Forests: Temperate Evergreen and Deciduous

Abstract

Temperate forests represent one of the major biomes on Earth. They are most common in eastern North
America, western and central Europe, and northeastern Asia, where the climate is defi ned by warm
summers, cold winters, and intermediate levels of precipitation. To a lesser extent, they are also present
in this same climate in Australia, New Zealand, South America, and South Africa. Temperate forests are
dominated by either deciduous or evergreen canopies. In temperate deciduous forests, more commonly
found in the Northern Hemisphere, plants drop their leaves in autumn, allowing for high seasonal variation
in light availability to the understory. By contrast, in temperate evergreen forests, which are more commonly
found in the Southern Hemisphere, plants keep their leaves year round. The temperate forest biome is
rich in geologic and anthropocentric history, with much of the land shaped by glacial and human activity.
Located in some of the most industrialized places in the world, temperate forests have been instrumental to
socioeconomic development in these regions. As a consequence, these forests have been heavily impacted
by expanding human populations, which remain the primary continuous threat to the structure and function
of these forests.

Full text

Encyclopedia of Natural Resources DOI: 10.1081/E-ENRL-120047447
214 Copyright © 2014 by Taylor & Francis. All rights reserved.
Ecotone—
Fragmentation
Forests: Temperate Evergreen and Deciduous
Lindsay M. Dreiss
John C. Volin
Department of Natural Resources and the Environment, University of Connecticut, Storrs, Connecticut, U.S.A.
Abstract
Temperate forests represent one of the major biomes on Earth. They are most common in eastern North
America, western and central Europe, and northeastern Asia, where the climate is defi ned by warm
summers, cold winters, and intermediate levels of precipitation. To a lesser extent, they are also present
in this same climate in Australia, New Zealand, South America, and South Africa. Temperate forests are
dominated by either deciduous or evergreen canopies. In temperate deciduous forests, more commonly
found in the Northern Hemisphere, plants drop their leaves in autumn, allowing for high seasonal variation
in light availability to the understory. By contrast, in temperate evergreen forests, which are more commonly
found in the Southern Hemisphere, plants keep their leaves year round. The temperate forest biome is
rich in geologic and anthropocentric history, with much of the land shaped by glacial and human activity.
Located in some of the most industrialized places in the world, temperate forests have been instrumental to
socioeconomic development in these regions. As a consequence, these forests have been heavily impacted
by expanding human populations, which remain the primary continuous threat to the structure and function
of these forests.
INTRODUCTION
Temperate forests represent one of the major biomes on
Earth, covering ~14% of Earth’s terrestrial land surface[1]
(Fig. 1). Along with boreal and subtropical/tropical wet
and dry forests, temperate forests are one of the Earth’s
dominant forest types. In comparison to other types, tem-
perate forests are intermediate in latitude, temperature, and
precipitation. By contrast, they generally exhibit much
stronger seasonality, favoring those plant and animal spe-
cies that were able to adapt to short-term climatic variation.
Environmental stresses from seasonal change include
extreme temperatures and variable access to moisture (in
the form of rain and/or snow), light, and nutrients (due to
the deciduous nature of some canopies). As a result, sig-
nifi cant changes in microclimates and growing conditions
throughout the year are common, creating adaptive oppor-
tunities and resulting in unique structure and composition
of ecosystems.[2–4]
Temperate forests are located in some of the most heav-
ily populated and developed regions on Earth, including
much of eastern North America, western and central
Europe, northeastern Asia and, to a lesser extent, Australia,
New Zealand, South America, and South Africa[5–7] (Fig. 1).
Temperate forests are an established part of the history and
culture of these regions and continue to play an important
role in providing ecosystem services to human popula-
tions, including timber products, carbon storage, clean
drinking water, erosion prevention, recreation, tourism,
aesthetics, property value security, and others. On the other
hand, the unsustainable use of such services has led to
negative consequences for temperate forest ecosystems
worldwide.
In this entry, we provide an overview of temperate for-
ests from both an ecological and societal perspective. We
describe the unique characteristics of temperate deciduous
and evergreen forests, the major anthropogenic agents that
have impacted them in the past and some of the emerging
threats they face today.
CLIMATE
Climate is a fundamental factor in defi ning the temperate
forest biome. Generally, a forest is considered temperate
if it is located in a region with hot summers and cold win-
ters. The annual range in temperature can be ±30°C, and
the mean annual temperature is between 3 and 18°C, with
mean midwinter temperature below 8°C and mean mid-
summer temperature about 18°C.[8] In the Northern Hemi-
sphere, the length of the frost-free period ranges from 120
to over 250 days, with growing season and temperature
increasing closer to the equator. As a result of less land
mass in the Southern Hemisphere, the climate is milder
than at similar latitudes in the Northern Hemisphere. Tem-
peratures in the Southern Hemisphere can be cold, but
typically far less than 1% of the hours of the year are

Forests: Temperate Evergreen and Deciduous 215
Ecotone—
Fragmentation
subject to frost, while in the Northern Hemisphere
subfreezing temperatures are typically reached many days
during the winter.[6]
Temperate forests receive even precipitation year round
averaging 750–1500 mm/yr. Snowfall can range from non-
existent in southern regions to extremely heavy in northern
regions. In the Northern Hemisphere, temperate forests are
geographically located between the boreal forest/tundra
and the tropical/subtropical forests while in the Southern
Hemisphere they are found south of the tropical/subtropi-
cal forests up to tree line in some areas (Fig. 1). Corre-
spondingly, the climate, productivity and species diversity
of temperate forests are intermediate in relation to other
biomes.[9]
The seasonal climatic regime in the temperate forest
regions has changed very little in the past 1000–2000 years.
This is refl ected in the structure and ecosystem dynamics
of the forests.[10] Winter deciduous trees or coniferous
evergreen species that are physiologically dormant during
winter months dominate the majority of the forests. Genera
differ between Northern and Southern hemispheres.
Examples include Acer, Fagus, Tilia, Quercus, Carya,
Populus, Ulnus, Betula, Fraxinus, Magnolia, Cornus,
Robinia, and Juglans in the Northern Hemisphere and
Eucalyptus, Acacia, Quercus, and Nothofagus in the
Southern Hemisphere.
At higher latitudes and altitudes, where temperatures
range from –30 to 20°C annually, temperate forests become
more dominated by coniferous evergreen tree species.
These regions are generally drier, with annual precipitation
from 300 to 900 mm. Seasons are defi ned by cold, long
snowy winters and warm, humid summers with at least
four to six frost-free months.[11] Evergreen temperate for-
ests may also occur in regions with a stronger maritime
climatic infl uence. Precipitation can be very high (typically
700–1000 mm) and snow is uncommon.[12,13] Summer fog
is often an important contributor to moisture uptake, reduc-
ing stresses from evapo-transpiration. In addition, the
growing season is much longer and sometimes continuous
in forests closer to the equator. In the midst of global
climate change, however, a shift is occurring in the location
of ecotone and temperate forest species.
