LETTER

Optimal design of agricultural landscapes

for pollination services
Berry J. Brosi1 , Paul R. Armsworth,2 & Gretchen C. Daily1
1
2

Center for Conservation Biology, Department of Biological Sciences, Stanford University, 385 Serra Mall, Stanford, CA 94107, USA
Department of Animal and Plant Sciences, University of Shefeld, Alfred Denny Building, Western Bank, Shefeld S10 2TN, UK

Keywords
Pollination; agriculture; ecosystem services;
bees; spatial modeling; landscape design.
Correspondence
Berry J. Brosi, Center for Conservation Biology,
Department of Biological Sciences, Stanford
University, 385 Serra Mall, Stanford, CA 94107,
USA. Tel: +1-650-450-3715; fax:
+1-650-723-5920.
E-mail: bbrosi@stanford.edu
Received: 18 October 2007; Revised 26 October
2008; accepted 6 January 2008.
doi: 10.1111/j.1755-263X.2008.00004.x

Abstract
Developing landscape design principles for the provision of ecosystem services is crucial to efficient and widespread implementation of environmental
service-based projects. We investigate optimal farm design for agricultural pollination services from bees nesting in native habitat, integrating ecological and
economic approaches in a spatial modeling framework. We evaluate the simplest case, and then add consideration of bee metapopulation dynamics and
heterogeneity in farmland productivity. We find that the need for spatially
even pollination coverage across farms means that bee habitat is often denser
at the edges, rather than the centers, of optimally designed farms, and also
highly constrains the ability of farmers to site bee habitat in less-productive
areas of farms with spatial gradients in agricultural fertility. Optimal farm configuration is not purely a matter of uniform size and spacing of bee habitat: in
some circumstances, farms combine large parcelsto ensure bee population
persistencewith smaller, dispersed patches to provide spatially continuous
pollination services. The highest-yield farm designs are those with a relatively
small (but non-zero) area of pollination reservoirs, suggesting a conservation
strategy of small parcels of service-providing habitat interspersed throughout
working landscapes. The design principles outlined here are likely general and
applicable to other ecosystem services supplied at local scales, such as agricultural pest control.

Introduction
Nature provides life-sustaining benefits to people in many
forms, including water purification, flood control, carbon sequestration, and insect pollination of crops (Daily
1997). In many places, these ecosystem services are becoming scarce relative to demand (Millennium Ecosystem Assessment 2005), motivating investment in new
mechanisms for their conservation (Chichilnisky & Heal
1998). For example, China is conserving forests for flood
control, New York City is investing in sustainable farming and forestry for water purification, and Costa Rica
pays landowners for biodiversity, scenic beauty, carbon storage, and water purification (Daily & Ellison
2002).

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Conservation Letters 1 (2008) 2736 

To realize the conservation potential of ecosystem services, we must better understand how to align conservation with economic incentives while replicating and
scaling up such model projects. At present, however,
there is little understanding of how to design ecosystem
service projects in a spatially explicit and economically efficient manner (Kareiva & Marvier 2003; Kremen 2005;
Kremen & Ostfeld 2005).
Here we develop a modeling framework for optimizing landscape design for the provision of pollination
services that integrates basic economic trade-offs. Previous works have applied spatial models to the mapping of ecosystem services in extant landscapes, including changes in service delivery under different land-use
change scenarios (e.g., Bodin et al. 2006; Chan et al.

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Landscape design for pollination services

