecological niche and roles and how the characteristics of todays forests
will be explained.

6. High Elevation Forest Community Analysis
A sampling method was used in order to establish a sound reasoning and analysis for what species
may be discussed as dominant vegetation. A line transect crossing the highest elevation of each mountain
was used, spanning only the highest area and beginning and ending once the elevation started to slope
down from the summit. These transects were evaluated to determine a) if any vegetational change could
be found that can be attributed to slope or aspect b) Which species were encountered in a frequency or
basal area that determines whether or not it is a dominant characteristic species.
The fact that the sample plots are extremely small when compared to the vegetational area and
ecosystems of the mountains has been recognized. The objective of this analysis was to determine what
vegetational systems are dominant and functioning on the highest elevations only. The hypothesis used
during sampling and analysis treats the high elevation assemblage as a sub-system of the entire forest
ecosystem. The natural history and present day condition of the Catskill high elevation forest community
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can be discussed in terms that are consistent with the entirety of the region. A uniform environmental
system has been met with adaptations from species that are best suited for unique physiographic regions.


6a. Slide, 4182 [figure 6-1]
The summit of Slide Mountain is dominated
by balsam fir, growing to their maturity and there
are abundant saplings to continue its dominance.
There were no seedlings of other tree species
present. A dense mat of moss was the primary
understory, while the typical understory herbaceous
species were scattered. Those species have an
affinity for undisturbed humus, and highest summit
area of Slide Mountain is disturbed. The entire length
of the summit transect, leaving out the side slopes, ran
across some form of soil compaction. The actual
highest point is bare substrate with the cement
remnants of an observation tower base, and a well-
worn hiking trail runs across that point. A small
clearing with a sandstone outcrop exists in one portion
of the summit [figure 6-2]. Balsam fir surrounds the
meadow that is grown in with grasses and scattered
paper birch and mountain maple saplings. In the
surrounding forested area, numerous campsites that show
evidence of past heavy use have left compacted soil and
burnt micro sites. However, areas with tighter balsam fir
growth have hindered the development of campsites. The
balsam fir canopy shades these areas and provides a
relatively undisturbed micro site where there is extremely
thick branch growth. Inspection of these more protected
6-1: Slide Mountain as seen from Cornell Mountain
6-2: Clearing in the summit area of Slide.
6-3: Natural disturbance area on Slide.

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area shows that even in these sites there is no well-developed understory herbaceous layer.
Slide Mountain has a very long southwestern spur ridge on a gradual grade. The summit forest
continues down this ridge, and is much more intact than the highest portion summit area. However, there
is intrusion by hardwoods where paper birch and mountain ash have established themselves in the lower
canopy. The understory herbaceous community here is well developed and healthy.
Within close proximity there is a blowdown area where most of the mature balsam fir have been
killed off [figure 6-3]. There is very thick growth of balsam fir saplings, and mountain ash and paper
birch saplings are also widespread in this area and will likely grow to their maximum. Slide Mountain is
very prone to such events, as its summit is the highest and most exposed in the region. It also receives
over 60 inches of annual precipitation a year, far higher than the surrounding lowlands (USDA/NRCS).
This contributes to the extremely lush undisturbed sections of the summit forest.

6b. Hunter, 4046[figure 6-4]
The summit of Hunter, like Slide, is
extremely disturbed. Numerous clearings of the
summit and the construction and maintenance of a
fire tower have left noticeable impacts on the forest
community. The actual summit, marked by a
boulder, lies within a cleared field for a fire tower.
A ranger cabin is present, while grasses and
spruce/fir saplings are reclaiming the clearing. The
transect line for the forest analysis was started on
the edge of the meadow where the summit
continues along a plateau, while the other side of
the meadow would dip down eventually sloping
towards the valley bottom.
Balsam fir, paper birch, and red spruce are
the three tree species present on the summit [figure
6-5]. Their relationship is evident in their
distribution. Some mature balsam firs are present,
but across the summit there are mostly medium
sized red spruce. Red spruce has the tendency to act
6-4: Hunter Mountain as seen from Plateau Mountain
6-5: Hunter Mountain summit forest.
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as a pioneer species in the event of a clearing if soil
conditions permit its establishment. Many of this red
spruce are dying off, with their top halves blown off
by wind. The middle story is made up of very thick
balsam fir saplings, which were able to germinate in
the partial shade created by the open red spruce
canopy. Spread throughout this area are occasional
paper birch, which are dominant trees all throughout
the slopes of Hunter mountain and have been able to
successfully establish themselves on the summit. It is likely that the balsam fir will grow to maturity and
dominance, red spruce will maintain its presence, and paper birch will remain as a codominant member of
this summit forest.
The understory communities that are typical to the Catskill higher summits, whose species are
described in the following sections, were abundant on Hunter Mountain. Unlike Slide, the anthropogenic
disturbance has not left a long impact on the soil conditions for the large forested summit area, and the
moss and humus layers have been able to develop successfully
[figure 6-6].

6c. Blackdome, 3985 [figure 6-7]
Blackdome, Thomas Cole, and Black Head are a part
of the same escarpment and are devoid of red spruce. This is
likely due to unique successional development of these
mountains that has resulted in the dominance of balsam fir
over red spruce, the dynamic of which is described in the
following sections.
There is evidence on Blackdome of periodic single-tree
mortality events, where a gap along the transect line in the
dominant balsam fir canopy is filled with mountain ash or
mountain maple saplings [figure 6-8]. However, seedlings of
balsam fir almost completely cover the mossy forest floor.
They have not yet reached a height that can be considered
6-6: The ground stratum on Hunter is healthy with typical
Catskill herbaceous species over moss stratification.
6-7: Blackhead Range, left to right, Thomas Cole,
Blackdome, Black Head.
6-8: Mountain ash with mountain maple seedlings
below growing in balsam fir canopy openings.
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established, but such a masting event will likely result in a
thick sapling layer similar to that on Hunter Mountain and can
tolerate the partial shade created by dispersed mountain ash.
Some small paper birches are present, taking advantage of the
thin soils. No paper birch seedlings or saplings were observed
that would indicate their eventually increase in frequency. The
forest floor is very lush, with moss mat over 3-4 inches deep covering most of the ground. Dominant
herbaceous species grow in great abundance with the balsam fir seedling mast [figure 6-9]. Overall, the
canopy was not closed, but it will likely become a closed canopy balsam fir stand if the seedlings are
established.

6d. Thomas Cole, 3943
The summit of Thomas Cole was dominated by a
healthy stand of mature balsam fir [figure 6-10]. Like
Blackdome, there was a field-like spread of balsam fir
seedlings. No seedlings or saplings of paper birch or mountain
ash were noted although there were dispersed medium size
occurrences. This further indicates their tendency as
opportunistic colonizers of an area, where a tree mortality and
canopy opening can result in the germination of their seed.
Scattered small hobblebushes were present in the understory.
These are young, and
have the potential to
spread rooting branches
to create a thick local
understory. Dominant
understory species
composition is similar
among the high summits, but Thomas Cole did
have a higher number of mountain woodferns
present. Thomas Coles understory community is unique; the moss layer has developed densely only on
6-9: The understory on Blackdome is extremely lush
with great species biodiversity.

