Plant pollination by animals is an essential eco-

system service. It is estimated that 60 to 80

percent of the worlds 250,000 flowering
plants depend on animalsmostly insectsfor
pollination (Kremen et al. 2007). Eighty-seven
of the worlds 124 most commonly cultivated
crops are animal pollinated, and insect-
pollinated forage plants such as alfalfa and clo-
ver provide feed for livestock. Roughly thirty-
five percent of global crop production is de-
pendent on pollination by animals (Klein et al.
2006). Pollinators also sustain wildland plant
communities that provide food and shelter for
myriad other wildlife. As one of the most
widespread and important ecosystem services
in terrestrial environments, plant pollination by
insects is essential to human health, global
food webs, and protection of biodiversity.
Annas northern blue. Photograph Kim Davis and Mike Stangeland (www.kimandmikeontheroad.com)
Written by
Scott Hoffman Black,
Nathan Hodges,
Mace Vaughan, and
Matthew Shepherd
The Xerces Society
for Invertebrate
Conservation


www.xerces.org
INVERTEBRATE CONSERVATION FACT SHEET
Pollinators in Natural Areas
A Primer on Habitat Management

Despite the recognized importance of pollina-
tion services, there is a growing body of evi-
dence that suggests pollinators are at risk. In
the United States, the National Research Coun-
cil (2006) reported that both managed honey
bee colonies and wild pollinators showed sig-
nificant declines. The causes of decline are dif-
ficult to pinpoint, but loss of floral diversity
and habitat due to increasing urbanization, ex-
pansion of intensive agriculture, invasive
plants, widespread use of pesticides, climate
change, and disease and parasites have all had
a negative impact on pollinator populations
(National Research Council 2006). As pressure
on pollinators increases from human activities
and other factors, undeveloped habitat and
natural areas can play a substantial role as
long-term refugia for these animals.

Pollinators in
natural areas
support di-
verse plant
communities,
wildlife food
webs, and ad-
jacent farms.

Incorporating
pollinator
needs into a
site manage-
ment plan will
result in excel-
lent habitat for
wildlife of all
types.

2
POLLINATORS AND THEIR IMPORTANCE
Importance of Natural Areas for Native Pollinators and
Agriculture
Native pollinators clearly help with crop pollination, yet
many agricultural areas lack the habitat necessary to support
native pollinators. The role that natural habitat within the
agricultural landscape plays in providing crop pollination
services is increasingly well understood. Proximity to natu-
ral or seminatural land is often an important predictor of
pollinator diversity in cropland (Bergman et al. 2004; Kre-
men et al. 2004; Morandin & Winston 2006; Hendrickx et
al. 2007; Winfree et al. 2007b). For agricultural areas that
have lost native pollinators due to habitat modification or
pesticide treatments, adjacent natural areas provide two
valuable benefits. First, they are a source of pollinators for
crop pollination (Kremen et al. 2004). Second, they act as
refugia for pollinators that can recolonize degraded agricul-
tural areas (Ockinger & Smith 2007).
Along with on-farm habitat, nearby natural areas
are integral to maintaining long-term populations of native
pollinators in agricultural landscapes; however, it is impor-
tant that the management of these natural areas takes native
pollinators into account.