2006; Olschewski et al. 2006). Morandin and Winston

(2006) optimized the quantity of native habitat for pollination services to canola, without considering spatial
configuration. By contrast, the work presented here is,
to our knowledge, the first effort to develop spatially
explicit landscape design principles for any one ecosystem
service.
Insect-mediated pollination services are essential for
harvests of two-thirds of all crop varieties (Roubik 1995)
and for the reproduction of up to 98% of plants in natural systems (Bawa 1990). This vital ecosystem service
is threatened, however, by pollinator declines (Watanabe
1994; Allen-Wardell et al. 1998; Biesmeijer et al. 2006;
NRC 2007). Recent concern over Colony Collapse Disorder in managed honeybees underscores the gravity of this
problem (Armas 2007; Cox-Foster et al. 2007). Nevertheless, wild bees residing in native habitat can provide pollination services to crops (Kremen et al. 2002; Klein et al.
2003; Ricketts 2004). For example, Ricketts et al. (2004)
found a 20% increase in coffee yields near two patches
of native forest in Costa Rica, which generated approximately US$60,000/year for one coffee farm.
These findings point to a strategy of managing what
we call pollination reservoirs, parcels of pollinator habitat
integrated into agricultural production systems in order
to provide crop pollination services. To implement such a
strategy, we need to understand when pollination reservoirs will be an economically attractive option; how big
they need to be to support pollinators; and their optimum
spatial configuration on a farm. Here we develop a spatial
model to configure pollination reservoirs for maximizing
crop yield at the scale of a single farm.

Models
We developed a simple, general modeling framework to
understand essential trade-offs and design principles for a
range of pollinators and farming systems, while allowing
future modeling efforts that add more detail to understand and isolate better the sources of results. Our focus
on generality and simplicity, however, comes at the necessary expense of some realism and immediate applicability.
We considered linear farms divided into cells of either
agriculture or native habitat, where bees nest only in native habitat cells and forage in both native and agricultural cells. We initially assumed constant bee population
density in native habitat cells (e.g., Bodin et al. 2006;
Cane et al. 2006; Brosi et al. in press) and homogeneity
in underlying agricultural fertility; we later relaxed these
assumptions. Ultimately, we optimized the configuration

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B. J. Brosi et al.

of agricultural and pollination reservoir cells within the

farm to maximize crop yields.
We assumed that bees fly in a straight line either left
or right from a reservoir cell R of native habitat with a
fixed probability q f of stopping to forage in each cell encountered; we refer to (1 q f ) as foraging dispersal. The
probability  of a bee from cell R foraging in cell i is:
 = 1/2 q f (1 q f )(|iR|)

(1)

where (|i R|) is the distance traveled. Summations yield

the total number of bees from a given reservoir cell foraging in cell i and from all reservoir cells. This movement
model fits the trends compiled in a global meta-analysis
of studies of bee movement from native habitat in agricultural lands (Ricketts et al. in press).
Bee density does not translate linearly into seed set
because once a sufficient number of pollen grains have
been deposited onto a plants stigma, adding more does
not benefit plant reproduction (e.g., Free 1993; Morandin
and Winston 2006). Following these works, we related
bee density to crop pollination () assuming:
= b + z(1 e dc )

(2)

where d is the density of bees, z controls the maximum

level of insect-mediated pollination, and c controls the
slope of the saturation curve. The baseline level of selfpollination in the absence of bees is b, which is zero for
obligately outcrossing crops such as almonds, and positive for a range of crops including coffee (Free 1993).
We call the proportion of crop yield from pollinators [i.e.,
( b)/] pollinator dependence and the saturating
relationship we modeled allows for assessment of a range
of hypothetical crops or cultivars with different pollinator
dependencies. Default values for all parameters are given
in Appendix S1.
To focus on farm design for pollination services, we assumed that yield was a function solely of crop pollination
(including baseline self-pollination); we later relaxed this
assumption to assess the effects of heterogeneity in underlying agricultural fertility. We optimized farm configuration to maximize crop yield using the MATLAB Genetic Algorithm from the GADS toolbox version 2.0 (The
Mathworks, Inc., Natick, MA, USA); see Appendix S2 for
more information.

Basic model results, trade-offs and intuition

The results from the basic model demonstrate an inherent
trade-off: efficient farm plans must balance the foregone
cost of not farming areas of bee habitat with the benefit of
higher yields from enhanced crop pollination. Pollination
reservoirs occur as single cells within larger areas devoted
to agricultural production (Figure 1) because croplands

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B. J. Brosi et al.