6-10: Balsam fir stand on Thomas Cole, with
mountain woodferns beneath.
6-11, 6-12: Dense moss developing particularly on decaying wood
debris and not the ground. Note the ferns over moss-less ground.
16
rotting stumps and wood debris, and not in thick ground mats like the rest of the sampled high elevation
forests. This could be a competitive dynamic with the also unique fern cover. Perhaps the ferns interact
with the humus in a way that hinders the development of a moss mat, restricting the moss to decaying
wood. [figures 6-11, 6-12].

6e. Black Head, 3942
Of the five mountaintops sampled, Black Heads
summit forest was the densest [figure 6-13]. This is
common among the mountains situated in higher elevation
zones where orographic precipitation collects over a central
point. Black Heads steep slopes dropping from its summit
to the Hudson valley floor, near sea level, are part of a
continuous mountain wall that makes up the eastern
Hudson Valley escarpment. The same ridge of mountains
that contains Hunter extends to this wall perpendicularly,
and the mountains positioned over the valley floor possess
extremely dense forests of balsam fir.
Balsam fir, which dominates Black Heads summit,
grows in great frequency. The first steep slopes from the
summit are a cover of balsam fir with an exceptionally thick
layer of moss seemingly pouring over the ledges [figure 6-
14]. When the fir grows in such a thick manner, the trees often
suppress each other and the overall rate of growth is slowed. Many of
the Catskill Mountains that receive greater orographic precipitation
share this variety of over story cover.
On the opposite side, where the summit slopes away from the
escarpment wall, the forest is more open although the moss mat
remains just as thick. Many of the balsam fir along the sample here
had the top halves of their trunks snapped off by wind. In the
openings, young mountain ash, paper birch, and mountain maple were
6-13: Extremely thick growth on Black Head.
6-14: The slopes over the Hudson Valley receive
greater precipitation. Balsam fir grows at its thickest
and moss at its densest.
6-15: Mountain ash, mountain maple,
and paper birch establishing a small,
dominant stand.
17
growing in greater frequency than openings on the other 4 samples, indicating that they might create
short-term small canopy dominance before the balsam fir saplings break suppression [figure 6-15].

6f. Main Analysis Results
The dominant over story trees are balsam fir and red spruce. In the highest elevations, balsam fir
dominates over red spruce in both frequency and basal area. On Hunter, the only stand sampled where red
spruce are present, the spruce were in decline and balsam fir is positioned to dominate in the future. This
does not indicate that red spruce throughout the Catskills is in decline, but that physiographic conditions
in the five highest elevations are more favorable to balsam fir than red spruce.
Paper birch, mountain ash, and mountain maple are present as opportunistic species, growing
where a canopy disturbance is evident. Mountain ash and Mountain maple do not create dominant
overstory stands in the Catskills and both are a sub-canopy species. However, if paper birch establishes
itself and is able to reproduce it may contribute to significant organic accumulation, furthering soil
development and expediting the migration of deeper-rooted hardwoods into the spruce-fir zone.
The understory stratum has overall higher biodiversity than the overstory. Dominant herbaceous
plants include mountain wood sorrel, clintons lily, sharp-leaved aster, starflower, and goldthread. These
tend to create small but very dense colonies, and their presence usually indicates stable, less disturbed
soil. A large array of mosses, lichens, and woodferns also make up the lush lower strata. Many rotting
pieces of tree debris lie on the ground, covered by a sheet of moss, lichen, and other herbaceous species.
The understory species are mostly small, less than 6 inches in height. Their development occurs on acidic
unfertile organic matter, mainly needles. Mosses are on elevated portions, such as logs or rocks. If higher
elevation hardwoods are able to establish themselves, like sugar maple or beech, the nature of leaf litter
changes, as will the conditions that understory species must be adapted to. Plants must have the ability to
grow above the taller layer of leaf litter, and the dominant understory species of the spruce-fir forests
would be replaced by taller, hardwood associated herbaceous species.

7. High Elevation Forest Community Dominant Species Biogeography
In this section the biogeography of each dominant species is discussed, including general growth
characteristics, morphological adaptations to available sunlight, soil depth, acidity and composition, seed
dispersal, and disturbance dynamics. Through these topics an understanding of the ecological niche and
seral stage each species inhabits will be established to further support the objective of this study. First
each species and forest stratification group and how they interact with their environment must be
18
understood, then a comprehensive discussion is presented that defines and explains the high elevation
forest community as a whole.