Climate-linked range shifts have been observed in the
northern hardwood–boreal forest ecotone where there has
been a decrease in boreal and an increase in northern hard-
wood basal area.[14] The velocity of temperature change is
projected to be much faster in temperate broadleaf and
mixed forests than in temperate coniferous forests.[15]
Temperature being considered the primary control on
treeline formation and maintenance, global warming is also
facilitating treeline advances to greater altitudes and
latitudes.[16,17] The synergistic effects of warming tempera-
tures and intensifying droughts have also triggered con-
cerns. The sensitivity of tree mortality and range shifts can
result from extreme water stress and lead to other ecologi-
cal consequences including changes in carbon stores and
dynamics, changes in microclimate, and changes in future
production of important habitat and food sources.[18,19]
LAND FORMATION AND HISTORY
On the geological time scale, climate has shaped the land
on which temperate forests now grow. Between 110,000
Fig. 1 Global temperate deciduous and evergreen forest biomes.

216 Forests: Temperate Evergreen and Deciduous
Ecotone—
Fragmentation
and 10,000 years ago, during the Late Pleistocene age, gla-
cial ice sheets left large parts of the Eurasian and North
American continents as treeless, frozen tundra.[20,21] Below
the tundra stretched the temperate forests, and beyond
these, temperate, grassy plains. As the glaciers retreated,
they left behind glacial till and lake-bottom sediments. As
the zones of tundra followed the shrinking glaciers, the
boreal and temperate forests encroached on the tundra belt
until the matrix of ecoregions looked much like it does
today. Since the end of the last ice age, human activity has
become a primary factor in shaping temperate forests.
The three largest temperate forest regions, eastern North
America, western and central Europe, and northeastern
Asia, have played an important role in the development of
human society. In North America, temperate forests were
heavily impacted by the spread of European settlements in
the 1700s. Much of the temperate forest was cleared,
logged, and/or burned to make way for agricultural needs
and human settlements.[22,23] In the 18th century, the temper-
ate forests of Europe were the birthplace of new lines of
scientifi c study and management, becoming what is now
modern forestry.[24,25] These practices were applied in
Europe and North America, where temperate forests pro-
vided the wood for fuel, building materials, and raw mate-
rial in industrial processing during the Industrial Revolution
of the 19th and 20th centuries.[26,27] With the spread and
development of forestry practices, large amounts of tem-
perate forest were logged for the production of timber,
pulp, paper goods, and other forest products. During the
19th and into the 20th century, much of the agricultural land
in the temperate forest range in North America was aban-
doned and the temperate forest grew back, peaking in its
coverage in the mid-twentieth century.[28] Today it is once
again under threat from continuous loss and fragmentation
primarily through human developments.
In Europe, many native temperate deciduous forests
have been extirpated and replaced with planted monocul-
tures, many of which are not representative of the native
vegetation.[29] Most of the remaining European temperate
forests display a simplifi ed structure and composition
resulting from centuries of tree harvesting and forest graz-
ing. In other places, land use change and conversion still
continue to affect forest ecosystems. For example, the tem-
perate forests of Asia, which cover parts of China, Japan,
and Korea, were largely unaffected by glaciation through
the Pleistocene.[30,31] Because of this, Asian temperate for-
ests have higher levels of biodiversity than their counter-
parts in Europe or North America and are home to species
with ancestries that extend much further back in geologic
history. However, more recent heavy logging and fi res have
reduced forested land area and caused species such as the
red-crowned crane (Grus japonensis) and the red panda
(Ailurus fulgens) to be listed as endangered.[32] Collectively,
temperate forests across the globe may have experienced
more severe impacts over broader areas than any other
large forest biome.[33]
STRUCTURE AND COMPOSITION
In contrast to grasslands or desert ecosystems, temperate
forests have a more complex structure composed of different
layers of vegetation. Temperate forest layers typically
include a layer of mosses and lichens on the forest fl oor (and
often on tree trunks and limbs), herbaceous and shrub lay-
ers, a subcanopy of trees, and a taller dominant canopy tree
layer.[34] The complexity of the subcanopy and canopy layers
can vary and often depend on the light requirements of the
dominant tree species. Trees are the dominant life form of
these ecosystems and can be either deciduous or evergreen.
Temperate forests undergo different stages in ecosystem
succession, allowing for the coexistence of plant species
with different life history strategies. These adaptations are
often linked to specifi c growing conditions associated with
different stages of forest growth. As a result, gradual transi-
tions in plant community composition and forest structure
change over time.[35,36] A classic example of succession in
northeastern temperate forests in North America is when
pioneer tree/shrub species such as birch (Betula spp.) and
poplar (Populus spp.), which are fast growing and require
high light availability, are often the fi rst to establish follow-
ing a disturbance. As trees grow and the canopy begins to
close, conditions become more favorable for more long-
lived, shade-tolerant species such as maple (Acer spp.), oak
(Quercus spp.), beech (Fagus spp.), and hemlock (Tsuga
spp.). These later-successional trees become dominant or
codominant and eventually out-compete early successional
species, reaching heights of 35–40 m.[37] This process takes
place over hundreds of years and can be reset at any time
by a disturbance of natural or human origin. Indeed, within
a relatively large and healthy naturally occurring forest
area, all of these successional stages are present at once.
Disturbance regimes, which refer to the type, intensity,
and frequency of disturbances in a particular region, have a
substantial impact on the composition and age structure of
vegetation.[38–40] In temperate forests, eolic events such as
hurricanes, tornadoes, and thunderstorm downbursts are a
major contributor to forest dynamics. Fire was once also an
important natural regulator, but with land use changes and
suppression of fi re by humans, many temperate forests
have become increasingly dominated by late successional
species.[41] Other natural disturbances include pest/disease
outbreaks, fl ood or drought, and snow and landslides.
Temperate Deciduous Forest
The temperate deciduous forest canopy is composed mostly
of broadleaved angiosperm tree species with the inclusion of
some coniferous tree species. Temperate deciduous broad-
leaved forests are often defi ned by the associations of spe-
cies. Association names are most useful for distinguishing
broad differences in forest type and are often representative
of variation in soils, topography, and climate. For example,
in eastern North America, some common associations are

Forests: Temperate Evergreen and Deciduous 217
Ecotone—
Fragmentation
heating/cooling). However, with the onset of global climate
change, recent studies recognize that temperature plays a
more important role in many phenological events.[48,49]
Plant hormonal signals to leaves cause chlorophyll, the
compound that absorbs light for photosynthesis and makes
the leaves look green, to degrade. Some nutrients from
the leaves are partially recovered and moved (i.e.,
retranslocated) to stem and trunk tissues before the leaves
fall off.[50–53] Buds are formed and go into dormancy until
the following spring, when the accumulation of degree
days over a temperature threshold triggers leaf growth.
Canopy tree species differ in their timing of bud break and
leaf expansion, with the development of the forest canopy
taking place over a period of a month or more.[54] Once
intact canopies are in full leaf fl ush and the growing season
is underway, typically only 1–5% of sunlight reaches the
forest fl oor.[55–57] Therefore, the portion of the year when
canopy trees are leafl ess creates a unique opportunity of
increased light availability for understory vegetation.