30%

75%

2
5

50%

20%
3

% of farm area
in pollination
reservoirs
(solid line)

% increase in
crop yields over
baseline
(dashed line)
25%

10%

0%

0%
0
0.1
short
dispersal

0.2

0.3

0.4

0.5

0.6

0.7

bee foraging dispersal (1-q f)

0.8

0.9

1
long
dispersal

1
2
3
4
5

Figure 1 Area coverage of pollination reservoirs and crop yield with

bee foraging distance. In the upper graph, each point on the x-axis corresponds to a single optimal farm design; the two lines show the proportion
of native habitat and crop yield for each farm. Designs of ve such farms
are shown in the lower portion, corresponding to the ve numbered points
on the graph. In farm design depictions, gray shading represents a pollina-

tion reservoir and white represents cropland. In the upper graph, the crop
yields for any farm can be found by connecting vertically to the dashed
line; farm 3 represents the point of maximum farm yield. Note that pollination reservoirs are concentrated in the centers of farms only with large
bee foraging distances.

adjacent to native habitat cells receive the greatest pollination benefit, and because bee density within a reservoir
cell is independent of habitat area under the assumption
of constant bee population size.
The quantity and spacing of pollination reservoir cells
in a farm depends on bee foraging (Figure 1). When foraging dispersal (1 q f ) is zero, bees never leave their
home cell, rendering them useless to farmers. At some
minimum bee foraging distance (Figure 1, farm 1), it becomes profitable to forgo agricultural harvests in some
cells and instead manage them as bee habitat. At such
a point, bees forage over short distances and a relatively
large proportion of the farm must be allocated to bee
habitat to provide pollination services to the remaining
croplands.
As foraging dispersal increases, bees are able to pollinate crops over a greater area, and there are fewer, more

widely spaced native habitat cells (Figure 1, farms 24).

Only at large bee foraging distances does the farmer concentrate bee habitat in the center of the farm (Figure 1,
farms 45) to prevent excessive spillover of bees foraging off-farm, and because bees from the center of the
farm forage far enough to pollinate croplands near the
edges. With even greater foraging distances, the farmer
must allocate more land to pollination reservoirs to keep
enough bees foraging on the farm (Figure 1, farm 5).
When bees forage over more extreme distances, most
bees are foraging off-farm and it is no longer profitable
to manage any bee habitat. These situations produce
contrasting farm yields (Figure 1, dashed line). Yield
is maximized when bees forage widely enough to provide pollination services to all of the cropland with the
smallest area of native habitat on the farm (Figure 1,
farm 3).

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Landscape design for pollination services

50%

B. J. Brosi et al.

short-dispersal bees
(dotted line); q f = 0.95
40%
2
4

medium-dispersal bees
(solid line); q f = 0.75

30%

%farm area
as pollination
reservoirs

5
7
3

20%
8

long-dispersal bees
(dashed line); q f = 0.1

10%

0%

20

no selfpollination

40

60

% yield from self-pollination

(inverse pollinator reliance)

80

100
complete selfpollination

short-dispersal bees
1
2
3
medium-dispersal bees
4
5
6
long-dispersal bees
7
8
9
Figure 2 Area coverage of pollination reservoirs with declining pollinator reliance. This graph depicts the interaction between foraging distance
and pollinator dependence. Each of the three responses lines represents a
different, but constant, bee foraging dispersal distance. Every point along
each of the lines comprises a unique farm design. Eight of these are shown

below, corresponding to the numbered points on the graph. As pollinator dependence decreases (moving right along the x-axis), the proportion
of native habitat in farms decreases, with the response dependent on
foraging distance.

As pollinator dependence decreases (as for different

crops), the optimal proportion of bee habitat in a farm
declines (Figure 2), in a manner dependent on foraging dispersal. Bees with short foraging distances require
farms with larger proportions of native habitat, making
the trade-off of forgoing cropland for bee habitat less at-

tractive as pollinator dependence declines (Figure 2, dotted line). In converse, with wide-ranging bees, each cell
of native habitat can provide pollination services to a
larger area of crops, so the total area of pollination reservoirs is initially smaller but its loss is slower as pollinator
dependence decreases (Figure 2, dashed line).

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B. J. Brosi et al.