7a. Trees
Balsam Fir Abies balsamea [figure 7-1]- Balsam fir is distributed throughout Southeastern Canada and the
northeastern United States. In general, regions with cooler temperatures
with abundant moisture allow its growth, although it is tolerant of harsher
conditions. Annual precipitation during its growing season can vary from
6 to 25 inches and there can be between 80 to 180 frost-free days in the
year, with optimum growth occurring when there are 110 frost-free days
(Frank). This characterizes the balsam fir zones in the Catskill region,
where the areas balsam fir is distributed vary in annual precipitation
amounts.
Balsam fir is well adapted to the shallow glacial till soils atop the
Catskill mountains, as the most important determining factor for growth is
soil depth. It is able to survive in soils rich or poor in organics, but where
its stands are dominant the soils have a thick layer of mor humus and a
predominant gray A soil horizon (Frank). Nutrient leaching through abundant orographic precipitation,
the cooler temperatures, and the acidic coniferous litter layer contributes to the spodosol and inceptisol
soil formation. Cycling of organics and the maintenance of acidic spodosol conditions allow balsam fir to
uphold its dominance. It can tolerate a wide range of acidity, and once established it becomes a major
constituent of the forest cover.
Processes for reproduction and early growth of balsam fir allow it to become a dominant species
in the higher elevation Catskills wherever soils are shallow. Its sprawling coniferous growth shades out
deciduous seedlings and allows it to absorb all available canopy sunlight. The sunlight exposure then
triggers flowering in trees 20-30 years old. 83% of trees in a New Brunswick stand were able to flower
where balsam fir was dominant, 59% in a codominant stand, 6% in the intermediate layer, and none in a
suppressed layer (Frank). This shows the importance for balsam fir to establish itself as a dominant
canopy species wherever it grows in order to further reproduce and how it may become eliminated from
stands if shaded out by competition. Large seed crops are produced on 2 to 4 year intervals, and the
amount of time that seeds may be spread is long while the distances a seed may fall vary. September and
October are peak months for seed distribution and continues on through November. The flat, wind-prone
7-1: Medium sized balsam fir.
19
Catskill summits allow seeds to be distributed over great distances, but the most effective distances are 80
to 200 feet. The seed embryos lay dormant, and the optimal location for development is in a mixed
stratified layer of moist organics and coarse textured minerals (Frank).
Soil moisture and temperature within a tolerable range
are the two most important factors for seedling development.
The dense moss layer, thick organics, and year round leaf
cover attributed with mature coniferous forests have a
compound effect to retain soil moisture in the Catskills,
helping the seedlings germinate and survive. Germination in a
mixed mineral-duff soil is more favorable, as a purely leaf
litter soil tends to desiccate faster and the heavy central
root can extend through the humus. Drought, frost
heaving, and burial by organic litter contribute to
seedling morality before the seedlings can be
considered established. Once the seedling is about 6 inches tall it can be
considered established, and even more so if any branching has occurred
[figure 7-2]. After this point, the trees growth is influenced by the stands
unique competition. Hardwood sprouts, especially mountain maple in the
Catskills, are the main competition for a balsam fir seedling. Developing
balsam fir requires at least 50% full sunlight, and at that level deciduous
seedlings may also sprout in the same stand, bringing competition
(Frank).
The thin layer of organics atop mineral soil found in the Catskills
provides a good location for balsam fir root systems, which are most
commonly in the organic litter layer and the uppermost inches of the
mineral layer, provided there is satisfactory weathering to allow root
growth. Because the root systems are restricted to growing in such a
shallow and poorly developed area, older dense stands are especially
at risk to wind damage or kill after heavy rainfall and winds loosen
the root systems.
Depending on the sites physiography, a mature balsam fir is
7-2: This masting of seedlings can be considered
established. From here, intraspecific competition for
moisture, nutrients, space, and sunlight will occur, killing
off many of the future saplings, to the point where a few
young trees are left to possibly grow to maturity.
7-3: A mature, healthy balsam fir
just below the summit of Hunter.
7-4: This overly crowded stand on
Hunters summit has been stunted from
intraspecific competition.
20
small to medium size ranging from 40 to 60 feet with diameters ranging from 12 to 18 inches. Balsam fir
has a tendency to become well established in the shade of larger trees, waiting for a canopy opening to
grow to its maximum. Some may reach 90 to 100 feet and 30 inches in diameter with a maximum age of
100-200 years, and the extent to which a balsam fir grows is very closely related to its unique site factors,
such as soil depth and mineral layer characteristics [figure 7-3, 7,4] (Frank).
Fire is a major damaging agent for balsam fir, but it is not a natural component of the Catskill
vegetational system and does not play a role in the development of balsam fir stands. Insects such as the
spruce budworm and balsam woolly adelgid are able to defoliate balsam fir and attack the stems, but have
not yet been introduced to the Catskills. Multiple species of
fungi cause decay within a living balsam fir, causing wood
rot by the time they are 70 years old. Currently the stands
of balsam fir in the Catskills are healthy. Instead of
introduced insects and fungi wiping out balsam fir, stands
are slowly being replaced as the soils become increasingly
thicker [figure 7-5]. The buildup of leaf litter allows
deciduous trees to become better established in the event of
a canopy opening. Balsam fir seedling are intolerant of
heavy leaf litter attributed with a deciduous stand, and
can then be eliminated as mountain paper birch,
mountain ash, mountain maple, black cherry, beech and
yellow birch become dominant species.

Red Spruce Picea Rubens [figure 7-6]- The native range of red spruce
is in eastern North America, along the Appalachian Mountains
northward through northern New Jersey, central New York, New
England, and the maritime regions of southeastern Canada. Optimal
growth occurs in cool, moist climates with 36 to 52 inches of annual
precipitation, 100 to 140 days with snow cover, a maximum January
temperature of 20 to 30 F, a maximum July temperature of 70 to 80
F and 90 to 150 frost free days for its growing season (Blum, 1992).
The maximum development for red spruce occurs in higher elevations
in eastern North America mountain chains where there is more
7-5: Just below the summit of Sugarloaf Mountain,
mature paper birch and mountain ash are beginning to
gain supremacy. These balsam firs are still able to flower
as they are not yet in shade, but seedlings will have
difficulty germinating and establishing themselves in the
increased amount of organic matter.
7-6: Healthy, mature red spruce grow to their maximum on the
expansive summit of Plateau Mountain.
21
humidity and rainfall is heavier during the growing season.
Like balsam fir, red spruce grows in mostly acidic soils comprised of spodosols and inceptisols. Its
wider distribution in the Catskills outside of higher elevations is attributed to its ability to grow on sites
unfavorable for many other tree species. A thin organic layer overlying unbroken rock on steep, sloped or
poorly drained valley bottoms with limited oxygen circulation limits the growth of many deciduous
species throughout the Catskills. Red spruce can often be found in such areas growing to its maximum
surrounded by saplings and seasonal streams [figure 7-7].
The maximum growth red spruce is 110 feet tall with a d.b.h of around 34 inches inches (Blum,
1992). Red spruce is able to grow close to that size on slope forests in the Catskills but atop to mountains
within the sample plots of this study it is a medium size tree at maturity, with a 12 to 24 inch d.b.h and 60
to 75 ft height. The strongest factor in determining growth rate is the sunlight condition; red spruce is able
to live in shade for many years but once it grows beyond the sapling stage it requires nearly full sunlight
(Blum, 1992).
Male and female flower buds will grow and open during May, located on the axil from the
previous years growth. Cone buds will separate by July, and mature between September and October to
grow 1.3 to 1.5 inches long. Once mature, the cones will be fully open for only a few days to receive
pollen. Every 3 to 8 years there will be a large seed crop with lighter crops between. Most seeds are
dispersed by wind and are germinated within 350 from the parent stands boundaries (Blum, 1992).
Following seed dispersal and deposition on a favorable seedbed, most of the seeds will germinate
next late may into early July with some germinating in the same autumn as dispersal. On desiccation
prone leaf litter, some seeds may lose their viability or delay germination into August. After one year it is
rare for a seed to remain viable (Blum, 1992). Like balsam fir, the main determining factor for
germination is adequate soil moisture, but unlike balsam fir, red spruce is able to germinate on almost any
soil medium. Mineral soils are the most favorable seedbed, while duff and humus are not as favorable
because they are more prone to heat and drying. Germination requires a light intensity of at least 10% full
sunlight but as the seedlings develop they require at least 50%, the same as balsam fir (Blum, 1992). The
seedlings are prone to the same damaging agents as balsam fir, including drought, frost heaving, and
crushing by hardwood leaf litter or snow. Red spruce seedlings and balsam fir seedlings are controlled by
the same factors but red spruce is much weaker and slower growing during its period of establishment.
While the parameters for germination allow a wide array of conditions, the reproduction ultimately
depends on seedling survival. The root system of red spruce seedlings is exceptionally slow growing for
trees and is very fibrous. This places a critical emphasis on the conditions of the top organic layers of the
22
soil profile. Because the roots grow so slowly, if the duff layer exceeds two inches the seedling root may
not reach moist mineral soil to supply water during a dry period in its growth (Blum, 1992).
If the seedling manages to survive and grow to a height of 6 inches, it can be considered
established. It can meet competition from deciduous trees and brush once established, but it can survive
close to 150 years under suppression as a smaller to medium sized tree until it is finally released. It is also
the last of its associated tree species to begin height and radial growth in the spring and early summer
(Blum, 1992), and this may allow red spruce to grow in a pattern that is responsive to how earlier growing
trees have reached towards the canopy openings. Both spruce and fir are shallow-rooted, but red spruce
roots grow shallower than balsam fir. Red spruce is more susceptible to drought and requires a higher soil
moisture retention capacity. The bulk of feeding roots are located in the humus and in the top inch of
mineral soil.
Whether or not red spruce is more shade tolerant than balsam fir is an unsolved question (Blum,
1992), and the answer may lie in the varying soil conditions and climates that can better suit one species
over the other. The main competition for stand dominance is from balsam fir and other established
hardwoods that produce heavy shade like beech and maple. As soil depths increase, beech, maple and
other hardwoods are able to establish themselves in canopy openings. Beech and maple in particular
produce heavy shade and leaf litter, both of which are
detrimental to red spruce development. In the
Catskills, it is common for scattered, mature red
spruce to grow among a dominant overstory of yellow
birch and paper birch [figure 7-7]. As stated earlier
red spruce has the ability to live suppressed, but the
vigor of its recovery declines with age. In a stand
where this is dynamic takes place, balsam fir is able to
outgrow red spruce once the canopy opens. It will
take about 5 years before red spruce shows accelerated
growth once the opening occurs (Blum, 1992).
Like balsam fir, red spruce is especially prone
to fire damage but this is of little natural concern to the
Catskill Mountains, and forest fires are extremely rare.
Red spruce shares the same biotic damaging agents as balsam fir in the Catskills but has not yet been
ravaged by introduced insect populations. High elevation red spruce growth rates have declined
7-7: On the wetter middle slopes of Hunter Mountain,
~3,000, yellow birch is the dominant species. There are
multiple hollowed-out drainage routes for rain and
snowmelt from the summit, and red spruce takes
advantage of the abundant soil moisture in these small
coves. Its tolerances for optimal growth in high elevations
can be traced to where it is able to grow successful stands
in lower elevations.
23
throughout the Appalachian mountain chain, and this is attributed to an interaction between insects,
diseases and the weakening by anthropogenic air pollutants. Studies have not yet indicated a significant
decline in Catskill stand growth rates, and the influences of anthropogenic pollutants are not yet
understood in the Catskill high elevation forests.