With the beginning of the warm season, herbaceous
plants, shrubs and small trees begin growth before the can-
opy does. Because maximum incident solar radiation fl ux
and zenith angle of the sun are larger in the spring months,
there is greater transmission of light and a greater likeli-
hood for understory plants to reach photosynthetic capacity
before canopy leaf fl ushing.[58,59] In an evolutionary adapta-
tion designed to maximize the amount of light received,
some plants, known as spring ephemerals, complete most
of their annual growth cycle in the few weeks between
snowmelt and the closure of the tree canopy. Other more
shade-tolerant species begin or continue to grow after can-
opy closure.[60]
Animal life
Many animals in temperate deciduous forests are mast-
eaters (nuts and acorns) or omnivores. Herbivores (e.g.,
deer, hare, and rodents) perform important tasks in the eco-
system, such as aiding in plant seed dispersal, but may
cause stress when their populations are overly abundant.
Small rodents, such as squirrels and chipmunks (Tamias
spp.), and some birds, such as blue jays (Cyanocitta cris-
tata), cache large seeds/nuts in the autumn for sustenance
in the winter.[61] Seeds that animals fail to recover may
grow in their buried location. Omnivores include raccoon,
opossum, skunk, fox, and black bear in North America and
badger, mink, and marten in Europe. Asian forest fauna
also include primates such as macaques. In temperate for-
ests around the world, humans have drastically reduced the
populations of many large carnivores such as wolves in
North America and Europe and tigers in Asia, and now
smaller carnivores such as coyote (Canis latrans) and lynx
(Lynx spp.) fulfi ll part of this niche.[62,63]
Some animals have adapted unique behaviors to better
suit them for the varied seasons. For example, some
Oak–Hickory and Beech–Maple.[42] The temperate decidu-
ous broadleaved forest that extends across East Asia (from
30° to 60°N) displays the greatest diversity of vegetation.
The greater diversity in Asian temperate deciduous forests is
likely the result of being less severely glaciated during the last
ice age. This was largely a result of the more connected land-
mass, which allowed temperate forests to retreat from the
advancing glaciers. For instance, in North America, the Flor-
ida peninsula and Gulf of Mexico coastal region was the
southern limit for temperate forest refuge, while in Europe
the east to west mountain range of the Alps limited southern
migration. Other locations of deciduous broadleaved forests
include areas around the eastern Black Sea, mountainous
regions in Iran and Caspian Sea, and a narrow strip of South
America including southern Chile and Argentina.[34,43–45]
The composition of deciduous temperate forest depends
on latitude. In North America, birches (Betula spp.), aspen
(Populus spp.), and maples (Acer spp.) are common, along
with needle-leaved conifers such as pines (Pinus spp.).
Coniferous species tend to decline in abundance further
south, but they remain in localized areas that exhibit drier,
more nutrient-poor conditions and higher altitudes.
In Europe, forests at higher elevations are composed
mainly of conifers (Picea spp. and Abies spp.) and Fagus
spp., as well as larch (Larix decidua).
In central temperate deciduous forests, broadleaved
deciduous species in Europe include oaks (Quercus spp.),
European beech (Fagus sylvatica), and hornbeam (Carpinus
betulus) [with Fraxinus spp. and Castanea spp. often just as
common as Carpinus]. Common trees in North America
include sugar maple (Acer saccharum), American beech
(Fagus grandifolia), basswoods (Tilia spp.), oaks, and hick-
ories (Carya spp.). In Asia, Japanese beech (Fagus crenata),
oaks, maples, ashes (Fraxinus spp.) and basswoods are most
common. At lower latitudes, broadleaved deciduous species
are joined by broadleaved evergreen angiosperm trees, such
as oaks, beeches (Northofagus spp.), eucalyptus ( Eucalyptus
spp.), and pines (Podocarpus spp.).
Plant life
Deciduous leaves are the most distinctive feature of tem-
perate deciduous broadleaved forests. Autumn changes in
leaf color and senescence have become very important to
these regions both economically and ecologically. “Leaf-
peeping” tourists travel to deciduous forest destinations to
view displays of reds, oranges, and yellows and invest in
local markets.[46,47] The dropping of leaves in the fall is also
a phenological event or recurring biological cycle that
changes the seasonal microhabitat, creating unique under-
story conditions for other plants and animals.
The onset of plant phenophases such as spring leaf fl ush
and autumn leaf senescence are controlled by two major
environmental signals: photoperiod (relative length of
days/nights) and total degree days (relative measure of

218 Forests: Temperate Evergreen and Deciduous
Ecotone—
Fragmentation
Plant life
While broadleaved trees in the Northern Hemisphere are
exposed to variable environmental stress, broadleaved trees
in the Southern Hemisphere are under weaker seasonal
forces. Because of milder climates and less extreme vari-
ability in the Southern Hemisphere, it is more advanta-
geous for trees to expose leaves all year round in order to
maximize photosynthesis and carbon gain.[66,67] Temperate
broadleaved evergreen tree leaves are generally thick with
smooth margins as opposed to deciduous leaves, which by
comparison are generally thin and lobed. These differences
are in part due to higher levels of rainfall and humidity.
Southern Hemisphere species maintain a waxy outer layer
on leaves to repel water. In these mild temperate evergreen
forests, species compositions are mainly from the fami-
lies of Lauraceae, Fagaceae, Theaceae, Magnoliaceae,
and Hamamelidaceae, all of which have a relatively low
resistance to cold and freezing temperatures.[68,69] In the
Southern Hemisphere, the very dry-adapted forests of
Australia are also found. Fire disturbance in this dry region
has allowed for the domination of Eucalyptus spp. which
have thick bark for fi re-resistance.[70,71]
Also relatively common in many temperate evergreen
forests are narrow-leaved evergreen trees. These species
produce their seeds in compact structures called cones,
which allow for greater protection of reproductive material
from predation as well as some unique adaptations to envi-
ronmental conditions.[72] Some evergreen conifers have
fl at, triangular scale-like leaves, such as found in the
Cupressaceae and some Podocarpaceae, while many ever-
green trees also produce needle-like leaves, which increase
the leaf area for photosynthesis and are thicker and smaller
to avoid desiccation.[73,74] Rates of area- or massed-based
leaf photosynthesis in temperate needle-liked evergreen
trees are typically much lower than for leaves of temperate
deciduous tree species, except when amortized over the life
of the leaf.[75] Temperate deciduous trees only keep their
leaves for one growing season, while temperate evergreen
forests keep their leaves for much longer.
The moist temperate coniferous forests dominated by
tree species such as giant sequoias (Sequoiadendron gigan-
teum), coastal redwoods (Sequoia sempervirens), Douglas
fi rs (Pseudotsuga menziesii), and kauris (Agathis australis)
sustain the highest levels of biomass in any terrestrial eco-
system.[76] These forests are rare and known for a rich vari-
ety of plant and animal species. Typically, two or more tree
layers of large broadleaved, dome-shaped evergreen angio-
sperms are present. These forests, which grow in moist
regions, are also known for higher abundance and diversity
of epiphytes and vines.