Population dynamics and sustainability

of pollination services
In the simple case outlined above, we assumed constant
bee population densities in native habitat cells. We relaxed this assumption to evaluate how bee population dynamics affect optimal farm design, which is of particular
importance when considering the persistence of pollinator populations in a landscape and thus the sustainability
of pollination services. We modeled simple metapopulation dynamics using a discrete approximation of logistic
growth of bee colonies, assuming colony dispersal as for
bee foraging but controlled by a separate reproductive
dispersal parameter (q r ). We iterated growth and dispersal dynamics for 500 generations, which consistently
yielded equilibrium population numbers. At each time
step, colonies first reproduced and then the newly produced propagules dispersed; propagules that stopped to
nest in pollination reservoirs survived, while those that
settled in agricultural cells did not. We used a seeded version of the genetic algorithm for optimization routines
with population dynamics (see Appendix S2 for more
detail); optimization routines were run after population
equilibrium was reached. We assessed 10,000 combinations of reproductive (q r ) by foraging (q f ) dispersal parameters (Figure 3).
When reproductive dispersal is zero, new colonies do
not leave their natal landscape cell, so each cell reaches its
carrying capacitybehaving exactly as for the simple case
without population dynamics (Figure 3, bottom edge of
graph). For long-distance bee reproductive dispersal, all
newly formed colonies disperse out of the farm boundaries, making pollination reservoirs worthless to farmers
(Figure 3, top edge). Similarly, when bees forage over
extremely small or large distances, farmers have no economic incentive to manage for pollination reservoirs, as
for the simple case (Figure 3, left and right edges).
Both the size and isolation of native habitat patches affect bee populations. For a pollination reservoir to sustain a bee population, it must be situated close enough
to other reservoirs to recruit new colonies from them
(Figure 3, farms 1 and 2) or else of sufficient size to retain enough of its own newly produced colonies (e.g.,
Figure 3, farms 37). Thus, in many cases farmers will
maintain larger pollination reservoirs, in contrast to the
simple case where all optimal farms have single-cell
reservoirs.
Farm design with bee population dynamics is largely
determined by interactions between bee dispersal for foraging and reproduction (Figure 3), and a wide range of
these parameters occurs in natural bee populations. For
bees with low foraging dispersal, farmers must closely
space pollination reservoirs to ensure effective pollina-

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Landscape design for pollination services

tion across their farm. This enforced reservoir proximity

reduces the minimum reservoir size needed to sustain
bee population, leading to small, closely spaced reservoirs (e.g., Figure 3, farm 1). In contrast, as reproductive dispersal increases (y-axis in Figure 3), patch size
requirements become larger (Figure 3 and Appendix S3
Figure A, farms 5, 7, 9) and thus must be balanced by increased foraging dispersal. Intermediate levels of foraging
and reproductive dispersal generate intermediate patch
size and spacing (Figure 3, farms 34).
Under some parameter combinations, the conflicting
demands of reservoir size and spacing lead to a hybrid
farm plan design, consisting of one or more relatively
large reservoirs to ensure a sustainable population of bees
and smaller satellite parcels sited between them to provide spatially even crop pollination (Figure 3, farms 46).
The conditions with the highest crop yields (Appendix S3,
Figure B) occur with short bee reproductive dispersal and
large foraging dispersal, leading to small, widely spaced
reservoirs (Figure 3 and Appendix S3 Figure B, farm 10).
A wide range of these foraging and reproductive dispersal parameters is reasonable for natural bee populations.
For many organisms (not just pollinators), reproductive
dispersal occurs over longer distances than foraging dispersal (i.e., Figure 3, above and left of the x = y diagonal).
The majority of bee species, including Apis mellifera (Apidae), the European honey bee (Gould & Gould 1988)
likely fit this pattern. Other bee species, however, have
different reproductive and foraging movement patterns.
The meliponines, social stingless bees of the tropics (Apidae: Meliponini), are instrumental in pollinating coffee
in both Asia and Latin America (Klein et al. 2003; Ricketts
2004). In meliponine colony reproduction, workers make
multiple trips between the old and new nest sites, meaning that new colonies cannot disperse much further than
normal foraging distances (Roubik 1988; Figure 3, areas near the x = y diagonal). Some ground-nesting solitary bees nest mere centimeters away from their natal
nest, as part of a nesting aggregation. Such bees, including some species of Perdita (Andrenidae; e.g., Danforth
1999) and the Alkali Bee (Nomia melanderi, with nesting beds managed for alfalfa pollination, Bohart 1972),
forage over larger distances than they typically disperse
reproductively (Figure 3, below and right of the x = y
diagonal).