Mountain Paper Birch Betula cordifolia [figure 7-8]- Moist, moderately drained soils at higher elevations
and particularly north facing slopes provide a good habitat for paper
birch. It has a very wide native range across northern North America
wherever physiographic conditions favor its germination and
development, from northwestern Alaska, southeast to British Columbia,
and the northern Rocky Mountains states, and eastward through Canada
and the Great lakes region into New England extending south through
the Appalachian Mountains. It is essentially a northern species adapted
to cold climates, growing well in the harsher more exposed Catskill
summits, tolerating wide variations in precipitation from 12 inches
during the growing season in some areas in Alaska to 60 inches in
eastern mountains (Safford, 1992). Generally, a short cool summer and
a long cold winter during which snow covers the ground for a long
period of time is favorable for paper birch, and these climatic patterns are common to the Catskill
Mountains.
Because paper birch has ample genetic diversity and a very wide range, it is able to tolerate many
soil and topographic conditions. Deep well drained spodosols and inceptisols are the best sites for its
growth, which are common features of vegetational development on glacial deposits throughout its range
(Safford, 1992). In the Catskills, it is more abundant on drier sites with rocky slopes. It grows within a
mixture of other tree species within its forest cover type but because of its shade intolerance it grows in
openings and occupies positions in the upper canopy.
From the middle of April through early June a paper birch at least 15 years of age, optimally
between 40 and 70, can enter its flowering period (Rich, 1906). The seeds are matured from early August
until the middle of September and are then dispersed shortly after ripening. Some seeds may be dispersed
earlier as a result of birds feeding on the developing fruits. A good crop of seeds occurs every other year
and tends to vary in amount from one region to another. If a crown grows too many catkins, the seed
7-8: Paper birch and its unique bark.
24
bearing fruit, the crown may deteriorate and foliage quality will decrease and growth will be stunted
(Safford, 1992).
Paper birch seeds are light and wing shaped, lending them to wind dispersal over great distances.
Despite the ability for far travel, most seeds fall under the stand in which they were produced (Safford,
1992). As a consequence of their small size paper birch seeds are extremely fragile. They are very
sensitive to moisture, temperature, light, and the soil medium of the seedbed. Mineral soil offers the best
site for germination, rates of germination on humus are reduced by 50%, and germination rates on
unbroken leaf litter are 10% of that in mineral soil, and twice as many seeds will germinate in a shaded
site than a full sun site (Safford, 1992). Although paper birch colonizes openings in the Catskill spruce/fir
forests, they do not germinate well in the full sun wind thrown openings. Instead, the established
seedlings can be found around overturned tree pits, mounds, and log micro sites where there is a shaded
dip and soil nutrients have been upturned and recycled (Kudish, 1971). Of these germinated seeds, those
located on humus will fare better than those germinated on leaf litter because of greater nutrient
availability. The heights of the seedlings after the first year of growth indicate paper birchs preference for
germination in disturbed partially shaded areas, where taller growth was found in areas with 45% full
sunlight, rather than 100%, or even 25% or 13% (Safford, 1992).
Thin soils atop the Catskill Mountains allow paper birch to establish itself with its shallow root
system. Within the top 24 inches of soil most of the feeding roots can be found, and taproots do not form
(Kudish, 1979). If an extension of the root system grows to a diameter of the stem diameters, it will
become part of the permanent woody root system. Although the roots do not extend deep into the soil,
they do not contribute to wind damage. High wind will break the trunk of a paper birch more often than it
will uproot it, and these broken stems will generally sprout (Safford, 1992).
Young paper birch will grow rapidly, often reaching a diameter of 8 inches after 30 years. This
growth rate will decline with age, and at an older age its growth rate is insignificant. In a mature stand the
average d.b.h. will be 25 to 30 cm and 70 ft in height. Under optimal conditions a tree in an old growth
stand may exceed 76 cm d.b.h. and 100 feet in height. After 60 to 70 years the tree will mature, and few
will live over 200 years (Safford, 1992). Mortality rates are high within a paper birch stand. An individual
tree will express its role in the community by dispersing seeds early in its life, while older trees grow
slowly and do not reproduce as vigorously. If a tree is suppressed, it will shortly die unless released early
in its life.
During the late 1930s and 1940s, large portions of paper birch were killed off or severely damaged
all throughout its eastern range. From the crown back, the twigs and branches died or lost vigor,
25
eventually killing off the entire tree during a period of 5 to 6 years (Safford, 1992). Shallow rooted trees
were most often damaged, and the root systems displayed symptoms before the crown. Many of these
trees sprouted new branches and eventually recovered, while the dieback subsided and is not considered a
current threat. The dieback may be attributed to acid deposition from fog. Although current populations in
the Catskills remain healthy, it is important to consider the possibility for unexpected ecological cascades
of degradation.
The forest floor is enriched with nutrients when paper birch leaf litter is in abundance. Calcium,
potassium, magnesium, phosphorus, and boron increase while manganese, aluminum, iron, and zinc
levels decrease. The nutrient enrichment can extend through thin organics to the top inch of mineral soil
where pH is also increased (Safford, 1992). Alteration of nutrient levels in soil previously occupied by a
coniferous forest with different nutrient parameters contributes to the decline and replacement of
spruce/fir stands (Kudish, 1971). The increases in nutrients, alkalinity and leaf litter contributes to the
process by which an increased number of hardwood species are able to colonize and establish themselves
in areas previously out of range, thus crowding out the spruce/fir stands.
Paper birch will either form a pure stand following a clearing by wind and ice in the Catskills or it
will establish itself in a mixture of varying extent within the spruce/fir community. More commonly it
will become an intimate feature of the Catskill spruce/fir forest, where it occupies openings in the canopy,
slowly altering the nutrient and organic composition of the forest
floor while burying conifer seedlings with its duff. Higher
elevation hardwoods, including beech, maple, yellow birch, and
black cherry are then able to establish themselves after a
considerable amount of time.

