3794 Ecological Processes | Wind Effects
Further Reading
Braun CE (ed.) (2005) Techniques for Wildlife Investigations and
Management, 6th edn. Bethesda, MD: The Wildlife Society.
Garshelis DL (2000) Delusions in habitat evaluation: Measuring use,
selection, and importance. In: Boitani L and Fuller TK (eds.) Research
Techniques in Animal Ecology: Controversies and Consequences,
pp. 111164. New York: Columbia University Press.
Krebs CJ, Boutin S, and Boonstra R (eds.) (2001) Ecosystem Dynamics
of the Boreal Forest: The Kluane Project. New York: Oxford
University Press.
Leopold A (1933) Game Management. New York: Charles Scribner
Sons.
Manel S, Schwartz MK, Luikart G, and Taberlet P (2003) Landscape
genetics: Combining landscape ecology and population genetics.
Trends in Ecology and Evolution 18: 189197.
Wind Effects
W Eugster, ETH, Zurich, Switzerland
2008 Elsevier B.V. All rights reserved.
Introduction
Mean Wind Speed and Turbulence
Wind over the Vegetated Surface
Changes in Surface Roughness: The Edge Effect
Turbulent Mixing and Trace Gas Exchange
Dispersal of Pollen, Spores, and Microorganisms
Influence on Small Animals and Seed Dispersal
Introduction
There are two categories of wind effects in ecology:
(1) the effect of the vegetation surface on the wind, how
it lowers wind speed near the ground, shelters niches from
strong winds where small animals and plants can establish
and live; and (2) the effect that wind and turbulence
excert on many aspects of animal behavior, plant growth
and survival, and the overall metabolisms of organisms.
Studies on forest recovery in North America have
pointed to the important role of high winds in temperate
forests. Although such catastrophes are rare, they could
be instrumental in the creation and maintenance of
mosaic patterns and hence the diversity of these woods.
Some attempts to reconstruct the history of winds during
the past glacial maximum (about 18 000 years ago) indi-
cate that tropical storms generating winds of hurricane
force were scarser, less intense, and shorter than those of
the present day, with important consequences for forest
ecology which include the influence on development,
structure, and composition of the migrating and reassem-
bling forests of the mid- and higher latitudes. However,
direct evidence of the effects of wind on forests is hard to
Morrison ML, Marcot BG, and Mannan RW (2006) WildlifeHabitat
Relationships: Concepts and Applications. Washington, DC: Island Press.
Powell RA (2000) Animal home ranges and territories and home range
estimators. In: Boitani L and Fuller TK (eds.) Research Techniques in
Animal Ecology: Controversies and Consequences, pp. 65110.
New York: Columbia University Press.
Sinclair ARE, Fryxell JM, and Caughley G (2006) Wildlife Ecology,
Conservation, and Management, 2nd edn. Malden, MA: Blackwell
Publishing.
Vucetich JA and Peterson RO (2004) Long-term population and
predation dynamics of wolves on Isle Royale. In: Macdonald D and
Sillero-Zubiri C (eds.) Biology and Conservation of Wild Canids,
pp. 281292. New York: Oxford University Press.
Waits LP and Paetkau D (2005) Noninvasive genetic sampling tools for
wildlife biologists: A review of applications and recommendations for
accurate data collection. Journal of Wildlife Management
69: 14191433.
Influence on Bird and Insect Migration
No Wind Effect?
Wind Chill and Heat Index
Metabolic Stress by Wind
Wind Throws and Wild Fires
Further Reading
come by, and also other aspects of wind effects in a wide
variety of ecologically relevant topics are rather scarcely
covered in the scientific literature. Almost all studies
reviewed in this article are based on an ecological ques-
tion that suggest some partially unknown dependence on
environmental variables. Mostly, temperature, precipita-
tion, and humidity are considered important variables in
such investigations, and wind effects are rather considered
a possible or likely additional side effect of the overall
ecological process under investigation. It is therefore not
surprising that there are almost no systematic studies in
the scientific literature that cover all aspects of wind in all
details.
Some specific aspects, where the ecological impor-
tance of the wind is rather obvious, are covered in much
greater detail in other topics (see Wind Shelterbelts).
Thus, here we focus on the very general physical rela-
tionship between wind (and thus turbulence) and other
environmental factors such as thermal heat loss and meta-
bolic rates of organisms, and the exchange of trace gases
such as CO2 shall be addressed before summarizing the
most relevant specific aspects of wind effects in the eco-
logical literature.
 Ecological Processes | Wind Effects 3795

Mean Wind Speed and Turbulence

Wind is a vector variable, but in many scientific applica-

tions only the scalar wind speed is investigated or of
interest. Since wind that is the term for the atmospheric Laminar
motion over the solid surface of the Earth with respect to sublayer
the surface itself is mainly driven by pressure gradients Laminar flow Turbulent flow
on relatively large scales over the globe, this term is often Distance from leading edge
used as a short-cut for horizontal mean wind speed. Over
sufficiently long observation intervals and over large sur- Figure 1 Transition from laminar to turbulent boundary layer as
wind blows over a vegetation surface changing from smooth to
face areas, there is no net loss or gain of air due to vertical
rough. Modified from Grace J (1977) Plant Responses to Wind.
motion; thus, the vertical component of the wind vector is London: Academic Press.