In maritime climates, fog can provide a signifi cant
amount of the annual water available to vegetation.[77]
Forming over adjacent oceans and water bodies, the moist
air travels with both humidity and nutrients that can be
mammals store fat and hibernate or go into torpor during
the cold months. Some birds take shelter in the trees by
becoming cavity-nesters, while others are migratory and
avoid the cold winters by fl ying to more southerly climes.
Soils and nutrients
Temperate deciduous forest soils show variation with lati-
tude and altitude, as well as with moisture. At the northern
or higher extent of the range, where there is a greater pres-
ence of needle-leaf evergreen tree species, soils tend to be
relatively acidic and nutrient poor. By contrast, soils are
strongly leached and less productive in the southern or
lower ranges. Temperate deciduous forest soils, however,
tend to be deep and fertile and do not freeze year-round.
Brown forest soils (alfi sols, in the American soil taxon-
omy) develop under the nutrient-demanding broadleaf
trees whose leaves bind the major nutrient bases.[64,65] This
causes leaf litter to be less acidic and provide a rich humus
layer. Along with the favorable summer growing season,
this is one of the reasons why this biome has historically
been popular for agricultural use.
Temperate Evergreen Forest
Temperate broadleaved evergreen forests occur at the
warm, moist end of the latitudinal gradient of temperate
forests and are characterized by long, hot, humid growing
seasons, and relatively milder winters. Precipitation is high
(1000–1750 mm) and the forest is dominated by broad-
leaved evergreen angiosperms. These forests occur in east-
ern Asia, coastal regions of New Zealand and Australia,
western coastal regions of North America and parts of Chile,
South Africa, and warmer European regions.[34, 43–45] There
are several forest variants that occur in regions of maritime
and dry-season climates. Maritime broadleaved forests are
dominated by tall, broadleaved evergreen angiosperms and
narrow- to broadleaved conifers and exhibit very high pre-
cipitation (2000–3000 mm). Maritime needle-leaved forests
occur only in a narrow band along the coastline of North
America from California to Alaska and throughout coastal
southern Europe and are known for needle- and scale-leaved
evergreen conifers such as giant sequoias in North America
or angiosperms such as evergreen oaks in Europe. Dry-
season broad-sclerophyll forests occur in areas with periodic
drought, exhibiting more of a Mediterranean climate, with
hot dry summer and cool wet winters.
Needle-leaved conifers can be interspersed within
deciduous canopies or occur in patches. They tend to grow
at higher altitudes near treeline or in drier regions. Condi-
tions can be highly seasonal, with severe winters and warm,
humid summers. Conifer tree species include fi r (Abies
spp.), spruce (Picea spp.), pine (Pinus spp.), cedar (Thuja
spp.), hemlock (Tsuga spp.), and larch (Larix decidua)
among others.

Forests: Temperate Evergreen and Deciduous 219
Ecotone—
Fragmentation
temperature and metabolism without eating, drinking, def-
ecating, or urinating.[92] In austral regions, temperate ever-
green forests are also home to many endemic species such
as Leadbeater’s possum (Gymnobelideus leadbeaten),
Parma wallaby (Marcopus parma) and Albert’s lyrebird
(Menura alberti).
Soil and nutrients
Soils of temperate evergreen forests are well leached and
low in productivity. Ultisols represent older, unglaciated
soils that have been weathered to a much greater degree
than those of temperate deciduous forests. Ultisols are gen-
erally less fertile than most temperate deciduous forest
soils and were further degraded under plantation and sub-
sistence agriculture in both the colonial and postcolonial
periods.[65,93]
THREATS
Threats to the temperate forest biome stem from direct
or indirect human activity. Historically, farming has been
one of the main reasons for deforestation in temperate
biomes. Today, an increasing number of roads, residential
and other developments have left forests highly frag-
mented, raising concerns over the healthy functioning of
these ecosystems.[94] Many of the plant species living in
temperate forests are well-adapted and function optimally
in soils with fairly low nutrient availability. However, more
impervious surfaces as well as modern agricultural and
industrial practices can alter nutrient availability and con-
tribute to a decline in forest health. Atmospheric deposition
of nitrogen, acid deposition and fertilizer applications are
changing the rate of input of nutrients along with the growth
and dynamics of temperate forest communities.[95–97] This
also leads to alterations in soil properties, nutrient cycling
and microbial community composition as well as nutrient
imbalances in plants and increases in forest mortality.
These nutrient imbalances in plants make them more vul-
nerable to other outside stressors such as diseases and
introduced invasive species.[98–100]
Human activities are also altering global climate, a
foundation for the characterization and distribution of tem-
perate forests. For deciduous canopies, observational evi-
dence has linked global climate change to differences in the
timing of spring leaf growth and autumn leaf drop. Warm
springs cause leaves to grow earlier, sometimes by up to
one month (http://www.usanpn.org), lengthening the dura-
tion of the growing season in some temperate forests.[101–103]
Climate variation may also result in a shift in fl owering
periods, potentially causing disruption in timing with pol-
linators/dispersers. In Japan, cherry blossom (Prunus
serrulata) trees are blooming earlier than they did histori-
cally, causing concerns regarding the traditional festivals
and tourism that surround the annual event.[104] Evidence of
intercepted by plants, reducing water stress caused by
transpiration. Some studies suggest that some leaf shapes
have evolved for more effi cient collection of fog drip.
It has also been hypothesized that fog in dry summer
months is essential in supporting the sustained growth of
large trees such as the redwoods of California.[78,79]
Temperate coniferous forests at higher altitudes often
have narrow tree morphologies which allow heavy snow to
slough off branches before breaking them.[72,73] These for-
ests are dense, with trees growing close together to mini-
mize wind damage; wind minimization is important
because conifers often have very shallow root systems due
to higher allocation of nutrients in the top layers of soil.[80,81]
Evergreen trees in higher, drier areas allocate carbon to
thick bark for greater protection from low-heat summer
fi res. Some conifers have adapted to take advantage of fi re
events, producing serotinous cones in which seeds are
maintained in a dormant state until a fi re occurs, opening
the cones and releasing the seeds for germination.[82,83]
Because the canopy of these temperate forests is ever-
green, the vegetation in the understory experiences low light
availability. Understory plants have adapted to maximize
their light/energy capture as well as their photosynthetic
capacity under such conditions. Shaded plants tend to allocate
more growth to leaves, displaying larger, thinner leaves
which take up more horizontal area for light capture.[75,84]
Physiologically, shade leaves saturate at lower light levels
and have lower light compensation points, which allows
them to maintain positive rates of net photosynthesis at low
light levels. Understory plants also adapt to respond quickly
to environmental changes in order to make use of sunfl ecks
or brief, unpredictable periods of high sunlight usually lasting
only seconds.[85–87] Instantaneous adjustments in photosyn-
thetic rates allow plants to be effi cient with the limited light
they receive to put toward carbon gain.