Mountain Ash Pyrus Americana [figure 7-9] American mountain
ashs native range occurs in northeastern North America, spanning
from Newfoundland and Nova Scotia south to New Jersey
and Pennsylvania. It appears in mountainous areas through
South Carolina and Georgia, and in the west it can be found
in Minnesota and eastern North/South Dakota where its
environmental parameters are met. It is a component of the
spruce-fir ecosystem as a member of the lower canopy, and
is codominant with balsam fir (Sullivan, 1992).
7-9: Young group of mountain ash.
7-10: Mountain ash branches grow spreading,
rounded crowns.
26
Mountain ash grows as a shrub or small tree 10 to 30 feet tall with an average d.b.h. of 10-25 cm.
The trunk is slender and short with spreading trunks and branches, growing to a narrow rounded crown
[figure 7-10]. If it grows in a closed canopy system the trunk grow longer, splitting higher along the tree
to maximize sunlight exposure. The trunk, crown, and fibrous root system are very slow growing and
short lived (Sullivan, 1992).
Numerous birds eat the mountain ash berries, and this contributes to the main method of seed
dispersal. Flowering occurs from May through July, and the fruit ripens in August. The berries remain on
the tree throughout the winter. The distances seeds are dispersed are usually that of a few hundred feet
and germinate best in paper birch-spruce-fir cover (Sullivan, 1992). Rocky hillsides or thin soils atop the
Catskill Mountains provide the best habitat, and is very common in canopy openings. It grows well in
poorly developed shallow spodosols and inceptisols where the climate is cool, windy, and humid. These
features are common between Catskill summits and their tree constituents. If mature or semi mature trees
are scattered in a coniferous forest, mountain ash is able to establish itself well as codominant small tree.
Mountain ash is identified as a facultative seral species (Sullivan, 1992). It is shade intolerant, and
it is more abundant in early seral spruce-fir communities where a disturbance has set back succession and
less abundant in old growth spruce-fir communities. Along with mountain paper birch, mountain ash is a
factor in the transition from spruce-fir forest to a hardwood forest through canopy opening colonization
and deciduous duff accumulation.

Mountain Maple Acer spicatum [figure 7-11]- Mountain maples native range extends through
southeastern Canada and the northeastern United
States, and in mountainous areas throughout North
Carolina and eastern Tennessee. It is a member of the
spruce-fir ecosystem in commonly recognized plant
associations (Sullivan, 1992). It occupies a niche in the
understory sub canopy layer, scattered throughout the
mature sub-climax forests and will occasionally form a
dense shrub layer in a very disturbed forest. It grows
opportunistically where red spruce and balsam fir are
removed from the canopy layers.
Maximum height can be up to 33 feet and is
usually smaller, around 20 feet, with a maximum d.b.h
7-11: Multiple, mature mountain maples grow on the
slopes around Black Domes summit, with occasional
seedlings and saplings on the summit.
27
of 15-20 cm. The trunk often grows short and crooked with a clumped bushy growth form. It grows a
shallow root system, with the majority of the roots located close to the soil surface. When the plant is
between 5 and 10 years of age it can reach maximum growth rates of about 1 foot per year, becoming
very slow growing once it reaches 40 to 50 years (Sullivan, 1992).
Through insect pollination the seeds are distributed by wind and require stratification throughout
the ground layer for best germination. In undisturbed soils where there has been no upturn or disturbance
the seeds will germinate at the highest rates. Once established, it can tolerate deep shade but grows best in
sun, using its bushy layered growth form to take advantage of periodic sunlight penetration through the
upper canopy (Sullivan, 1992).
In the spruce-fir community mountain maple grows in scattered groups in the understory sub-
climax community. It does not slowly invade as leaf litter accumulates; rather it is a feature of the
understory small tree or shrub layer. If it grows to a significant proportion it can suppress spruce or fir
seedlings, but ultimately it will give way to conifers, as it is short lived.

7b. Understory Shrub & Herbaceous Layer
Clintons Lily Clintonia borealis [figure 7-12]- Native to
North America, this perennial grows at a moderate rate
during the spring from older rhizomes. Although it is
characteristic of shady coniferous forests, it is full shade
intolerant. Clintons lily spreads extremely slowly, mostly
by vegetative clones through rhizomes. It can reproduce
through seeds via its small berries, but it takes up to 10
years for the perennial clone to establish itself to the
point where it can successfully produce seeds
(USDA/NRCS). The seedlings have low vigor and are
usually not successful in establishing a new clone
colony. At maturity, it will grow to a maximum height
of just over one foot. The long period of time required for a colony to establish dominance indicates its
sensitivity towards a site disturbance, such as upheaval of a nearby tree or smothering by fallen organic
debris during an ice or wind storm.