considered to be zero. Hence, the wind vector is approxi-
mated as a two-dimensional entity that can be described
by the scalar horizontal wind speed and the wind direc-
turbulent kinetic energy dissipation that ends up in the
tion. All standard weather stations use this basic concept
Brownian motion of single molecules and thus dissipates
for recording wind. This is however not necessarily the
kinetic to thermal energy.
best possible simplification for small-scale and short-term
Wind has a kinetic energy,
investigations, and thus has important implications for
ecological processes. As an example, three-dimensional m 2
Ek V 1
wind gusts in autumn can easily pick up fallen leaves from 2
the ground, despite the fact that the leaves are relatively with m the unit mass of the unit air volume, and V
heavy and the mean vertical and horizontal wind speed the scalar wind speed of the three-dimensional wind
over an hour or longer may be rather low. But on the vector, that is composed of mean kinetic energy and
timescale from tenths of a second to several minutes, turbulent kinetic energy. At moderate to high wind
turbulence which includes such wind gusts can be speeds the mean kinetic energy is by far greater than
the most relevant wind effect. The turbulent timescale the turbulent kinetic energy. In addition, wind carries a
typically extends up to 1 h, whereas longer timescales momentum,
are associated with mean wind effects. There is not a
sharp separation between turbulence and mean wind,  mV 2
although a spectral gap between the two timescales has
been postulated by some scientists. In reality, there is a
confounding effect with the diurnal course of wind speeds
that show different and locality-specific conditions during Turbulent
the day as compared to the night. boundary
layer

Laminar Flow and Turbulent Winds Laminar

sublayer
Turbulence is generated inside a laminar flow when there
ew
is mechanical friction or thermal convection that perturbs c
the flow. Figure 1 illustrates this for a laminar flow with a ec
certain wind speed that moves from a smooth onto a rough
surface. At a certain distance downwind of the leading edge
of this increased roughness, the laminar flow becomes sc
chaotic, that is, turbulent. However, this turbulence does
not completely reach the surface, a minute laminar surface
layer always exists, over the surface of any object, including gc 20 m
s
plant leaves (Figure 2). Turbulent exchange of heat, moist- mc
ure, CO2, and other trace gases is by far more efficient than p
exchange in laminar flows (see below). Once the air is
turbulent the flow does not easily become laminar again Figure 2 Diffusion pathways at a leaf surface on a windy day: c,
cutin; ec, epidermal cell; ew epicuticular wax; gc, guard cell; mc,
since the transition from turbulent to laminar flow is not as mesophyll cell; p, pore; s, substomatal cavity; sc, subsidiary cell.
clearly defined at it is in the opposite direction. The decay Modified from Grace J (1977) Plant Responses to Wind. London:
of turbulence in the air is subject to the physical rules of Academic Press.
3796 Ecological Processes | Wind Effects
At high wind speeds, especially during storms, a
very high kinetic energy (both mean and turbulent com-
ponents increase with increasing wind speed) may result
from the wind, which is responsible for the devastating
damages by hurricanes and other storm events, that how-
ever are also important for producing gaps (wind throws)
in forest ecosystems and thus for these ecosystems overall
life cycle. Turbulent motions that are most relevant at
lower wind speeds do not have such a devastating effect
when mean wind speed is low. The kinetic energy of the
mean wind is what wind mills profit from, and also
migratory birds benefit from this component. Near the
ground, the vegetation has to absorb both the kinetic
energy of the wind and its momentum, but in this case
the momentum absorption is by far the more relevant
process and as a first approximation only momentum
transfer by the vegetation is considered, neglecting the
additional effect of energy absorption.
For flying birds it is mostly the kinetic energy of the
wind that influences their daily life. It has been shown
for Sandwich Terns (Sterna sandvicensis) on the isle of
Griend, the Netherlands, that their loss of prey to the
competing kleptoparasitizing Blackheaded Gulls (Larus
ridbundus) significantly increases with wind speed. Thus,
wind directly reduced the ability of Sandwich Terns to
defend their prey (mostly fish) against attacks of
Blackheaded Gulls. This had a negative effect on the
amount of food transported to the colony, while klepto-
parasitism increased. Therefore, wind speed severely
affected energy intake of the chicks and had strong
negative effects on chick growth. During the first two
weeks posthatching, kleptoparasitism was relatively low
and had only small effects on chick growth, even under
unfavorable weather conditions. From then on, the nega-
tive effects of kleptoparasitism on growth became
considerable.
Table 1 Relations between canopy heights (m) and aerodynamic roughness length (m) for different vegetation types
Vegetation Type
Forest Tropical
Coniferous
Pine
Woodland Trees
Savannah
Crops Vines
Beans
Corn
Wheat
Wheat stubble
Grass Thick/thin
Sparse
Soil Bare
After Garratt JR (1992) The Atmospheric Boundary Layer. Cambridge: Cambridge University Press.
Wind over the Vegetated Surface
Vegetation is the most important interface between the
atmosphere and the solid ground in terrestrial ecosystems.