Animal life
The fauna of the temperate evergreen forest is similar to
that of the deciduous forest ecosystems, with many smaller
mammals, large ungulates and a few larger omnivorous
mammals contributing to biological interactions. Many of
these species, mostly birds and small rodents, have adapted
skills to compensate for the scarcity of food in these nutri-
ent poor systems. The red squirrel (Sciurus vulgaris)
removes cone scales to reach the nutrient rich seeds of the
coniferous trees.[88,89] They often hoard these seeds, many
of which they do not recover, resulting in seed dispersal.
In dry forests at higher latitudes, large ungulates such as
moose (Alces alces) browse nutritious spring vegetation
and wetland foliage in warmer seasons and on young
tree shoots when the ground is frozen and covered with
snow.[90,91] Some fauna in the coniferous evergreen forests
located at higher latitudes have also adapted to the cold,
snowy winters. For example, bears (Ursus spp.) are able to
go into long periods of dormancy, maintaining both body

220 Forests: Temperate Evergreen and Deciduous
Ecotone—
Fragmentation
sustainable use of these resources and proper protection
and management of temperate forests will be important in
ensuring the health and productivity of the biome.
REFERENCES
1. Thomas, S.C.; MacLellan, J. Boreal and Temperate For-
ests, in Forests and Forest Plants. In Encyclopedia of Life
Support Systems (EOLSS); UNESCO, Eolss Publishers:
Oxford, UK, 2004.
2. Chabot, B.F.; Mooney, H.A., Eds.; Physiological Ecology
of North American Plant Communities; Chapman and
Hall: New York, NY, 1985.
3. Small, C.J.; McCarthy, B.C. Spatial and temporal varia-
tion in the response of understory vegetation to distur-
bance in a central Appalachian oak forest. J. Torrey Bot.
Soc. 2002, 129 (2), 136–153.
4. Gendron, F.C.; Comeau, P.G. Temporal variation in the
understory photosynthetic photon fl ux density of a decidu-
ous stand: the effects of canopy development, solar eleva-
tion and sky conditions. Agric. Forest Meteorol. 2001,
106, 23–40.
5. Woodward, F.I. Climate and Plant Distribution; Cam-
bridge University Press: New York, NY, 1987.
6. Axelrod, D.I. Origin of deciduous and evergreen habits in
temperate forests. Evolution 1966, 20, 1–15.
7. Food and Agriculture Organization of the United Nations.
State of the World’s Forests, 2009.
8. Vankat, J.L. Boreal and Temperate Forests. In eLS; John
Wiley & Sons Ltd: Chichester, England, 2002.
9. Reich, P.B.; Bolstad, P. Temperate Forests: Evergreen and
Deciduous. In Terrestrial Global Productivity; Mooney,
H., Roy, J., Saugier, B., Eds.; Academic Press: San Diego,
CA, 2001.
10. Graumlich, L.J. A 1000-year record of temperature and pre-
cipitation in the Sierra Nevada. Quarter. Res. 1993, 39, 249.
11. Elsevier. Coniferous Forests. In Ecosystems of the World,
Vol. 6.; Elsevier: Amsterdam, the Netherlands, 2005.
12. Ovington, J.D. Temperate Broad-Leaved Evergreen For-
ests; Elsevier: Amsterdam, the Netherlands, 1983.
13. Ohte, N; Tokuchi, N. Hydrology and biogeochemistry of
temperate forests. In Forest Hydrology and Biogeochemistry:
Synthesis of Past Research and Future Directions; Levia,
D.F; Carlyle, D; Tanaka, T., Eds.; Springer Science+Business
Media: New York, NY, 2011.
14. Beckage, B.; Osborne, B.; Gavin, D.G.; Pucko, C.; Sic-
cama, T.; Perkns, T. A rapid upward shift of a forest eco-
tone during 40 years of warming in the Green Mountains of
Vermont. Proc. Nat. Acad. Sci. 2008, 105 (11), 4197–4202.
15. Loarie, S.R.; Duffy, P.B.; Hamilton, H.; Asner, G.P.; Field,
C.B.; Ackerly, D.D. The velocity of climate change.
Nature 2009, 462 (24), 1052–1055.
16. Holtmeier, F.K.; Broll, G. Sensitivity and response of
northern hemisphere altitudinal and polar treelines to
environmental change at landscape and local scales.
Global Ecol. Biogeogr. 2005, 14, 395–410.
17. Harsch, M.A.; Hulme, P.E.; McGlone, M.S.; Duncan,
R.P. Are treelines advancing? A global meta-analysis of
treeline response to climate warming. Ecol. Lett. 2009,
12, 1040–1049.
warming temperatures also suggests a shift in species dis-
tributions toward the poles, which could result in the
extinction of some plant and animal species.[105,106]
Rapid global climate change creates conditions less suit-
able for many species adapted to their particular environ-
ment, increasing the amount of stress put on individuals as
well as their vulnerability to certain pathogens or pests.[107]
Climate also has a strong infl uence on the success of cer-
tain fungi, bacteria, and viruses, affecting infection sever-
ity and spread. For example, in the northwestern United
States, heavy rains and wet weather during warm periods
are optimal conditions for sudden oak death infection.[108]
In many cases, humans accidentally introduce these dis-
eases. Chestnut blight (Cryphonectria parasitica), a fun-
gus introduced to America from Europe in 1904, is one of
the best-known examples. In four decades, the chestnut
blight largely eliminated American chestnut (Castanea
dentata) from the continent, one of the most important and
widespread tree species across the eastern temperate forest
of North America.[109,110]
Other introductions to temperate forests include nonna-
tive plant invaders that have, in some cases, taken over large
portions of forested land, changing species, and nutrient
compositions as well as the identity of the landscape.[111–113]
In some cases, invasive species have the potential to reduce
biodiversity, extirpate native species and negatively impact
ecosystem structure and services, thereby negatively affect-
ing the economy and human health and well-being. Species
such as garlic mustard (Alliaria petiolata), native to Europe
and introduced to forests of North America, exude chemi-
cals that may disturb natural plant–plant and plant–soil
processes and interactions.[114] Many invasives may be
unpalatable to native fauna such as Japanese knotweed
(Fallopia japonica)[115] which is nonnative to forests of
Europe and North America or even poisonous like
St. John’s Wort (Hypericum perforatum) in Australia.[116]
Negative effects of invasive plants on human populations
include economic losses[117] and even disease as in the case
of Japanese barberry (Berberis thunbergii), a shrub intro-
duced to North America that creates favorable habitat for
vectors of Lyme disease (Borrelia burgdorferi).[118]
CONCLUSION
Temperate forests are ecologically distinct biomes and pro-
vide important regional and global ecosystem services. The
species in these ecosystems have evolved unique physio-
logical traits to function under a variety of seasonal stresses.