7-12: An established Clintons Lily colony. This species
is intolerant of soil disturbances, indicating that its
position on Hunter Mountain is stable. Hunter Mountain
has had patches cleared for construction in the past, but
the presence of this species shows that the larger forest
ecosystem is intact and healthy.
28
Goldthread Coptis groenlandica [figure 7-13]-
Goldthread can be found throughout coniferous
forests of North America in cool, moist climates,
growing best in moderately drained spodosols
(Sullivan, 1992). It dominates the understory strata in
nutrient-poor areas or sites with abundant mor-humus,
characteristic of the micro site directly underneath
conifer trees. The damp, limited sunlight, mossy sites
where it can be observed atop the Catskill Mountains
are characteristic of its growing sites throughout its
entire range. Because of its preference for moist sites it is full sun intolerant, but it does require some sun.
Consequently, when a canopy disturbance occur,s it is intolerant of the new open conditions and is unable
to regrow (Sullivan, 1992).
Goldthread is an evergreen hemicryptophyte, growing from a creeping rhizome beneath the
surface as a perennial. It does not have a main stem, but rather grows layered branches very close to the
ground (Sullivan, 1992).

Mountain Wood Sorrel Oxalis Montana [figure 7-14] - Wood sorrel is a dominant understory species
throughout the eastern North American coniferous
mountain forests, primarily in spruce-fir forests. It
grows perennially in well-developed colonies comprised
of vegetative clones as geophytes, with its main point of
growth coming from beneath the ground (Pavek, 1992).
Because of its wide range and tolerance for an array of
site conditions, wood sorrel is not attributed with a
particular set of site characteristics. However, its habitat
in the Catskill Mountains is usually under a mature balsam
fir canopy. This implies a shady area with a thin organic
layer lacking a smothering annual leaf litter. The soil under
a pure balsam fir stand often has a lower water retention capacity, and this can affect the growing zones of
mountain wood sorrel that prefer mature balsam fir stands.
7-13: A very dense Goldthread colony in a moist, mossy site.
7-14: Wood sorrel grows close together but not in
dense clusters. Here it is seen on the moss layer,
indicating its preference for moister sites.
29

Starflower Trientalis borealis [figure 7-15]- Throughout eastern North America, starflowers may be
found in cool, moist, humus rich soils. It grows to a maximum
height of under one foot as a perennial and is most commonly
spread through seeds (USDA/NRCS). Its characteristic flower
consists of 7 white pedals, and colonies will spread to form a
mat on the forest floor. When observed in the Catskills it is
typically found growing out of dense moss mats. It is able to
grow in mixed high elevation forests where deciduous trees
contribute to the leaf litter and establish mature stands.
Starflower then has a higher tolerance for germination through
deeper leaf litter, but its primary habitat in the Catskill forest is in dense coniferous forests.

Hobblebush Viburnum lantonoides [figure 7-16]- Organic rich, moist forests in higher elevation
northeastern North America are the habitat for hobblebush. It
grows best in acidic soils with a mixed soil medium. Growing
up to 12 feet high, it tends to sprawl over wide distances by
growing pendulous outer branches (USDA/NRCS). These
branches may root once they make contact with the ground,
further establishing the colony. This tangle is a typical image
of the higher elevation Catskill forests where navigation in a
straight line becomes cumbersome, leading to its name. Fall
foliage is a brilliant red with fruits attracting birds.

7c. Mosses
A major characteristic of the high
elevation Catskill forest is a dense moss mat
7-15: Starflower grows mixed with other
species on the ground stratum.
7-16: Hobblebush creates thick spreads in the
forest. Its growth is associated the growth of
other high elevation understory species, and does
not inhibit their development.
7-17: The organic matter that this moss is growing from is
visible in the upper portion of this picture. Organic debris,
mainly pine needles, is visible, tangled underneath the moss
and scattered lichen. This eventually becomes part of the
humus, furthering the growth of this small area.
30
covering the forest floor [figure 7-17]. These mats usually cover boulders, fallen logs, and grow up the
trunks of living trees [figure 7-18]. Upon close inspection, multiple species of moss grow over one
another and usually prefer one micro site. The life forms these different species will take serve a different
purpose that can be indicative of what kind of microenvironment it must contend with throughout its life
cycle. The growth forms ultimately serve a common purpose, which is water retention (Glime, 2010).
Reducing air exposure, thickening the boundary
layers vegetative thickness, and growing on top of
one another to protect each other can achieve the
water retention needed to sustain the moss mat.
Water retention and availability of moisture
is the primary determining factor in the distribution
of mosses. The boundaries for each species are
ultimately set by temperature, but growth within
that boundary is limited by availability of water.
The ecosystem where a moss mat grows is altered
through the retention of water (Glime, 2010). The
soil temperature under moss is reduced, which then
slows the rate of decomposition and nutrient cycling rate.
While nutrient cycling is slowed, mosses also tend to retain
nutrients absorbed from the soil within their younger tissue
for future use. Spruce-fir forests in the Catskills thrive
where there is a thick moss layer, and the slow growing
spruce roots are able to survive well under the moss. This
suggests that the nutrients that mosses retain are not in the
soil but are stored in a way that is available to feeding
roots (Glime, 2010).
Mosses play an important role in the Catskill
high elevation forest community. Spruce and fir
seedlings can often be observed growing directly from the moss mat [figure 7-19], benefiting from the
nutrient and water retention. Mosss tendency to store nutrients and transport those nutrients throughout
the living tissue depending on moisture availability indicates that their biomass is an important feature
when weighing the nutrient availability and fertility of the soil.
7-18: This lump of different moss species on a log collectively
retains moisture and builds organics. Wood sorrel is growing
from the organics that have accumulated. Moss contributes its
properties to help cycle, maintain, and grow the forest floor.
7-19: Balsam fir seedlings growing towards
establishment. Organics accumulated on the moss to the
point where the seeds could germinate, and now the
retained soil moisture and available, retained nutrients aid
the seedlings.
31

7d. Lichens
Lichens are a symbiotic life form between fungal and algal
cells [figure 7-20]. They grow spreading colonies on bare
substrate or an organic surface such as tree bark or a log,
dissolving and drawing out nutrients and furthering the
chemical and physical weathering processes of mineral
layers [figure 7-21].

The distribution of lichen species and forms
throughout the Catskill forest depends on extreme
microclimate conditions that do not affect other species.
The texture of bark, temperature, moisture, acidity, and
availability of bare substrate are factors that determine
what sites are best suited for a specific variety of lichen
(Larson, 1980).
These life forms developed in response to the micro
site and what kind of substrate the lichen is feeding from.
The internal morphology however, is consistently similar
across all species, where the bulk of the tissue is a fungal
component of the lichen colony [figure 7-22] (Waggoner). The
cortex is where the lichen colony comes in direct contact with
the outside environment. Here, the filaments are packed tightly
to keep other organisms and to minimize the inner filaments
direct exposure to sunlight, which will damage algal cells.
Below the cortex the fungal filaments lose their
density. The algal partner cells are then able to
live and distribute themselves just below the
cortex in the symbiont layer, in an arrangement similar to leaf cells in order to maximize photosynthetic
production. Below that algal layer is the medulla layer, where fungal filaments are tightly woven and
attached to the underlying substrate. Enzymes are emitted to break down the substrate, or if the lichen is
growing on organic matter it is using the host as a bed for photosynthesis.
7-20: Various species and growth forms of lichen
growing atop a sandstone cliff on Hunter Mountain.
7-21: The dark, loosely attached lichen shown here is
not the only lichen in this picture. The discoloration
on this rock is actually complete lichen cover.
Enzymes are released that slowly chemically erode
the rock, contributing to the breakdown process of
the Catskills.
7-22: Internal lichen morphology. Source: Waggoner, B. with
Berkeley University.