Importance of Pollinators to Natural Areas
By aiding in wildland food production, helping with nutri-
ent cycling, and as direct prey, pollinators are important in
wildlife food webs. For example, many migratory songbirds
require a diet of berries, fruits, and seeds from insect-
pollinated plants. Additionally, pollinator larvae are an im-
portant component of the diet of many young birds (Buehler
et al. 2002). Summerville and Crist (2002) found that forest
moths play important functional roles as selective herbi-
vores, pollinators, detritivores, and prey for migratory song-
birds. Belfrage et al. (2005) demonstrated that butterfly
diversity was a good predictor of bird abundance and diver-
sity, apparently due to a shared requirement for a complex
plant community.
Pollinator insects are a diverse component of the
wildlife of natural areas. They include butterflies and moths
(Lepidoptera), bees and wasps (Hymenoptera), beetles
(Coleoptera), and flies (Diptera). There are tens of thou-
sands of pollinator species worldwide representing these
different taxonomic groups. Bees alone account for approxi-
mately 20,000 of the pollinating species worldwide
(Michener 2000), with an estimated 4,000 species in the
United States (Winfree et al. 2007a).
Pollinators perform such a range of ecological ser-
vices in natural ecosystems that they are clearly a keystone
group in nearly all terrestrial ecosystems, necessary for
plant reproduction and in forming the basis of an energy-
rich food web (Kearns et al. 1998).
Bees and other pollinator insects are not only important for natural
areas, but also bring great benefits to nearby farms and gardens.
Photograph of mining bee (genus Andrena) by Bruce Newhouse.
Habitat Needs of Pollinator Insects
Using pollinator conservation as a framework for managing
natural areas will result in diverse plant communities and
excellent habitat for all types of wildlife. In managing for
insect pollinators, the first step is to understand the habitat
needs of bees, butterflies, moths, and other species. These
needs can be divided into two main categories: a diversity
of native flowers available throughout the season, and egg-
laying or nesting sites. Land managers can play a key role
in protecting and enhancing pollinator habitat through the
careful application of ecosystem management tools. More-
over, they can provide nesting materials for bees, and where
appropriate, can increase floral resources through revegeta-
tion, as well as provide hostplants for butterflies or moths.
Diversity of native flowers. Flowering plants that
benefit from insect pollination produce energy-rich nectar
and nutrient-rich pollen to attract pollinators. While forag-
ing for these, insects unwittingly transfer pollen grains
within individual flowers, among flowers on the same plant,
and between flowers on plants in the same general area,
fertilizing the flowers and increasing genetic outcrossing of
the species. Forage resources are necessary throughout a
pollinators adult life, and most species benefit from a suc-
cession of diverse blooming plants that provide adequate
forage (Bowers 1985). A diverse community of insect polli-
nators, therefore, requires a diversity of native flowers
(Dramstad & Fry 1995; Holzschuh et al. 2007).

Nesting and egg-laying resources. In addition to
food, insect pollinators require egg-laying sites. For exam-
ple, butterflies and moths require the appropriate hostplants
for laying eggs and for their larvae to eat. The majority of
bee species nest in the ground, digging narrow tunnels that
lead to a small number of brood cells. Most of the remain-
ing bee species occupy existing tunnels in large, dead,
woody vegetation, though some do chew out the center of
pithy twigs (Michener 2000). Bumble bees and honey bees
are the main exception to this. Bumble bees require a small
cavity such as an abandoned rodent nest for their colony
(Kearns & Thomson 2001), and feral honey bees usually
occupy large cavities, such as a hollow tree (OToole &
Raw 1999). Some wood-nesting species also need materials
such as mud, leaf parts, or tree resin to construct brood cells
in their nests (OToole & Raw 1999). It is also important
that nest sites are close enough to sources of nectar and pol-
len (Cane 2001).
Generalist and specialist species. The diverse
habitat requirements suggested above are most appropriate
for conserving generalist pollinator species, those species
that can forage from a wide range of plant sources. In con-
trast, specialist pollinator species use limited sources of
nectar and pollen, or have specific hostplants for their
young. Some studies have found that management tech-
niques that emphasize the broad habitat requirements of
pollinators may preferentially select for generalist species,
while ignoring the more specific requirements of specialist
species (Swengel 1996; Swengel 1998; Winfree et al.
2007a).
3
Livestock grazing in natural areas is a common practice
throughout the United States. While unmanaged grazing can
damage ecosystems (Bilotta & Brazier 2007), there is evi-
dence that carefully managed grazing not only minimizes
negative impacts but can provide positive benefits to floral
resources in some rangeland settings, especially where
shorter flowering plants are suppressed by taller grasses and
buildup of thatch. Like many of the management tools dis-
cussed in this briefing, there is an ecological cost to intro-
ducing grazing into a natural system, especially in areas
where herbivory by large ungulates did not occur. In some
sites, it is entirely unsuited for protecting pollinators, while
in other cases it can be a useful tool. For grazing manage-
ment to be used effectively, the correct combination of tim-
ing, intensity, and duration of grazing, and class of livestock
must be found that is best suited for the site. Finding the
ideal permutation is not easy, and relies on an in-depth
knowledge of the local ecosystem along with well-
developed livestock management skills. Developing a com-
GRAZING
While the ecological impact of overgrazing can be severe, moderate
levels of rotational grazing may provide some positive benefits.
Photograph by Dr. Lloyd Glenn Ingles, California Academy of Sci-
ences.
Each of the techniques considered in this primergrazing,
fire, mowing, herbicides, and insecticidescan be used to
manage habitat to benefit pollinators. Each can also have
damaging, at times severe, impacts on pollinators if they are
not used carefully. Theres no single management plan that
can provide ideal habitat for all pollinator taxa, but there are
some general considerations that apply to all situations.
As with any management activity, biological in-
ventories should be done to first identify important plant
resources and pollinator habitat. Inventories should pay
close attention to occurrences of rare or specialist pollinator
species and their life cycle and habitat requirements. In
some cases, specialist species, especially those with limited
distributions, will become the priority consideration in plan-
ning management to ensure that they receive adequate pro-
tection.
In all of these management techniques, it is impor-
tant to leave some patches untreated. Mowing or burning
HABITAT MANAGEMENT: GENERAL CON-
SIDERATIONS
the entire habitat, for example, could severely impact polli-
nators and leave them with limited opportunities to recolo-
nize the site. It is generally better to treat separate parts of
the site in a multi-year cycle, retaining refugia from which
pollinators can spread.