Being closely related to variation in environmental condi-
tions, temperate forests may act as an ecological indicator
of global climate change, changes in nutrient cycling and
other environmental alterations brought on by human
activities. Temperate forests have been and are of impor-
tant use to industry and tourism. As they continue to play
an important role in regional cultures and economies,

Forests: Temperate Evergreen and Deciduous 221
Ecotone—
Fragmentation
37. Allaby, M. Temperate Forests. Biomes of the Earth Series;
Chelsea House: New York, NY, 2006.
38. Oliver C.D.; Larson, B.C. Forest Stand Dynamics. In
Biological Resource Management Series; McGraw Hill:
New York, NY, 1990.
39. Schelhaas, M.; Nabuurs, G.; Schuck, A. Natural distur-
bances in the European forests in the 19th and 20th centu-
ries. Global Change Biol. 2003, 9 (11), 1620–1633.
40. Lorimer, C.G.; White, A.S. Scale and frequency of natural
disturbances in the northeastern U.S. implications for
early successional forest habitats and regional age distri-
butions. Forest Ecol. Manag. 2003, 185, 41–64.
41. Aber, J.D.; Nadelhoffer, K.J.; Steudler, P.; Melillo, J.M.
Nitrogen Saturation in Northern Forest Ecosystems. Bio-
Science 1989, 39, 378–386.
42. Braun, E.L. Deciduous Forests of Eastern North America;
Hafner Publishing Co.: New York, 1972.
43. Röhig, E.; Ulrich, B. Temperate Deciduous Forests. In
Ecosystems of the World; Röhig, E.; Ulrich, B. Eds.; Else-
vier: Amsterdam, the Netherlands, 1991.
44. Archibold, O.W. Ecology of World Vegetation; Chapman
and Hall: London, UK, 1995.
45. Sayre, A.P. Temperate Deciduous Forest. Twenty-First
Century Books: New York, NY, 1994.
46. Beckley, T.M. The nestedness of forest dependence: a con-
ceptual framework and empirical exploration. Soc. Nat.
Resour.: An Int. J. 1998, 11 (2), 101–120.
47. Norris, G.A. The Economical Impact of Climate Change
on the New England Region. New England Regional
Assessment 2008.
48. Mahall, B.E.; Bormann, F.H. A quanititative description
of the vegetative phenology of herbs in a northern hard-
wood forest. Bot. Gazette 1978, 139, 467–481.
49. Korner, C.; Basler, D. Phenology under global warming.
Science 2010, 327 (5972), 1461–1462.
50. Havrenek, W.M.; Tranquillini, W. Physiological processes
during winter dormancy and their ecological signifi cant.
In Ecophysiology of Coniferous Forests; Smight, W.K.,
Hinckley T.M. Eds.; Springer-Verlag: Berlin, 1995.
51. Hutchison, B.A.; Matt, D.R. The distribution of solar radi-
ation within a deciduous forest. Ecol. Monogr. 1977, 47,
185–207.
52. Reifsnyder, W.E.; Furnival, C.M.; Horowitz, J.L. Spatial
and temporal distribution of solar radiation beneath forest
canopies. Agric. Meteorol. 1971, 9, 21–38.
53. Hoch, W.A.; Singsaas, E.L.; McCown, B.H. Resorption
protections. Anthocyanins facilitate nutrient recovery in
autumn by shielding leaves from potentially damaging
light levels. Plant Physiol. 2003, 113 (3), 1296–1305.
54. Dreiss, L.M.; Volin, J.C. Infl uence of leaf phenology and
site nitrogen on invasive species establishment in temper-
ate deciduous forest understories. Forest Ecol. Manag.
2013, 296, 1–8.
55. Gill, D.S.; Amthor, J.S.; Bormann, F.H. Leaf phenology,
photosynthesis and the persistence of saplings and shrubs
in a mature northern hardwood forest. Tree Physiol. 1998,
18, 281–289.
56. Sefcik, L.T.; Zak, D.R.; Ellsworth, D.S. Photosynthetic
responses to understory shade and elevated carbon dioxide
concentration in four northern hardwood tree species. Tree
Physiol. 2006, 26, 1589–1599.
18. Breshears, D.D.; Cobb, N.S.; Rich, P.M.; Price, K.P.;
Allen, C.D.; Balice, R.G.; Romme, W.H.; Kastens, J.H.;
Floyd, M.L.; Belnap, J.; Anderson, J.J.; Myers, O.B.;
Meyer, C.W. Regional vegetation die-off in response to
global-change-type drought. Proc. Nat. Acad. Sci. 2005,
102 (42), 15144–15148.
19. Adams, H.D.; Guardiola-Claramonte, M.; Barron-Gaf-
ford, G.A.; Villegas, J.C.; Breshears, D.D.; Zou, C.B.;
Troch, P.A.; Huxman, T.E. Temperature sensitivity of
drought-induced tree mortality portends increased regional
die-off under global-change-type drought. Proc. Nat.
Acad. Sci. 2009, 106 (17), 7063–7066.
20. Denton, G.H.; Hughes, T.J. The Arctic ice sheet: an outra-
geous hypothesis. In The Last Great Ice Sheets; Denton,
G.H., Hughes, T.J., Eds.; John Wiley: New York, NY,
1981.
21. Cox, C.B.; Moore, P.D. Biogeography: An Ecological and
Evolutionary Approach; John Wiley: New York, NY, 2010.
22. Foster, D.R. Land Use History (1730–1990) and Vegeta-
tion Dynamics in Central New England. J. Ecol. 1995,
76, 135–151.
23. Foster, D.R.; Motzkin, G.; Slate, B. Land-use history as
long-term broad-scale disturbance: regional forest dynam-
ics in central New England. Ecosystems 1998, 1L, 96–119.
24. Ferrow, B.E. A Brief History of Forestry in Europe, the
United States and Other Countries; University Press:
Cambridge, MA, 1991.
25. Volin, V.C.; Buongiorno, J. Effects of alternative manage-
ment regimes on forest stand structure, species composi-
tion, and income: a model for the Italian Dolomites. Forest
Ecol. Manag. 1996, 87 (1–3), 107–125.
26. Perlin, J.A. Forest Journey: The Role of Wood in the
Development of Civilization; Harvard University Press:
London, UK, 1991.
27. Lehtinen, A.A.; Donnor-Amnell, J.; Saether, B., Eds. Poli-
tics of Forests: Northern Forest-Industrial Regimes in the
Age of Globalization; Ashgate Publishing: Great Britain,
2004.
28. USDA Forest Service. U.S. Forest Facts and Historical
Trends; FS-696. USDA Forest Service: Washington, DC,
2001.
29. Zerbe, S. Restoration of natural broad-leaved woodland in
Central Europe on sites with coniferous forest plantations.
Forest Ecol. Manag. 2001, 167 (1–3), 27–42.
30. Grichuk, V.P. Late Pleistocene vegetation history. In Late
Quaternary Environments of Soviet Union; Vwlichko,
A.A., Eds.; University of Minnesota Press: Minneapolis,
MN, 1984.
31. Ehlers, J; Gibbard, P.L. The extent and chronology of
Cenozoic global glaciations. Quarter. Int. 2007, 164–165,
6–20.