32
In boreal ecosystems, which include the coniferous forests of the high elevation Catskills, lichen is
an important factor in nutrient and biomass accumulation. Metabolism in lichen is opportunistic, and
whine moist climatic conditions are favorable lichen will absorb water, becoming dense and sponge like,
carrying out its metabolic processes for as long as possible and returning to a state of inactivity (Larson,
1980).

7e. Ferns
The dominant variety of fern found
throughout the high elevation Catskill forest is
mountain wood fern. They reproduce both by
spores and vegetative cloning through growing
male, female, and bisexual plants. Spores may be
dispersed by wind collect in great quantity in an
area where ferns are absent, colonizing a new area.
Vegetative reproduction through rhizomes is more
common, showing the importance for a colony to
become well established before it is able to spread
(USDA/NRCS). Mountain woodferns grow in a
very wide variety of physiographic sites, but in the
Catskill Mountains they are commonly found in
cool moist woodlands, both in shade and sun. They
thrive in the acidic, shallow, rocky, organic rich soils found throughout the mountains, with a strong
affinity for moist sites. Studies throughout the range of woodferns have mixed conclusions as to what
triggers their spread and eventual understory dominance in a mature forest (USDA/NRCS). In the Catskill
Mountains, mountain wood fern grows as a member of the mature forest community. There are many
areas known as fern glades, where wooderns grow to their maximum and cover dense patches of the
forest floor [figure 7-23] (Kudish, 1971). The presence of ferns does not indicate a disturbance or seral
stage. Instead, they are a very well developed fern colony that has out competed other members of the
understory strata, and have colonized the area independently of a facultative disturbance event.


7-23: Dense understory of fern under mature balsam fir
growth, on the first slopes below the summit of Thomas Cole
Mountain. The codominant understory species in this stand is
Clintons Lily, which is intolerant of site disturbances.
Because of the presence of a disturbance-intolerant species and
mature climax community tree species, it can be concluded
that this site is in its mature form. While mountain woodfern
was sparse in the high elevation samples besides Thomas
Cole, it is a component of the mature high elevation Catskill
forest.
33
8. Catskill High Elevation Forest Community Discussion
The high elevation Catskill forest is an ecosystem undergoing successional progression during the
present post-glacial period. The sedimentary bedrock, exceptionally weathered and carved during
complex periods of glaciations, has been broken down into mineral soil through physical and chemical
erosion. Derived nutrients fixed by early successional species and the decayed biomass have been cycled
to create the organics comprising the soil. This process began once glaciers retreated, and the few alpine
glaciers remaining kept the upper reaches of mountainous regions out of reach to the pioneer species
struggling to establish the new forest.
The process of an organic layer being thickened over continuously weathering mineral soil and
bedrock is ongoing, present as the high elevation spruce-fir and balsam fir forest assemblage of the
greater Catskill forest ecosystem. While many lower elevation shelved escarpments share the same
characteristic thin organic layer and bedrock weathering, these are topographical features within bounds
of the hardwood forest community. Slide, Hunter, and the Blackhead Range are the upper reaches for the
forest migration upwards. The forest analyzed on the five highest individual mountain elevations of the
Catskill Mountains is a representation of the biodiversity that is produced and adapted to the physical
limitations on survival currently present. It also provides insight into the dynamic forest succession of the
region. While it is true that the mature conifer forest is the climax community for its set of physical
limitations, the accumulation of organic matter alters the duff and soil composition to the point where
different tree communities have a chance to contend with the conifers.
Because the dominant tree species are balsam fir and red spruce in the region of this studys scope,
the forest is ecologically defined as a red spruce- balsam fir forest. The ecological definition for the region
of balsam fir forest is not appropriate. Even though Slide and other mountains are completely dominated
by balsam fir they are still a spruce-fir ecosystem. This is because the domination by balsam fir over red
spruce is a part of their seral dynamic. There are Canadian boreal forests dominated by balsam fir that are
a unique ecosystem apart from spruce, but in the Catskill mountains it is possible, through a set of
disturbances and seed dispersions, for conifer stands of either balsam fir or red spruce to be completely
replaced by one another. Several documentations by 19
th
century naturalists and early surveyor remarks of
the Catskill Mountains mention forest communities in local regions that are not present today (McIntosh,
1964). This shows how the forest is changing with every tree generation that becomes established under a
certain set of ecological circumstances that are altered as disturbances and organic accumulations
progress.
34
The field study conducted provides an insight into how this process is undergoing and what
environmental processes are at work behind the evolving forest succession. The over story red spruce-
balsam fir canopy is the catalyst and determining factor for the vegetational processes in the middle and
understory. Shade and the organics deposited by tree biomass sustain a large part of the environment
needed to secure the habitat for herbaceous and ground story vegetation. The two primary canopy trees,
balsam fir and red spruce, are in competition with one another for stand dominance. Where mature red
spruce dominates, it has taken advantage of a canopy opening after possibly having been suppressed by
shade or lack of space. Its fast growth once it achieved full sun enabled it to out grow and shade the
balsam fir around it, which will then not be able to enter its reproductive flowering stage.
Conversely, where mature balsam fir dominates a stand which is the case on Slide Mountain, the
balsam fir has been able to take advantage of the harsh environment that red spruce has failed to well
establish itself in. Balsam fir grows faster, and its roots grow deeper than red spruce. Deeper root growth
allows balsam fir to reach the top layer of mineral soil and spread farther than red spruce to find soil
moisture, and it is also able to tolerate much drier soil conditions than red spruce. Slide mountain was the
most recent to be exposed during deglaciation, and its thin soils provide a condition where balsam fir is
the only tree species present on the highest elevation. Mountain ash, mountain maple, paper birch, and
occasional lower elevation yellow birch grow along its slopes, and as the organic layer accumulates these
may migrate higher. Slide is also recovering from human disturbance on its highest elevation, but this
disturbance does not appear to have eliminated other tree species or have had a major alteration in forest
succession.
Disturbances will occur and the domination of one species over the other is temporary, as is the
presence of a strict spruce-fir forest. Paper birch will commonly move into disturbed areas and canopy
gaps. The summit of Hunter Mountain is a good example of this process, where the open young red
spruce canopy is in an area repeatedly cleared and disturbed by human activity. Paper birch dominates the
slopes and has high species frequency on the summit. Its shallow roots and fast growing saplings have
been able to colonize the area, and in time it will be seen if they will further open the area to existing
patches of mountain ash or mountain maple through soil stabilization. Alternatively, the exceptionally
dense balsam fir saplings in the area may mature and eliminate the chance for paper birch to spread.
These variables must be weighed and monitored if the complete process is to be understood.
The Blackhead range is an example of the Catskill spruce-fir forest in the mixed high elevation
hardwood stage discussed earlier, and is also a product of natural disturbance events, not anthropogenic.
The individual micro sites and dominances vary, but the three can be classed into one general category of
35
the Catskill high elevation forest succession. They each share the characteristic where paper birch,
mountain ash, and mountain maple are emergent lower canopy small trees. This occurs because of the
buildup of organics allowing the deeper root systems of deciduous trees to take hold, and their leaf litter
may contribute to the decline in conifer seedling vigor. This fate is not sealed, as a deciduous tree located
in this area may be an opportunist and will not contribute to any sort of masting event or slow takeover of
one particular species.
Beneath the over story canopy on these peaks is the understory herbaceous layer. The species
described previously are not restricted to this area because of an affinity for a particular conifer cover or
narrow edaphic tolerance. It is their seedling and germination requirements that result in their dominance
under spruce-fir cover. In lower elevation areas dominated by eastern hemlock and red spruce, the same
understory community is present. Their habitat seems to be that of coniferous cover. This means they are
best adapted to grow in an area consisting of partial sun, acidic soils, and non-smothering leaf litter. Their
absence in the hardwood forest surrounding low elevation coniferous areas suggests that the full sun in
spring before leaf development and the heavy leaf litter in the autumn prevents their growth while other
species well adapted to hardwood forest conditions dominate. Thus, the distribution of the dominant
herbaceous and understory species in Catskill coniferous forests is a function of what conditions are
created by the over story species that create shade and cycle organic matter.
It is appropriate to compare the high elevation conifer forest to a closed canopy boreal forest.
Both the Rocky Mountains of the west and the Appalachian mountain chains throughout the east contain
species of over and under story strata that are present throughout the Canadian closed canopy boreal
forest. The disturbance regimes in the Catskill Mountains are different; the absence of fire in the Catskills
creates a different succession pattern. Throughout the dense, closed canopy sections of the vast boreal
forests, a periodic fire every two to four hundred years will reset the forest to a species best suited for fire
reproduction, such as jack pine (Larson, 1980). This model of succession outlines that as the levels of
sunlight reaching the forest floor decrease when the trees grow older, each new generation is of a species
that can tolerate increasingly lower levels of sunlight through its germination and establishment phases. In
the Catskills, the forest is not reverted back to one species, but the forest ecosystem evolves to facilitate
the tolerances of different tree populations. Lower elevation hardwood forests have replaced all of the
spruce-fir that was once present in the lower elevations.
The main similarity between boreal forests and the Catskill forests that serves to characterize the
Catskill forest as partially boreal is the post-glacial revegetation dynamic. In the Bearing Straight, areas
experiencing glacial retreat and post-glacial vegetation development follow a pattern that can be
36
extrapolated into the Catskills during the close of the Pleistocene. The bare substrate that is left in the
wake of glacial retreat is weathered to the point where a rugged plant is able to establish itself. This
pioneer species eventually dies, and its organics, along with any dead insects, blown particles, and
continuously weathered rocks collect to form the first layer of organics and minerals that become the
future soils. Nitrogen fixing shrubs can then establish themselves in the organics, and their leaf litter
contributes to the soil. Peat mats accumulate, and the soil becomes suitable for small trees, then conifers
to colonize the area and grow to maturity. This population will then become a mature boreal forest,
subject to the disturbance regimes of the boreal ecosystem (Allen, 2010). It is likely that a closely similar
process occurred in the Catskill Mountains to create the forest present today. Pollen records in bogs
throughout the Catskill Mountains show that a conifer forest once existed at lower elevations that
resemble boreal conifer forests (Kudish, 1971). As organics accumulated, this area was able to support
hardwood species that have been spreading up the mountains ever since in a manner similar to that of
closed canopy boreal forests. At this point, the young infertile soils that the spruce-fir forest was able to
grow in have become more nutritive. Using this perspective, the high elevation spruce-fir forests are the
last refuge for these species and their intact ecosystem in the Catskill Mountains. High ledges that do not
accumulate soil well and heavy orographic precipitation zones may retain a coniferous assemblage
because of their harsh environment, but it is likely that throughout oncoming centuries hardwood
populations will encroach higher into the Catskill Mountains.