prehensive grazing and natural resource plan that includes
pollinator conservation practices as one of its goals will
help protect key communities of both plants and pollinators.
Livestock can greatly alter the structure, diversity,
and growth habits of a plant community, which in turn can
affect the associated insect community (Kruess &
Tscharntke 2002). For example, it has been shown that un-
controlled sheep grazing in mountain meadows removed
enough flowering plants to eliminate bumble bees from
study sites (Hatfield & LeBuhn 2007). Sugden (1985) hy-
pothesized that sheep grazing harms pollinator habitat in
four ways: destruction of potential nest sites, destruction of
existing nests and contents, direct trampling of adult bees,
and removal of food resources.
In Arizona, Debano (2006) conducted one of the
few studies focused explicitly on the impacts of domestic
livestock grazing on invertebrate communities in an area
that had not been grazed historically. The results clearly
show that invertebrate species richness, abundance, and
diversity were greater in ungrazed sites. The author sug-
gested that since insects in the Southwest had not evolved in
the presence of buffalo or other large ungulates, they had
not developed grazing pressure adaptations, which made
them more susceptible to being affected by the presence of
cattle.
Other research shows that managed grazing can
benefit insect communities by managing invasive plants and
allowing spring- and summer-blooming flowers to grow.
Controlled grazing has been shown to help maintain an
open, herbaceous-dominated plant community that is capa-
ble of supporting a wide diversity of butterflies and other
pollinators (Smallidge & Leopold 1997).