On the one hand, vegetation adapts to wind conditions
and special plant social community compositions are
found in the Arctic and Alpine environments where per-
sistant and strong winds influence exposed locations such
as hills, mountains, and crests. On the other hand, vegeta-
tion makes the Earths surface rougher than what would
be the case over bare soil (Table 1), and thus strongly
influences the wind speed (Figure 3) and direction in the
atmosphere near the ground. The wind is driven by
pressure and temperature differences on large scales,
whereas the Earths surface does not move and becomes
stationary under most occasions. Exceptions are very
special conditions during hurricane-force winds, and cer-
tain exposed locations with corresponding soil conditions,
where bluffs are created by the steady wind movement.
Under normal conditions, wind speed at some nonzero
height above the ground must be zero to fulfill the criter-
ion that the vegetation stays in place. Rough vegetation
such as forests excert a much higher roughness (1 m) to
the atmospheric wind motion than shortcut grass (on the
order of millimeters to centimeters; see Table 1). Based
on this roughness the increase in wind speed with height
above the vegetation depends strongly on vegetation type
and structure. This vertical wind speed profile tends to
increase logarithmically with height above the canopy
(Figure 3), and the physical process responsible for this
is momentum absorption. Tall vegetation such as forests
absorb momentum with their roots, and also in the
dynamic motion of the stems. Thus, under strong winds
it depends on the rooting type of the tree and the wood
quality whether a tree can be uprooted or whether the
stem breaks at a certain height above the ground.
Canopy height (m) Roughness length (m)
3235 2.24.8
10.427.5 0.283.9
12.415.8 0.320.92
1015 0.4
89.5 0.40.9
0.91.4 0.0230.12
1.18 0.077
0.8 0.064
0.25/0.4/1.0 0.005/0.015/0.05
0.18 0.025
0.1/0.5 0.023/0.05
0.025/0.015/0.45/0.65 0.0012/0.002/0.018/0.039
0.0010.01
 Ecological Processes | Wind Effects 3797

(a) 1.6 (b) based on pH readings along a transect from the shore of
Forest Maize the Belcher Islands off the Hudson Bay coast, and it is also
1.2
noted that the drying effect of the wind, possibly in com-
z /h

z /h
0.8 bination with salt spray and other factors (ice and sand
particles), may be as important.
0.4

0
0 0.4 0.8 1.2 1.6 0 0.4 0.8 1.2 1.6
u(z)/u(h) u(z)/u(h) Directional Growth Response

Figure 3 Wind profiles (a) in a pine forest canopy of 16 m height Besides pruning steady winds from a persistent wind
(h), and (b) in a maize canopy of 2.1 m height. Both profiles show direction can lead to directional growth response of
the mean horizontal wind speed, normalized for the wind speed trees, it is widely observed in coastal areas, in deep
at the top of the canopy (z h). In the case of the forest the mountain valleys with a well-developed valley wind sys-
specific wind profile inside the trunk space (z/h < 0.5) a secondary
tem (Figure 5), or on wind exposed crests, rims, and hills.
maximum of the wind speed can be seen. Maize profiles for light
winds of 0.88 m s1 (closed triangle) and strong winds of Since wind speeds generally increase with altitude from
2.66 m s1 (open triangle) at the top of the canopy are shown. the lowland to the mountains, it has even been argued that
From Raupach MR and Thom AS (1981) Turbulence in and above high-altitude plants in wind-swept mountains may be less
plant canopies. Annual Review of Fluid Mechanics 13: 97129. affected by global warming, and that the spread of low-
land plant species into uplands as predicted by some
Wind Pruning and Salt Spray
To unroot a tree normally requires strong gusts in heavy
mean winds, as for example during storms. If winds are
strong and steady, but not very gusty, then the energy may
not be sufficient to unroot a tree and thus the wind-pruning
effect may shape trees and shrubs (Figure 4). Along the
seacoast of British Guiana, along the subtropical shores of
the island and Trinidad and southern California, and the
subarctic shores of Hudson Bay and Labrador, the wind is
reported to result in pruned trees. It appears that the steady
subtropical winds have a similar pruning power as the icy
blasts of the subarctic. However, this is not necessarily an
effect of the wind alone. It has been argued that the proxi-
mity to the sea leads to a high load of salt spray in the wind, Figure 5 Trees growing under conditions with persistently high
wind speeds from a specific direction retain their asymmetric
and that the toxicity of that salt may be the true ecological
shape even when there is no wind. In this example from near
factor of wind pruning. Salt spray deposition on young Zweisimmen, Switzerland, the daytime up-valley wind (from right
shoots seems to actually kill many of them, thus causing to left) shaped the characteristic habitus of these trees.
the pruning. This observation is however only weakly Photograph by Werner Eugster.

Figure 4 Wind-pruning effect on Metrasideros polymorpha trees of a cloud forest on the Big Island of Hawaii (left). Under the strong
onshore winds, leafs are detached from branches until only clusters of leaves at the outer margin of the tree volume remain (right).
Photographs by Werner Eugster.
3798 Ecological Processes | Wind Effects
global warming scenarios may be strongly restricted in
higher altitudes due to the lack of adaptation of lowland
plants to such steady and comparatively strong winds.