32. http://www.iucnredlist.org (accessed March 2012).
33. Williams, M. Americans and Their Forests; Cambridge
University Press: Cambridge, MA, 1989.
34. White, M.A. Deciduous Forests. Encyclopedia.com 2001
(accessed January 2012).
35. Thompson, J.N. Treefalls and colonization patterns of tem-
perate forest herbs. Am. Midland Nat. 1980, 104, 176–184.
36. Rees, M.; Condit, R.; Crawley, M.; Pacala, S.; Tilman, D.
Long-term studies of vegetation dynamics. Science 2001,
293 (5530), 650–655.

222 Forests: Temperate Evergreen and Deciduous
Ecotone—
Fragmentation
75. Busing, R.T.; Fujimori, T. Biomass, productions and
woody detritus in an old coast redwood (Sequoia semper-
virens) forest. Plant Ecol. 2005, 177, 177–188.
76. Ewing, H.A.; Weathers, K.C.; Templer, P.H.; Dawson,
T.E.; Firestone, M.K.; Elliott, A.M.; Boukili, V.K. Fog
water and ecosystem function: heterogeneity in a Califor-
nia redwood forest. Ecosystems 2009, 12, 417–433.
77. Canadell, J.; Jackson, R.B.; Ehleringer, J.R.; Mooney,
H.A.; Sala, O.E.; Schulze, E.D. Maximum rooting depth
of vegetation types at the global scale. Oecologia 1996,
108 (4), 583–595.
78. Burgess, S.S.O.; Dawson, T.E. The contribution of fog to
the water relations of Sequoia sempervirens (D. Don):
foliar uptake and prevention of dehydration. Plant, Cell
Environ. 2004, 27 (8), 1023–1034.
79. Simonin, K.A.; Santiago, L.S.; Dawson, T.E. Fog inter-
ception by Sequoia sempervirens (D. Don) crowns decou-
ples physiology from soil water defi cit. Plant, Cell
Environ. 2009, 32, 882–892.
80. Enright N.J.; Marsula R.; Lamont B.B.; Wissel C. The
ecological signifi cance of canopy seed storage in fi re-
prone environments: a model for non-sprouting shrubs. J.
Ecol. 1998, 86, 946–959.
81. Echenwalder, J.E. Conifers of the World; Timber Press,
Inc.: Portland, OR, 2009.
82. Givnish, T.J. Adaptation to sun and shade: a whole plant
perspective. Aust. J. Plant Physiol. 1988, 15, 63–92.
83. Midgley, J.; Bond, W. Pushing back in time: the role of fi re
in plant evolution. New Phytol. 2011, 191 (1), 5–7.
84. Boardman, N.K. Comparative photosynthesis of sun and
shade plants. Annu. Rev. Plant Physiol. 1977, 28, 355–377.
85. Chazdon, R.L.; Pearcy, R.W. The importance of sunfl ecks
for understory plants. BioScience 1991, 41 (11), 760–766.
86. Brantley, S.T.; Young, D.R. Contribution of sunfl ecks is
minimal in expanding shrub thickets compared to temper-
ate forest. Ecology 2009, 90, 1021–1029.
87. Lertzman, K.P.; Sutherland, G.D.; Inselberg, A.; Saunders,
S.C. Canopy gaps and the landscape mosaic in a coastal
temperate rain forest. Ecology 1996, 77, 1254–1270.
88. Miyaki, M. Seed dispersal of the Korean pine, Pinus
koraiensis, by the red squirrel, Sciurus rulgaris. Ecol. Res.
1987, 2 (2), 147–157.
89. Wauters, L.A.; Gurnell, J.; Martinoli, A.; Tosi, G. Does
interspecifi c competition with introduced grey squirrels
affect foraging and food choice of Eurasian red squirrels?
Animal Behav. 2001, 61 (6), 1079–1091.
90. Hjeljord, O.; Sundstol, F.; Haagenrud, H. The nutritional
value of browse to moose. J. Wildlife Manag. 1982, 46 (2),
333–343.
91. Wam, H.K.; Hjeljord, O. Moose summer and winter diets
along a large scale gradient of forage availability in south-
ern Norway. Eur. J. Wildlife Res. 2010, 56, 745–755.
92. Hellgren, E.C. Physiology of hibernation in bears. Int.
Assoc. Bear Res. Manag. 1995, 10, 467–477.
93. Mbagwu, J.S.C.; Lal, R.; Scott, T.W. Effects of desurfac-
ing alfi sols and ultisols in southern Nigeria: I. Crop Per-
formance. Soil Sci. Soc. Am. J. 1984, 48, 828–833.
94. Foley, J.A.; DeFries, R.; Asner, G.P.; Barford, C.; Bonan,
G.; Carpenter, S.R.; Chapin, F.S.; Coe, M.T.; Daily, G.C.;
Gibbs, H.K.; Hellowski, J.H.; Holloway, T.; Howard,
E.A.; Kucharik, C.A.; Monfreda,C.; Patz, J.A.; Prentice,
57. Hattenschwiler, S.; Korner, C. Tree seedling responses to
in situ CO2-enrichment differ among species and depend
on understorey light availability. Global Change Biol.
2000, 6, 213–226.
58. Richardson, A.D.; O’Keefe, J. Phenological differences
between understory and overstory: a case study using the
long-term Harvard Forest Records Book. In Phenology of
Ecosystem Processes: Applications in Global Change
Research; Noormets, A., Eds.; Springer: New York, NY, 2009.
59. Augspurger, C.K. Early spring leaf out enhances growth
and survival of saplings in a temperate deciduous forest.
Oecologia 2008, 156, 281–286.
60. Kudo, G.; Ida, T.Y.; Tani, T. Linkages between phenology,
pollination, photosynthesis and reproduction in deciduous
forest understory plants. Ecology 2008, 89, 321–331.
61. Li, H.; Zhang, Z. Effects of mast seeding and rodent abun-
dance on seed predation and dispersal by rodents in
Prunus armeniaca (Rosaceae). Forest Ecol. Manag. 2008,
242, 511–517.
62. Nowak, R.M. Evolution and taxonomy of coyotes and
related Canis. In Coyotes: Biology, Behavior, and
Management; Bekoff, M., Ed.; Academic Press: New York,
1978.
63. Thurber, J.M.; Peterson, R.O. Changes in body size asso-
ciated with range expansion in the coyote (Canis latrans).
J. Mammal. 1991, 72 (4), 750–755.
64. Ulrichm B. Stability, elasticity and resilience of terrestrial
ecosystems with respect to matter balance. In Potentials
and Limitations of Ecosystem A nalysis; Schulze, E.D.,
Zwölfer, H. Eds.; Springer Verlag: Berlin, 1987.
65. Hendrick, R.L. Forest Types and Classifi cation. In The
Forests Handbook, Volume 1: An Overview of Forest Sci-
ence; Evans, J., Ed.; Blackwell Science Ltd: Oxford, UK,
2008.
66. Chabot, B.F.; Hicks, D.J. The ecology of leaf life spans.
Annu. Rev. Ecol. Syst. 1982, 13, 229–259.