Management Considerations of Grazing: Timing, Dura-
tion, and Intensity
A diverse pollinator population requires adequate nectar
and pollen sources from early spring to early fall, which
makes seasonal timing a key consideration for an effective
grazing plan. Ideally, management should be adjusted as
needed to maintain the majority of the floral resources in an
area throughout the seasons. Grazing during periods when
floral resources are already scarce (e.g., mid summer,
though this varies between regions) may result in insuffi-
cient forage available for pollinators such as bumble bees
which, in some areas, forage into late September (Carvell
2002). Likewise, grazing during spring when butterfly lar-
vae are active on hostplants can result in larval mortality or
remove important vegetation and nectar resources
(Smallidge & Leopold 1997). The most effective time to
graze varies from site to site, but would generally be after
the majority of the floral resources have died back, or when
many pollinators are in diapause (a state of dormancy) or
4
FIRE
Due to the importance of fire in many native ecosystems,
prescribed burning is a commonly used management tool.
Understanding how insects, including pollinators, respond
to fire is integral to designing a fire management strategy
that protects pollinators.
Many studies have found a negative or mixed re-
sponse of invertebrates to fire. In a study that examined
arthropods in prairies of the American Midwest, Harper et
al. (2000) found that overall species richness and the abun-
dance of all but one of the arthropod species measured de-
creased in burned sites. Their results suggest that burning a
small habitat fragment in its entirety could risk extirpating
some species because of limited recolonization from adja-
cent habitat.
Reviewing the literature associated with the effects
of various management techniques on insects, Swengel
(2001) found that fire is more detrimental than grazing,
mowing, or haying. Due to the loss of more than 99 percent
of the native tall-grass prairie, managing the existing rem-
nants based on historical fire disturbance regimes may not
be appropriate. Instead, Swengel advocates an approach
based on individual site characteristics and assessment of
how the plants and insects at that site will respond to fire.
Many other studies have found that fire benefits
insect communities. Hartley et al. (2007) found that fire was
an excellent management tool in prairies for decreasing
woody plants and simultaneously encouraging higher ar-
thropod diversity. The authors suggested, however, that
recolonization of the burned plots from nearby refugia is an
important factor in the recovery of insect populations.
In researching prescribed burns in western forests,
Huntzinger (2003) found that there was greater butterfly
species richness in areas where fire had been reintroduced.
The burns created large sunlit openings in the forest canopy
that were favorable for herbaceous plant growth, which in
have successfully laid eggs. This is usually in late summer
and fall.
With pollinator conservation as a goal, grazing
intensity and duration are also important to consider. Inten-
sive grazing using high livestock numbers has been shown
to be detrimental. However, moderate to light stocking lev-
els with herds rotated through the management area to cre-
ate a mosaic of grazing stages can be a useful method for
moderating succession in a targeted vegetation community.
The timing and frequency of rotation depends on both the
size and type of the herd and the size of the grazed area
(Schtickzelle et al. 2007). Generally speaking, grazing peri-
ods should be short, with comparatively long recovery peri-
ods for the habitat.

turn encouraged butterflies. The author recommends a cau-
tious approach to prescribed burning to ensure that a range
of habitat heterogeneity is maintained or restored.

Management Considerations of Fire: Scale and Refugia
Prescribed burning as a management tool is a two-edged
sword. It clearly has a role to play in long-term maintenance
of pollinator habitat, but can have catastrophic impacts on
pollinator populations. To avoid undue loss of insects, a
number of considerations should be integrated into fire
management protocols.
A single prescribed fire should not burn an entire
area of pollinator habitat. A program of rotational burning
in which small sections30 percent of a site or lessare
burned every few years will ensure adequate colonization
potential and refugia for insects. In addition, as a fire moves
through an area, skipssmall, unburned patchesshould
be left intact as potential micro-refuges. Periods between
managed burns over the same patch should be conservative.
Based on a variety of studies cited above, it appears that
three to ten years allows adequate recovery of pollinator
populations, depending on the ecosystem and specific man-
agement goals.
Unless the objective for a prescribed fire is for
brush or tree removal, (e.g., pinyon-juniper, chamise, or
mesquite), high-intensity (hot) fires should be avoided.
Low-intensity prescribed burns conducted early or late in
the day, or from late fall to early spring, are not only prefer-
able for pollinators but also reduce impacts on other wild-
life species such as reptiles and ground nesting birds.
5
Fire can be an important management tool for managing or restoring native prairies. Photograph courtesy of J eff Vanuga/USDA-NRCS.
MOWING
Mowing is commonly used in areas where site access and
topography permit equipment access, such as road margins
and powerline corridors, and is often used in place of graz-
ing or fire to manage vegetation. Like grazing, mowing can
alter grassland succession and species composition by sup-
pressing the growth of woody vegetation (Forrester et al.
2005). Mowing can have a significant impact on insects
through direct insect mortality, particularly for egg or larval
stages that cant avoid the mower (Di Giulio et al. 2001).
Mowing also creates a sward of uniform height and may
destroy smaller topographical featuressuch as grass tus-
socks (Morris 2000)when care is not taken to avoid these
features or the mower height is too low. Such features pro-
vide structural diversity to the habitat and offer potential
nesting sites for pollinators such as bumble bees.
Yet, there are some instances when mowing is
beneficial for pollinators. In a large-scale survey of prairie
and barrens butterfly species, Swengel (1998) found that
mowing benefited specialist butterflies.