Changes in Surface Roughness: The Edge
Effect
Sharp edges of vegetation which are less abundant and
less pronounced in natural ecosystems than in anthropo-
genically disturbed and shaped ecosystems such as
agroecosystems and managed forests are subject to
wind effects that depend strongly on the distance to the
change in roughness. When the wind first blows over a
smooth (e.g., grass) surface and then abruptly has to
change to a rough (e.g., agricultural crop or forest) surface,
additional turbulence is created within a relatively short
distance as wind passes over this roughness change
(Figure 6). This additional turbulence carries extra
momentum that has to be absorbed by the vegetation
downwind to obtain a new equilibrium with the rougher
surface. This leads to the phenomenon that in a wheat
field for example there may be a few rows of plants
directly at the roughness change that seem quite unaf-
fected even by strong winds, while only 1 m downwind
one or several rows may be completely flattend by this
additional momentum. In the case of forests, wind throw
often excludes the trees at the forest edge, partially due to
the same phenomenon. But trees also can adapt to
1.0
Height (m)
0.1
0.01
0.5 0.6 0.7 0.8 0.9 1.0
Wind speed (relative scale)
Figure 6 Effect of a change from smooth to rough terrain.
Fetch that is the distance of uniform surface in the upwind
direction over the rough terrain was: (), 0.32 m; (), 1.18 m; (),
2.32 m; (
), 6.42 m; (), 16.42 m. Modified from Grace (1977)
after Bradley (1968), Quart. J. Roy. Meteorol. Soc. 94, 361379.
constant wind pressures by building special cells to coun-
teract this pressure. This is best known for trees in
mountain valleys and along seashores with persistant
and sufficiently strong winds in specific directions. In
mountain valleys these are the up-valley (daytime) and
down-valley (nighttime) wind directions. Which one is
stronger depends on the complex combination of orienta-
tion of the valley, length, topographic differences in the
surroundings, and more. But by studying trees which are
leaning in the direction of the dominant strong winds
(Figure 5), it is easy to determine the locally dominant
wind system. Near costs it is the diurnal sea breeze that
dominates wind pressure on trees, while the nocturnal
land breeze is in most cases much weaker.
On smaller scales, linear landscape elements such as
hedgerows, tree lines, and tree lanes are ecologically
important surface roughness elements, especially in
otherwise rather smooth agricultural landscapes. In the
Netherlands, for example, it has been shown that among
other possible functions (orientation clues, foraging habi-
tat) such linear elements provide shelter from wind and/
or predators for the two bat species Pipistrellus pipistrellus
and Eptesicus serotinus.
Turbulent Mixing and Trace Gas Exchange
Turbulent exchange is roughly three to four orders of
magnitude more efficient than diffusive mixing in a lami-
nar airflow. For trace gas exchange between the
atmosphere and the plants, the tiny laminar layer sur-
rounding each leaf (Figure 2) is thus non-negligible.
Given this huge difference in effectiveness of turbulent
versus diffusive transport a laminar boundary layer of
0.11 mm provides a similar resistance against the free
exchange of CO2 between the atmosphere and the plant
stomates as does 1 m of turbulent air. In Figure 2 it is
clearly seen that the laminar layer separates the turbulent
atmosphere where CO2 is available in vast quantities
from the stomatal opening and the substomatal cavity, the
buffer from where CO2 is used for photosynthesis. Any
changes in turbulence, wind speed, and wind direction
will also affect the thickness of this laminar boundary
layer and thus have an effect on the exchange of trace
gases, heat, and momentum between plants and the atmos-
phere. Figure 7 shows that depending on plant leaf shape,
the laminar boundary layer and thus the wind speed profile
at varying distances from the leaf surface show a relatively
large microscale variation.
1.1
On much larger scales, as wind blows over oceans and
open water, it induces mixing of the surface layer, thereby
enhancing the exchange of gases across the water surface,
which is important for the oxygen content in the water
and uptake or release of CO2 and CH4 to and from water
bodies. Similar mixing occurs in the air above the surface

(a) Wind speed (m s1)
0 1.0 1.5
Wind direction
(b) Turbulence
0 0.5
30
Vertical distance (mm)
20
15
10
5 (9.81 m s2), cw the friction coefficient (1 for circular
10 20 30 40 50
Horizontal distance (mm)
Figure 7 The boundary layer over a Populus leaf. Profiles of
(a) mean wind speed and (b) turbulence, shown in transverse
sections in a laminar free stream. Modified from Grace J (1977)
Plant Responses to Wind. London: Academic Press. After Grace
and Wilson (1976) Journal of experimental Botany 27, 231241.
which reaerates the plant canopy and essentially is
responsible for resupplying photosynthetically active
plants with CO2 from the atmosphere, while at the same
time O2 produced by plants is carried away and mixed
into the surface layer of the atmosphere.
Evaporation and Transpiration
A widely investigated topic of wind effects on ecosystems
not covered in this article is found in the hydrological and
biophysical literature on evaporation of water from eco-
systems and transpiration from plants, either as a
component of the hydrological cycle (the viewpoint
taken by ecohydrologists), or in combination with CO2
exchange (the ecophysiological viewpoint). See
Evapotranspiration for more information.
Dispersal of Pollen, Spores, and
Microorganisms
The explosive pollen release from many wind-pollinated
plants, particularly tree species with copious pollen pro-
duction, is triggered by moderately gusty winds. Similarly,
spores from the Swiss fern Asplenium ruta-muraria are
released either by wind-induced shaking of the leaves
(ballanemochory) or by the physical energy of impacting
raindrops. In some palms (Chamaedorea pinnatifrons ( Jacq.)