Effect of cropland heterogeneity

In the simple case and with population dynamics, we
assumed equivalence of farm cells in potential agricultural yield (analogous to soil fertility). While such an assumption holds in industrial, large-scale agriculture, heterogeneity in fertility is present in many smaller-scale

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B. J. Brosi et al.

Proportion pollination reservoirs with population dynamics

varying reproductive & foraging dispersal, 50-cell landscape
1

50%

0.9

45%
9

40%

35%

0.7
7
6

0.6

30%

3
4

0.5

25%
20%

0.4
2

15%

0.3
1
8

0.2

10%
10

5%

0.1
0

proportion of pollination reservoirs

reproductive dispersal (1-q_repro)

0.8

0%
0

0.1

0.2

0.3

0.4
0.5
0.6
0.7
foraging dispersal (1-q_forag)

0.8

0.9

1
2
3
4
5
6
7
8
9
10
Figure 3 Area coverage of pollination reserves with population dynamics. When population dynamics are considered, farm design is driven
largely by the interaction between foraging distance (x-axis) and reproductive dispersal (y-axis). Each point on the graph corresponds to the farm
design that is optimal for that parameter combination, with color at that
point related to the area of pollination reserves on the farm. Lighter colors
represent farms with greater coverage of pollination reservoirs. Numbers

32

on the graph correspond to numbered farm designs below and are ordered by foraging dispersal (x-axis). Foraging distance and reproductive
dispersal distance interact to drive farm designs with different patch size
and spacing. In some cases, for example, farms 46, a hybrid design with
two different patch sizes results. For representation of other variables
from the same farm designs, see Appendix S3 for maximum patch size
(Figure A) and crop yields (Figure B).

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B. J. Brosi et al.

operations, particularly in the developing world. To address this issue, we relaxed the assumption of homogeneity in fertility by considering farms with a linear gradient
in cropland fertility, capturing a simple type of variation
found on hill slopes and along distance gradients to rivers.
We established a fertility gradient by multiplying the
self-pollination of each agricultural cell (variable b, Equation (2)) by a linear fertility factor. We set up paired
farms of equal average fertility in farms with and without fertility gradients by adjusting the baseline level of
self-pollination, allowing us to focus on the differences in
spatial configuration. For crop yields with fertility gradients, see Appendix S5.
Fertility gradients create an incentive for farmers to
shift bee habitat toward the less-fertile side of the
farm (Figure 4). Most such shifts are relatively modest
(Figure 4, farms 36), though they vary with the strength
of the fertility gradient and with bee foraging dispersal.
The subtlety of farm plan response to fertility gradients is
driven by the fact that the most fertile agricultural cells
are the most costly to trade off, but they also have the
most to gain from nearby bee habitat.
Surprisingly, the greatest changes in farm plans occur
with short bee foraging dispersal (Figure 4, dotted line) in
crops with low to intermediate pollinator reliance. With a
relatively strong fertility gradient, farmers do not trade off
any cropland for bee habitat on the higher-fertility side of
the farm (Figure 4, farms 12), but do on the less fertile
end of the farm where there is a lower cost to maintaining bee habitat. For crops with high pollinator reliance,
this pattern reverses because pollination services are necessary across the breadth of the farm.