Management Considerations of Mowing: Technique,
Timing, and Scale
The differences between an ultimately beneficial mowing
regime and a detrimental one are technique, timing, and
scale. Because mowing can completely remove floral re-
sources from the treated area, it should generally not be
conducted when flowers are in bloom. An exception to this
would be in a weed management program where there is a

6
HERBICIDES
INSECTICIDES
Insecticides can severely impact pollinator populations.
Nevertheless, insecticides are widely used in natural areas
throughout the United States to control both native and non-
native species. Many forestry insecticides have been found
to have lethal or sublethal effects on native pollinators
(Helson et al. 1994), and broad-spectrum insecticides used
for grasshopper control in rangelands impact nontarget in-
sects (Alston & Tepedino 2000). On farms, overspray and
drift of insecticides can also affect nontarget organisms in
field borders (ilgi & J epson 1995) or adjacent natural ar-
eas.

Management Considerations of Insecticides: Application
Method, Product Formulation, and Timing
In situations where insecticides must be used, there are a
number of things that can be done to minimize negative
effects on native pollinators (Riedl et al. 2006).
The easiest and simplest way to avoid large polli-
nator die-offs as a result of insecticide application is to
completely avoid treatment of areas that have flowers in
bloom. Because pollinators utilize pollen and nectar as a
food source, often exclusively, they will be active primarily
in and around areas where flowers are abundant (Johansen
& Mayer 1990). By avoiding applications in blooming ar-
eas, the risk to pollinators will be reduced, but not elimi-
nated. Pollinators often travel through or complete some
part of their reproductive process in areas where there are
narrow window of opportunity for optimum control of the
target species. Wherever possible, management areas
should be mowed during seasons when flowers have died
back or are dormant, such as in fall or winter (Munguira &
Thomas 1992). Mowing at these times will also reduce dis-
ruption to nesting bumble bees.
To minimize these effects and allow sufficient
space and time for pollinator populations to recover, mow-
ing a mosaic of patches over several years is better than
mowing an entire site; no single area should be mowed
more than once a year (Di Giulio et al. 2001). If weed man-
agement is the short-term objective, it may be necessary to
mow more frequently. In this case, try to limit mowing only
to patches of weeds. As with all management, carefully
consider the impact of mowing on the life cycle of known
rare or specialist species in the management area. Other
techniques that will benefit pollinators as well as other ter-
restrial wildlife are: use a flushing bar on mower/swather,
use a high minimum mower/swather height (twelve to six-
teen inches), use reduced mower speed (less than eight
miles per hour), and avoid mowing at night (Green 2007).
Wherever herbicides or insecticides are applied, use the most tar-
geted method possible. Photograph by Chris Evans, River to River
CWMA; Bugwood.org.
Herbicides are used to manage vegetation structure and
composition, especially in controlling invasive plants, and
thinning of small trees and shrubs (Miller & Miller 2004).
When applied with care, herbicides can be a useful manage-
ment tool. However, they can also dramatically change
plant communities and decrease the usability of habitat for
pollinators. For example, broadcast applications of non-
selective herbicides can indiscriminately reduce important
floral resources (Smallidge & Leopold 1997), leading to a
decline in pollinator reproductive success and survival rates.

Management Considerations of Herbicides: Application
Method and Active Ingredients
To avoid herbicide damage to nontarget plants and associ-
ated pollinators, some simple precautions should be taken.
In general, avoid broadcast spraying or pellet dispersal be-
cause large numbers of larval hostplants or adult forage
plants could be destroyed. Instead, spot treat with a back-
pack sprayer, weed wipe, or similarly well-targeted tech-
nique, allowing for selective control of undesirable plants
while avoiding nontarget species. In at least one study, tar-
geted spraying, combined with mechanical removal of lar-
ger shrubs, was found to be effective in maintaining butter-
fly habitat. At the same time, long-term management costs
were reduced because fewer visits were required to suppress
undesirable vegetation (Russell et al. 2005).
Herbicide applications should be specific enough
to avoid spraying nontarget forage plants and hostplants.