67. Hallik, L.; Niinements, U.; Wright, I.J. Are species shade
and drought tolerance refl ected in leaf-level structural and
functional differentiation in Northern Hemisphere temper-
ate woody fl ora? New Phytol. 2009, 184, 257–274.
68. Lieth, H.; Box, E.O.; Peet, R.K.; Masuzawa, T.; Yamada,
I.; Fujiwara, K.; Maycock, P.F. Handbook of Vegetation
Science; Kluwer Academic Publishers: the Netherlands,
1995.
69. Bannister, P. Godley review: a touch of frost? Cold hardi-
ness of plants in the southern hemisphere. New Zealand J.
Bot. 2007, 45 (1), 1–33.
70. Beard, J.S. Temperate forests of the Southern Hemisphere.
Vegio 1990, 89 (1), 7–10.
71. Fire in ecosystems of south-west Western Australia:
impacts and management. Symposium proceedings (Vol.
1): 225–268, Perth, Australia, 2002.
72. Leslie, A.B. Predation and protection in the macroevolu-
tionary history of conifer cones. Proc. R. Soc. B 2011,
278, 3003–3008.
73. Sprugel, D.G. The relationship of evergreenness, crown
architecture, and leaf size. Am. Nat. 1989, 133 (4),
465–479.
74. Brodribb, T.J.; Field, T.S.; Sack, L. Viewing leaf structure
and evolution from a hydraulic perspective. Func. Plant
Biol. 2010, 37, 499–498.

Forests: Temperate Evergreen and Deciduous 223
Ecotone—
Fragmentation
108. Kelly, M.; Meentemeyer, R.K. Landscape dynamics of the
spread of Sudden Oak Death. Photogrammatic Eng.
Remote Sens. 2002, 68 (10), 1001–1009.
109. Anagnostakis, S.L. Chestnut blight: the classical problem
of an introduced pathogen. Mycologia 1987, 79 (1),
23–37.
110. Paillet, F.L. Chestnut: history and ecology of a trans-
formed species. J. Biogeogr. 2002, 29, 1517–1530.
111. Walker L.R.; Smith S.D. Impacts of invasive plants on
community and ecosystem properties. In Assessment and
Management of Plant Invasions; Luken, J.O., Thieret,
J.W., Eds.; Springer: New York, NY, 1997.
112. Kohli, R.K.; Jose, S.; Singh, H.P.; Batish, D.R. Invasive
Plants and Forest Ecosystems.; CRC Press: Boca Raton,
FL USA, 2009.
113. Daehler, C.C. Performance comparisons of co-occuring
native and alien invasive plants: implications for conserva-
tion and restorations. Annu. Rev. Ecol. Evol. Syst. 2003,
34, 183–211.
114. Stinson, K.A.; Campbell, S.A.; Powell, J.R.; Wolfe, B.E.,
Callaway, R.M.; Thelen, G.C.; Hallet, S.G.; Prati, D.;
Klironomos, J.N. Invasive plant suppresses the growth of
native tree seedlings by disrupting belowground mutual-
isms. PLoS Biol. 2006, 4 (5), e140.
115. Krebs, C.; Gerber, E.; Mathies, D.; Schaffner, U. Herbi-
vore resistance of invasive species and their hybrids.
Oecologia 2011, 167 (4), 1041–1052.
116. Southwell, I.A.; Bourke, C.A. Seasonal variation in hyper-
icin content of Hypericum perforatum (St. John’s Wort).
Phytochemistry 2001, 56 (5), 437–441.
117. Pimentel, D.; Lach, L.; Zuniga, R.; Morrison, D. Environ-
mental and economic costs associated with non-indige-
nous species in the United States. BioScience 2000, 50
(1), 53–65.
118. Williams, S.C.; Ward, J.S.; Worthley, T.E.; Stafford, K.C.
Managing Japanese barberry (Ranunculales: Berberidacea)
infestations reduces blacklegged tick (Acari: Ixodidae)
abundance and infection prevalence with Borrelia
burgdorferi (Spirochaetales: Spirochaetaceae). Environ.
Entomol. 2009, 38 (4), 977–984.
I.C.; Ramankutty, N.; Snyder, P.K. Global consequences
of land use. Science 2005, 309 (5734), 570–574.
95. Vitousek, P.M.; Aber, J.D.; Howarth, R.W.; Likens,
G.E.; Matson, P.A.; Schindler, D.W.; Schlesinger,
W.H.; Tilman, D.G Human alteration of the global
nitrogen cycle: sources and consequences. Ecol. Appl.
1997, 7 (3), 737–750.
96. Gundersen, P.; Schmidt, I.K.; Raulund-Rasmussen, K.
Leaching of nitrate from temperate forests: effects of air
pollution and forest management. Environ. Rev. 2006,
14 (1), 1–57.
97. Aber, J.D.; Goodale, C.L.; Ollinger, S.V.; Smith, M.-L.;
Magill, A.H.; Martin, M.E.; Hallett, R.A.; Stoddard, J.L.
Is nitrogen deposition altering the nitrogen status of
Northeastern forests? BioScience 2003, 53 (4), 375–389.
98. Waring, R.H. Characteristics of trees predisposed to die.
BioScience 1987, 37 (8), 569–574.
99. Bytnerowicz, A.; Omasa, K.; Paoletti, E. Integrated effects
of air pollution and climate change on forests: a northern
hemisphere perspective. Environ. Poll. 2007, 14, 438–445.
100. Turner, J.; Lambert, M. Soil and nutrient processes related
to eucalypt forest dieback. Aust. Forest. 2005, 68 (4),
251–256.
101. Menzel, A.; Fabian, P. Growing season extended in
Europe. Nature 1999, 397, 659.
102. Cleland, E.E.; Chuine, I.; Menzel, A.; Mooney, H.A.;
Schwartz, M.D. Shifting plant phenology in response to
global change. Trends Ecol. Evol. 2007, 22 (7), 357–365.
103. Badeck, F-W.; Bondeau, A.; Bottcher, K.; Doktor, D.;
Lucht, W.; Schaber, J.; Sitch, S. Responses of spring phe-
nology to climate change. New Phytol. 2004, 162 (2),
295–309.
104. Aono, Y.; Kazui, K. Phenological data series of cherry tree
fl owering in Kyoto, Japan, and its application to recon-
structions of springtime temperatures since the 9th century.
Int. J. Climatol. 2008, 28 (7), 905–914.
105. Malcolm, J.R.; Markham, A. Global Warming and Terres-
trial Biodiversity Decline. World Wildlife Fund Report,
WWF U.S.: Washington, DC, 2000.
106. Parmesan, C. Ecological and evolutionary responses to
recent climate change. Annu. Rev. Ecol. Evol. Syst. 2006,
37, 637–669.
107. Ayres, M.P.; Lombardero, M.J. Assessing the conse-
quences of global change for forest disturbance from her-
bivores and pathogens. Sci. Total Environ. 2000, 262,
263–286.