Ecological Processes | Wind Effects 3799
Oerst. and Wendlandiella sp.) in Peruvian Amazonia the
release of the pollen is triggered by movements of insects
inside the flowers, and the term insect-induced wind pol-
lination has been suggested since these insects do not
normally also visit the female flowers of these palms.
On the ground, a wind gust can pick up small dust
particles, sedimented pollen and microorganisms such as
bacteria and mites from the surface or host organism. Once
in the air, moderately turbulent winds are already sufficient
to keep such small biotic and abiotic objects aloft. The
general concept of updraft of a voluminous body in the
atmosphere is described by Stokes law of sedimentation,
where the terminal falling velocity Vt of an object is
r
2mg
Vt 3
cw A
with m the mass (kg m3), g the gravitational acceleration
bodies, <1 for aerodynamically formed bodies),  the
density of the air (1.2 kg m3 at sea level), and A
the projected surface of the body (m2). Figure 8 shows
the terminal fall velocity for small organisms of 1 mm to
2.5 mm and how typical vertical wind speeds in the air can
counteract the falling of such objects, once they are
dispersed in the air. In this respect wind has almost
exactly the same effect as the water flow in rivers: under
high turbulence and horizontal speeds animals may find
sheltered spots where they are not picked up by the
motion of water or the air, but once they lose adhesive
contact, their body size and weight may be too small to
grasp ground again, and they become suspended in the
fluid until they happen to end up in a calmer area where
their settling velocity is greater than the wind (or water)
motion, which allows them to reach the ground surface
again. Figure 8 shows that bodies with a diameter smaller
than 10 mm are normally too small to return to the
5.0 Typical mean updrafts in convective clouds
Velocity (m s1)
2.0 ty
oci
vel
1.0 lfall
m ina
0.5 Ter
0.2
Typical turbulent updrafts
0.1
1 5 10 50 100 500 1000
Aerodynamic diameter (m)
Figure 8 The terminal fall velocity of ball-shaped objects
(pollen, seeds, bacteria, microorganisms) compared to typical
updrafts in the turbulent atmosphere (up to 0.2 m s1) and
typical mean updrafts in convective clouds (thunderstorms). The
thick line applies to objects that have a similar density as water. If
updrafts are stronger than the terminal fall velocity, then an
organism or particle in the atmosphere remains suspended in the
atmosphere and will only deposit on obstacles such as trees due
to impaction. Larger organisms that are subject to a high terminal
fall velocity need active means to keep themselves aloft.
3800 Ecological Processes | Wind Effects
ground in the turbulent atmosphere. Thus, for such small
bodies, impaction becomes the most relevant process how
they can be eliminated from the atmosphere, that is, when
they physically hit the surface of a tree or another plant.
The sticky stigma of a flowers pistil further helps to
capture pollen even when impaction is weak.
Wind pollination is considered inefficient compared
with insect pollination. This finding has led to the
hypothesis that the rise to dominance of the angiosperms
over gymnosperms at evolutionary timescales is due to
reproductive innovations, especially those involving co-
evolution with biotic gene dispersers. This has most likely
contributed to the present-day situation that conifers are
biogeographically restricted to stressful environments
where gymnosperms may suffer a comparative disadvan-
tage if pollinators face persistently high wind speeds.
Influence on Small Animals and Seed
Dispersal
Small insects need to adopt to wind speeds. Studies carried
out in a wind tunnel indicate that weak winds >0.2 m s1
already have an effect on the flight and landing behavior of
the bug Prostephanus truncatus (Horn). In the open land-
scape, mosquitoes (Anopheles marajoara in Brazil) can only
freely navigate in air with wind speeds below about
0.85 m s1 (3 km h1). From the aphid parasitoid Aphidius
nigripes, it is reported that males generally did not reach
females at wind speeds of 1.0 m s1, as the majority of
individuals taking flight in the pheromone plume (81.8%)
was unable to sustain upwind flight. The general picture is
that as wind speed increases these small animals increas-
ingly lose control over their flight trajectory and may no
longer target their prey or mate as desired. Swallows, for
example, are known to fly close to the ground before
thundershowers, where the prefrontal increase in wind
speed restricts the activity of small flying insects to the
few lowest meters close to the ground. In summer, when
mosquitoe abundance is enormous in the Arctic, reindeer
and caribou select windy locations for resting and rumina-
tion, preferably close to the seashore, on gravel pads in
large rivers with a well-developed diurnal valley wind
system, or on snow fields with thermotopographic wind
resulting from the contrasting surface temperatures
between snow and vegetation or rocky surfaces.
Some spider species benefit from this effect by letting
themselves drift away termed ballooning in the scientific
literature to explore new habitats using a short thread
that increases their updraft and thus drifting distance at
higher wind speeds. Although there are contradictory
views between authors about the importance of various
environmental factors, it is widely agreed upon the upper
wind speed limit of 3 m s1 for ballooning. Most work has
been focusing mostly on the meteorological conditions at
the time of take-off, whereas literature on the underlying
motivation of the spiders and instigation of pre-ballooning
behavior (climbing to a prominent point and silk release) is
very limited and largely considered supposition by some
experts. Since spiders are important polyphagous predators
on arable farmland, the high mobility of ballooning species
means that they are often the first to arrive in a crop newly
infested with pests, and have a role in controlling the out-
break until more specific predators arrive.