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insecticides on native bees. In Grasshopper Integrated Pest Management
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and M. W. Sampson. Washington, DC: United States Department of Agri-
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size and organic farming on diversity of birds, pollinators, and plants in a
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Bergman, K. O., J . Askling, O. Ekberg, H. Ignell, H. Wahlman, and P.
Milberg. 2004. Landscape effects on butterfly assemblages in an agricul-
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Bilotta, G. S., and R. Brazier. 2007. The impacts of grazing animals on the
quality of soils, vegetation, and surface waters in intensively managed
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Bowers, M. A. 1985. Bumble bee colonization, extinction, and reproduc-
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2002. Food supply and parental feeding rates of hooded warblers in forest
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7
REFERENCES
To learn more about providing habitat for pollinators,
please see our Pollinator Conservation Handbook. In addi-
tion, check the publications page of our website
(www.xerces.org) for detailed guidelines for wildland man-
agers, as well as guidelines for restoring and creating polli-
nator habitat on farms, parks, and golf courses.
FOR MORE INFORMATION
no blooms. For example, leafcutter bees harvest small leaf
pieces to construct their nests and can be exposed to insecti-
cide residues on contaminated foliage, butterfly caterpillars
will continue to feed on contaminated leaves, and many
bees nest in the ground where they may be oversprayed.
Another key method for reducing insecticide ef-
fects on native pollinators is to choose the insecticide for-
mulation and application method that is the least harmful.
Generally, dusts and microencapsulated insecticides are the
most dangerous formulations for bees, and aerial spraying is
the most harmful method of application (Riedl et al. 2006).
Dusts are difficult to control, can easily be blown out of the
target area, and readily adhere to the small hairs that cover
the bodies of pollinators such as bees. Micro-encapsulated
insecticides are of a similar size and electrostatic charge as
pollen grains, making them easily ingested or transported
back to the nest by foraging insects. Both dusts and micro-
capsules are collected along with pollen and used to provi-
sion brood, which can result in lethal or sublethal effects on
larvae in the nest (Johansen & Mayer 1990; MacKenzie
1993). In comparison, sprayed solutions and large granules
are not as readily incorporated into the foraging of pollina-
tors, and negative effects that do occur are more likely to be
limited to the adult bee. Aerial spraying almost always re-
sults in some degree of unintended drift into nontarget ar-
eas. Methods that increase the accuracy of targeted applica-
tion are recommended, such as ground application and
coarse sprays (Zhong et al. 2004).
Those applying insecticide should be aware of
butterfly hostplants in the management area and avoid
spraying on or around them. If a managed area is known to
host rare or specialist pollinators, ensure that adequately
buffered habitat refugia are available during and after insec-
ticide application.
Pollinators are a vital component of our ecosystems. Planning man-
agement with them in mind will help create a healthy environment.
Photograph of bumble bee (genus Bombus) by Mace Vaughan.

COPYRIGHT 2007 The Xerces Society for Invertebrate Conservation
4828 SE Hawthorne Blvd., Portland, OR 97215. Tel: 503-232 6639. www.xerces.org

The Xerces Society is an equal opportunity employer.
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Our thanks go to the Bradshaw-Knight Foundation, Bullitt Foundation, Columbia Foundation, CS Fund, Disney Wildlife
Conservation Fund, Dudley Foundation, Gaia Fund, Panta Rhea Foundation, and Richard and Rhoda Goldman Fund for
providing funding for this publication. Thank you, too, to Celeste Mazzacano (Xerces Society), Craig Thomsen (UC Davis),
Sean Tenney (Xerces Society), Vance Russell (Audubon California), and Wendell Gilgert (USDA-NRCS) for reviewing
drafts, and to Adair Law for editing.