Discussion
Model insights
Our model demonstrates three novel and nonintuitive insights into the design of agricultural landscapes for maximizing pollination services: (1) Rather than concentrating
pollination reservoir cells in the center of farms to minimize bee loss, most optimal farm designs have reservoirs
spread across their breadth. Counter-intuitively, some
farm plans have a greater density of reservoir cells at their
edges as compared to the center; (2) Optimal farm configuration is not a simple matter of uniform size and spacing
of pollination reservoirs. Instead, many optimal designs
include large patches to ensure bee population persistence, with smaller, dispersed patches to provide spatially
even pollination services; and (3) The need for spatially
continuous pollination services constrains the ability of
farmers to site pollination reservoir cells in less-fertile
areas of heterogeneous farms. Surprisingly, the largest

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Landscape design for pollination services

shifts in reservoir placement occur when bees forage over

small scales.
To achieve satisfactory crop yields, pollination must be
provided across the breadth of a farm. Croplands on farm
edges often need a greater density of nearby pollination
reservoirs than those in farm centers (e.g., Figure 1, farms
13, Figure 2, farms 16) due to the lack of bee input
from adjacent land and the leakage of bees foraging
off-farm, which is greater at farm edges. This is surprising because we had anticipated that farm designs with
bee habitat concentrated in their centers would minimize
bee lossa configuration that only occurs at large foraging distances (e.g., Figure 1, farm 5). For most parameter
combinations, the increased concentration of pollination
reservoirs is near, rather than directly on, the edges of
farms, and the differences in concentration are relatively
subtle.
Small pollination reservoirs spaced evenly across a farm
are not necessarily sufficient to ensure bee population
persistence. Our initial intuition was that inclusion of
population dynamics would uniformly increase reservoir
patch sizes. Instead, it sometimes leads to a hybrid design with one or more large reservoir patches (sources)
mixed with smaller, dispersed sink patches that allow
for spatially continuous pollination services. Thus, farm
design is not necessarily a simple matter of a uniform size
and spacing of bee habitat.
With heterogeneity in agricultural fertility, farmers
could ideally site pollination reservoirs in less fertile areas, freeing more fertile zones for crop production. With
linear gradients in fertility, however, pollination reservoirs cannot be too far from the most valuable croplands or they would suffer reduced pollination services
and smaller harvests. Therefore, shifts in the placement of
pollination reservoir cells toward the less fertile end of the
farm are typically small, constrained by the need for spatially continuous pollination services. Farm designs can
change strongly with fertility gradients, but surprisingly,
this is not with long-range bee foraging dispersal. Instead,
with low pollinator dependence and short-foraging bees,
farm plans place all pollination reservoirs on the less fertile end of the farm and manage the more fertile side
purely for agricultural production.
A more general result is that the conditions leading
to the greatest crop yields have relatively small amounts
of native habitat (e.g., Figure 1, farm 3; Figure 3 and
Appendix S3 Figure B, farm 10). This result may underpin management recommendations where some
land is removed from production to provide ecosystem
services. In many such cases, investing in environmental
services will be most profitable and commonplace
when landowners can set aside the smallest area
needed to realize ecosystem service benefits, suggesting

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B. J. Brosi et al.

reservoirs 25%
strongly
shifted

20%

% shift of
pollination
reservoirs
from farm
center

short-dispersal
bees; q f = 0.9

long-dispersal
bees; q f = 0.15

15%

10%

5%

reservoirs
completely
centered

0%

medium-dispersal
bees; q f = 0.6

all cells
equivalent

slope of fertility gradient

7
strong fertility
gradient

short-dispersal bees
1
2
long-dispersal bees
3
4
medium-dispersal bees
5
6
least fertile

intermediate fertility

most fertile

Figure 4 Shifts in the center of mass of pollination reservoirs with fertility gradients. This graph shows the extent to which native habitat on a farm
is shifted in response to gradients in underlying agricultural fertility. In the
farm design depictions below, the left side is less fertile and the right side
is more fertile. On the y-axis, the percentage shift from farm center is measured as center of mass (average position of reservoir cells), re-centered
at zero, and divided by farm length. Thus, in the 50-cell landscapes used
here, a 10% shift means that reservoir cell center of mass is 5 cells away

from the farms true center. The larger the value, the more shifted or
asymmetrical the farm plan is. Numbered circles correspond to sample
farm designs below. The largest shifts (e.g., farm 2) occur when bees have
relatively short foraging distances and when the farm has a strong fertility
gradient; in such cases, optimal farm designs do not include pollination
reservoirs on the more-fertile side of the farm. See Figure S2 for a graph
of crop yields from the same farm designs.