In a similar way as ballooning spiders take advantage
of the horizontal translocation by wind plants profit from
the wind to disperse their seeds. The parachute type seeds
of Asteraceae and the winged seeds of Acer, Fraxinus, Ulmus,
and many coniferous trees are good examples of how
plants benefit from available wind to spread out faster
than would be possible without the help of the wind. For
seeds that do not have wings, hairs, or parachute-type
annexes, the Stokes settling velocity (eqn [3], Figure 8)
applies and explains why in general small seeds are wind
dispersed because of their long residence time in the
atmosphere (the residence time is inversely proportional
to the terminal fall velocity) than large seeds, that lead to
small dispersial kernels downwind of the seeder plant
unless the seeds are dispersed by animals. A special case
exists for vegetation near open water bodies, where large
buoyant seeds can float on the water and be dispersed by
its currents, such that even large and heavy seeds can be
transported over long distance that would not be possible
by the wind alone.
Nature has brought about a wealth of shapes and
forms of seeds that do not correspond with the simplest
version of a spherical with cw 1. For some plants,
specific wind tunnel studies have been carried out to
determine the true dispersal capacity of seeds. For six
Canadian perennial grassland species with different seed
aerodynamic attributes, it was investigated how dispersal
distances vary with varying wind speeds and release
heights. Dispersal distances of long-range dispersed
seeds (99th percentile values) increased exponentially
with wind speed. At wind speeds of 14 m s1, predicted
maximum distances were 1015m for small and rela-
tively heavy spherical seeds and 2030 m for large and
relatively light cylindrical or disk-like seeds. In the study
area, wind gusts >10 m s1 at plant height occur at least
annually, and plants of the selected species live up to
several decades. This suggests a great potential for long-
range dispersal during the lifetime of a plant. It is argued
that plants may gain wider dispersal of seeds by increas-
ing the release height (e.g., taller infructescences) and by
requiring stronger winds to release seeds (e.g., dispersal
in autumn and winter).
Transport distance is one aspect, the other is the release
of seeds from the flower heads by wind speed. In a wind
tunnel study with flower heads of two thistle species,
Carduus nutans and Carduus acanthoides, with ripe seeds,

the effect of laminar versus turbulent flows of increasing
velocity was investigated. Seed release increased with wind
speeds of both laminar and turbulent flows. However, far
more seeds were released, at significantly lower wind
speeds, during turbulent flows. In other cases, the seeds
are primarily dispersed by the wind, followed by secondary
dispersal by rodents living on the ground which collect the
seeds and cache them in the soil. Treatment by rodents,
primarily yellow pine chipmunks (Tamias amoenus), of four
species of pine seeds, lodgepole pine (Pinus contorta, 8.7 mg
seed weight), ponderosa pine (Pinus ponderosa, 55 mg),
Jeffrey pine (Pinus jeffreyi, 157 mg), and sugar pine (Pinus
lambertiana, 213 mg), that vary in size and weight was stud-
ied in the Carson Range of western Nevada. For the
species examined, seed size appeared to have had little
effect on several other attributes, including mean dispersal
distance, substrate choice, and microhabitat choice. It was
found that although a larger seed size and weight decreases
primary wind dispersibility of pine seeds, the secondary
dispersal by scatter-hoarding rodents compensates for poor
wind dispersal so that total dispersibility of large-seeded
pines is not compromised.
Influence on Bird and Insect Migration
At a much larger scale many long-distance migratory bird
species have adopted flight tracks that best profit from
large-scale wind fields on the Earth, which saves energy
and thus increases the survival rate. On the other hand,
migratory birds that are facing strong headwinds, may
suffer severe losses if such an occurrence combines with
low temperatures, scarce food resources or the like. In
general, the nocturnal and diurnal wind directions and
speeds are not necessarily the same. Near the ground the
so-called low-level jet, a relatively strong wind with its
maximum speed at only 100300 m above the ground
surface is active at night, whereas the atmosphere may
be calm during the day, and wind speeds are higher aloft.
In a study on the migration patterns and environmen-
tal effects on stopover of monarch butterflies at Peninsula
Point, Michigan, it was found that wind direction had a
significant influence on the number of monarchs recorded
on each count over a 7-year period, with higher counts
during north winds.
On smaller scales, it has been shown for two dragon fly
species, Pantala hymenaea and Pantala flavescens, in natural
flight over a lake at ambient wind speed and direction,
that they are able to compensate at least partially for
crosswind drift, which shows evidence for use of a ground
reference to correct for drift when flying over water, and
their ability to cope with much higher wind speeds
(5.0 m s1) than small insects are able to.
Ecological Processes | Wind Effects 3801
No Wind Effect?
Although there are many studies that found ecological
effects of wind on animals, plants, and ecosystem pro-
cesses, it should be remembered that there are other
studies that were unable to find such effects. For example,
the bat Pipistrellus pipistrellus in Oxfordshire did not show
an apparent response to wind nor rain in the time spent
outside the roost. This is remarkable since it feeds on
insects and one might expect a behavior similar to the
one known from swallows. Since it is generally difficult to
publish negative results in the scientific literature, and
moreover wind as a three-dimensional vector variable
makes it particularly challenging to derive the relevant
information from simple measurements (e.g., if mean
wind speed was measured when actually turbulent kinetic
energy would have been the variable with higher predic-
tive power), the lack of clear statements on where wind
does not have an effect should not come as a surprise.