9. Conclusions
Geological, glacial, and vegetative succession histories intertwine in the Catskill Mountains to
explain the prevailing attributes of the highest elevation forest assemblage. The most basic form of the
region, the bedrock itself, is a link to an era where trees were first becoming dominant factors in the
biosphere. Fluctuating shorelines throughout a wedge of clastic material, protruding as a delta from the
eroding, towering Acadian Mountains, set the foundation for the current forest communities to develop
upon.
Over the vast time between the Devonian Catskill formation and the present, life on earth has
evolved with its changing environmental conditions. Genetic traits, successfully adapted to the physical
limitations on life within a region, live on as the species that are able to establish their dominance in a
community, becoming a defining and fundamental element of their ecosystem. Each species within the
community develops a niche, and when environmental conditions are favorable to their growth, survival,
and reproduction, the species composition of the community will change.
37
In the Catskill forest, the processes of environmental change and species composition change
intertwine, both affecting each other. As global temperatures increased, the last continental ice sheets
withdrew from the Catskill region, leaving a barren landscape to be colonized by new vegetation
following post-glacial, closed-canopy boreal primary succession. The longer growing season in the
Catskills, however, facilitated the establishment, reproduction, and subsequent spread of mid-continental
hardwood species. These follow the Catskill boreal forest remnants wake of organic matter accumulation
and soil development, replacing the conifers up the mountain slopes. Openings in high elevation summit
forests and the immediate slopes below summits host early facultative deciduous species, the pioneers for
the next high elevation community in areas that will support developing soils.
The Catskill forest is in a succession continuum. To define the spruce-fir forest as the ultimate
climax forest on the summits is assuming that there will be no changes. They are a climax community in
the sense that within their given ecological tolerances and limitations, balsam fir, red spruce, and their
associated dominant understory are the mature, old growth without significant disturbance. However, in
the perspective of natural history, the mature forests of the present are transitionary glimpses into the
processes of biodiversification.
Biogeography of the Catskill high elevation forests opens analysis to an ancient landscape.
Distribution of the dominant species is dependent on whether or not the seeds can germinate, and as
organic debris accumulates, the shallow rooted conifers create the conditions for their demise. As the past
glaciations show, these scenarios are not permanent, and future changes in climate will alter the
conditions in which life must survive. To look even further, eventually the mountains will have eroded to
the point where only small hills remain, washing back into the sand which once formed the once great
plateau. The high elevation Catskill forest may be spruce-fir today, but natural history tells us that new
worlds and their biospheres will manifest themselves. At that point, biogeography becomes fiction.









38
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