In forest ecology it has been postulated that there are
two main factors why biotic effects of wind have not been
well studied: (1) the difficulty of measuring wind in the field
and separating its effects from the confounding variables of
temperature and humidity; and (2) the expense involved in
carrying out wind tunnel experiments in the laboratory.
Wind Chill and Heat Index
The bioclimatic temperature sensed by an organism can
differ considerably from the absolute physical tempera-
ture that is measured by conventional instruments. For
humans, elaborate concepts to compute a wind chill tem-
perature have been established to account especially for
the effect of wind. The concept bases on the knowledge
that increasing mean wind speeds increase also turbu-
lence, and thus the heat transport away from an
organism that has a warmer skin temperature than the
atmosphere. Although it is widely known that the ambient
moisture or humidity in the air has an additional influ-
ence, for the sake of simplicity most approaches only
consider wind speed as a specific factor when considering
wind chill.
Controlled experiments with humans were carried out
to determine the functional relationship between wind
chill and preceived temperature. This was not possible
with primates, where thermoregulation is known to be an
important ecological constraint. Shade temperatures, solar
radiation, humidity, and wind speed all serve to alter an
animals perceived temperature. In a recent review, three
thermal indices currently available were compared. Black
bulb temperatures can account for the effect of solar radia-
tion, with wind chill equivalent temperatures and the heat
index providing quantifiable estimates of the relative
3802 Ecological Processes | Wind Effects
impact of wind speed and humidity, respectively. The
authors presented three potential indices of the perceived
environmental temperature that account for the combined
impact of solar radiation, humidity, and wind speed on
temperature, and performed a preliminary test of all of
the climatic indices against behavioral data from a field
study of chacma baboons (Papio cynocephalus ursinus) at De
Hoop Nature Reserve, South Africa. It was found that the
complexity of the interactions among environmental fac-
tors that influence thermoregulation in primates will
require the development of biophysical models of the
thermal characteristics of the species and its environment.
Until such models are developed, however, it is concluded
that wind chill and heat indices should permit a more
detailed examination of the thermal environment, allowing
thermoregulation to be given greater precedence in future
studies of primate behavior.
Another widely established approach is not to try to
compute a bioclimatic temperature or index, but relate
the metabolic energy consumption of an animal to envir-
onmental factors.
Metabolic Stress by Wind
Small animals can profit from the presence of a laminar
sublayer (Figure 1) even under highly turbulent condi-
tions. Due to the much lower heat exchange in that laminar
layer, they may avoid metabolic stress under high winds.
This is almost impossible for larger animals, such as breed-
ing arctic shorebirds. It was found that tarsus length in all
shorebirds breeding in the Canadian arctic shows an evo-
lutionary response to average metabolic stress encountered
across the breeding range, such that birds nesting in meta-
bolically stressful environments have relatively shorter
legs. Longer-legged birds living in colder environments
will experience greater metabolic costs because their torsos
are elevated farther away from the grounds wind-dampen-
ing boundary layer. It was suggested that the widely known
Allens rule that relates the metabolic rate of an organism to
its volume should be extended: body-supporting appen-
dages of homeotherms may be shorter in colder
environments so as to take advantage of a boundary layer
effect, thereby reducing metabolic costs.
Another study that investigated the effects of water
levels and weather on wintering herons and egrets found
that larger and longer-legged species tended to be found in
deeper water, although both species frequently were found
together in shallow water. Severe weather with high winds
caused the birds to suspend foraging and remain sheltered
from the wind. Consequently, a higher percentage of smal-
ler heron and egret species did not survive severe storms
since searching shelter from wind meant fasting. A 3-day
storm period was simulated to lead to >10% decline in
body mass of the smaller herons and egrets.
Wind Throws and Wild Fires
Extreme events with high wind speeds are important in
the life cycle of many ecosystems, especially forests.
Hurricanes in the tropics, tornadoes, and other windstorms
further north and south reshape forest ecosystems via wind-
throws that eliminate the weakest and thus most often the
oldest individuals in the forest canopy. For example, in New
England forests, leaning is the most prevalent damage to
young stands, whereas breakage and uprooting dominated in
older stands. Breaking was slightly more important in older
conifer than hardwood stands, comprising 614% of the
stems and generally occurring 15 m from the ground, but
numbers vary not only widely between species and stand
composition but also among storm events in the same stand.
Since heavy storms are often accompanied with severe
lightning strikes, the wind effect can easily be a combination
of wind and fire. When a wildfire starts then the wind
conditions will strongly determine how quickly the fire
advances with the wind, and what damage is done to the
ecosystem. In some cases, such as the Bishop pines, the fire
even is necessary to free the seeds in the cones and initiate
the life cycle of this forest type. In the gaps the new vegeta-
tion can resprout, and since more light and precipitation
reaches the ground, this provides niches and living space for
early successional plants. As a consequence also the fauna
may be affected. Organisms with a life size that is much
smaller than gaps in forests may only find a suitable ecolo-
gical niche in the gaps that shift their location over the years.
In subalpine forests of the Swiss alps, it was found that the
gaps created by windthrows add considerably to the species
diversity of macrofungi. Larger animals such as black bears
in southeast Alaska were found to react in just an opposite
way: 58% of the den sites were found in forests that were
most protected from catastrophic storm effects, and only 6%
in forests most exposed to storm damage. These results
suggest that the effect of catastrophic windstorm disturbance
on overwinter habitat for black bears is the key factor
influencing the site selection for black bear dens.
See also: Climate Change 1: Short-Term Dynamics;
DispersalMigration; Estuarine Ecohydrology; Fire;
Reaeration; Wind Shelterbelts.
Further Reading
Cartar RV and Morrison RIG (2005) Metabolic correlates of leg length in
breeding arctic shorebirds: The cost of getting high. Journal of
Biogeography 32: 377382.
de Gayner EJ, Kramer MG, Doerr JG, and Robertsen MJ (2005)
Windstorm disturbance effects on forest structure and black bear
dens in southeast Alaska. Ecological Applications 15: 13061316.
Doutt JK (1941) Wind pruning and salt spray as factors in ecology.
Ecology 22: 195196.
Ellenberg H and Strutt GK (1988) Vegetation Ecology of Central Europe.
Cambridge: Cambridge University Press.

Ennos AR (1997) Wind as an ecological factor. Trends in Ecology and
Evolution 12: 108111.
Foster DR (1988) Species and stand response to catastrophic wind in
central New England, USA. Journal of Ecology 76: 135151.
Garatt JR (1992) The Atmospheric Boundary Layer. Cambridge:
Cambridge University Press.
Geiger R, Aron RH, and Todhunter P (1995) The Climate Near the
Ground. Braunschweig: Vieweg.
Grace J (1977) Plant Responses to Wind. London: Academic Press.
Grace J and Wilson J (1976) The boundary layer over a populus leaf.
Journal of Experimental Botany 27: 231241.
Hill RA, Weingrill T, Barrett L, and Henzi SP (2004) Indices of
environmental temperatures for primates in open habitats. Primates
45: 713.
Moore PD (1988) Forest ecology: Blow, blow thou winter wind. Nature
336: 313313.
Myers RK and van Lear DH (1998) Hurricanefire interactions in coastal
forests of the south: A review and hypothesis. Forest Ecology and
Management 103: 265276.
Raupach MR and Thom AS (1981) Turbulence in and above plant
canopies. Annual Review of Fluid Mechanics 13: 97129.
Wind Shelterbelts
J-J Zhu, Institute of Applied Ecology, CAS, Shenyang, Peoples Republic of China
2008 Elsevier B.V. All rights reserved.
Introduction
Interactions between Wind and Trees
Shelterbelt Structures
Determination of Optical Porosity
Introduction
The quantity of falling solar energy and the proportion
that is absorbed by either the atmosphere or the surface
varies greatly from one place to another, which results in
regional variations in temperature both for Earths surface
and atmosphere. The variations of temperature cause a
difference in atmospheric pressures, which leads to the
movement of air from high-pressure area to low-pressure
area, that is, the wind. Wind moves horizontally, verti-
cally, and turbulently. It is affected by the conditions of
the surfaces it encounters. The surface wind influences
the habitats for wildlife, the growth of crop and livestock,
soil erosion, snow distribution, sand-blowing, etc., and
causes extreme damages when it is very strong.
Shelterbelt is always called windbreak (some authors
distinguish the usage between the two terms based on
their objectives; here no distinction is adopted), which
can be defined as the barrier used to reduce wind speed.
It usually consists of trees and shrubs, or even perennial
or annual crops, wooden fences or other materials
(Figures 1 and 2). Shelterbelts when they are reasonably
designed can provide large areas of reduced wind speed
because they increase the roughness. The areas of
Ecosystems | Wind Shelterbelts 3803
Senn-Irlet B and Bieri G (1999) Sporocarp succession of soil-inhabiting
macrofungi in an autochthonous subalpine Norway spruce forest of
Switzerland. Forest Ecology and Management 124: 169175.
Stienen EWM, Brenninkmeijer A, and Geschiere CE (2001) Living with
gulls: The consequences for Sandwich Terns of breeding in
association with black-headed Gulls. Waterbirds 24: 6882.
van Dorp D, van den Hoek WPM, and Daleboudt C (1996) Seed
dispersal capacity of six perennial grassland species measured in a
wind tunnel at varying wind speed and height. Canadian Journal of
BotanyRevue Canadienne de Botanique 74: 19561963.
van Gardingen P and Grace J (1991) Plants and wind. Advances in
Botanical Research 18: 189253.
Vonlanthen CM, Kammer PM, Eugster W, Buhler A, and Veit H (2006)
Alpine vascular plant species richness: The importance of daily
maximum temperature and pH. Plant Ecology 184: 19.
Weyman GS (1993) A review of the possible causative factors and
significance of ballooning in spiders. Ethology, Ecology and Evolution
5: 279291.
Woodward FI (1993) The lowland-to-upland transition modeling plant-
responses to environmental change. Ecological Applications
3: 404408.
Wind Profiles near Shelterbelts
Establishment and Management of Shelterbelts for
Wind Protection
Further Reading
reduced wind speed, especially in windy regions, are
generally called sheltered zones, which are very useful
for wildlife, agriculture, and for the people suffering from
the severe climates. In fact, there are many ecological
functions such as protecting against erosion, improving
crop production, filtering air and water, ameliorating cli-
matic extremes, improving ecological environmental
qualities, reducing and ameliorating potential conflicts
Figure 1 Farmland shelterbelt.