that environmental service-based projects may favor

widespread implementation of small-scale measures
interwoven throughout working lands.
The results of our model link to previous works in a
number of ways. In some respects, the crop pollination

system is most analogous to marine reserve models, some

of which have attempted to optimize the size and configuration of no-take reserves interspersed with fishable
areas (e.g., Neubert 2003). Such models, however, typically attempt to maximize the number of fish produced

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B. J. Brosi et al.

across a seascape, without regard to where they are produced. By contrast, in a crop pollination system the goal is
not maximizing bee populations; it is producing enough
pollinators to provide sufficient crop pollination (subject to the diminishing-returns relationship captured in
our Equation (2)). In addition, an optimal landscape is
also subject to the inherently spatial constraint of ensuring that the pollinators are spread across the breadth of
the farm so that they can reach as much of the crop as
possible.
Compared with other works on pollination, our findings are consistent with those of Bodin et al. (2006),
who showed that small forest reserves scattered across a
landscape could provide substantial pollination services.
They are also in accord with findings from Morandin and
Winston (2006) who found that landscapes with about
30% native habitat generated optimal farmer profit in a
canola agroecosystem. Our default parameters generated
similar estimates of about 25% native habitat, though for
a crop with about 40% of yield due to self-pollination
(which is likely higher than the percentage for canola
[Free 1993]).

Model extensions
Extensions of our model would contribute additional insights to the design principles presented here. One possibility would consider multispecies bee communities with
stochastic extinctioncolonization dynamics, linking pollination reservoir size with population persistence and
bee species richness. The availability of season-long foraging resources may be a limiting factor for bees in some
landscapes, and could change optimal landscape configurations. Considering several adjacent farms rather than
an isolated one with no bee input from surrounding areas could also change the design of optimal farms, especially at their edges. It would also be valuable to parameterize the models for a specific crop system, considering
economic variables like crop prices, farm inputs, and the
availability and price of managed honeybee pollination
services.

Achieving a pollination reservoir strategy

Implementing a pollination reservoir strategy would confer a range of additional benefits to society. Even small
tracts of native habitat in agricultural landscapes can provide habitat requirements for a diverse range of taxa
(Estrada et al. 1997; Daily et al. 2001; Perfecto & Vandermeer 2002; Brosi et al. 2007), and can contribute services
such as crop pest control (Thies & Tscharntke 1999) and
water quality improvement (Balvanera et al. 2001). Creating funding mechanisms for pollination reservoir strate-

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Landscape design for pollination services

gies could help capture these additional benefits, especially if they target farmlands with a high degree of congruence between pollination services and other conservation values (Balvanera et al. 2001).
Given the increasing investment in ecosystem servicebased conservation projects, it is imperative to develop
spatially explicit strategies for provisioning such services
in an economically efficient manner. The models presented here lay a conceptual foundation for strategies to
provide agricultural pollination services, which could easily be extended to other services that native habitats provision over small spatial scales, like agricultural pest control. We hope that this framework will assist in moving
ecosystem services toward becoming a mainstream, commonplace strategy for improving both biological conservation and human welfare.

Acknowledgments
We thank M. Boni, M. MacPherson, S. Ramachandran,
and J. Van Cleve for programming assistance and advice.
W. Murray gave helpful suggestions and advice on optimization methods. K. Al-Kafaji, M. Boni, S. Jackson, and
J. Van Cleve gave insightful and constructive comments
on the article. B. Brosi was supported during this work
by the Anne and Robert Bass Stanford Graduate Fellowship in Science and Engineering and by a Teresa Heinz
Scholarship for Environmental Research. We gratefully
acknowledge further support from the Winslow Foundation and Peter and Helen Bing.

Supplementary Material
The following supplementary material is available for this
article:
Appendix S1: Default parameter values and equations.
Appendix S2: Optimization methods.
Appendix S3: Maximum patch size and crop yields
with population dynamics, with figure.
Appendix S4: Crop yields with fertility gradients, for
three bee foraging dispersal patterns, with figure.
This material is available as part of the online article
from:
http://www.blackwell-synergy.com/doi/full/10.1111/j.
1755-263X.2008.00004.x
(This link will take you to the article abstract).
Please note: Blackwell Publishing are not responsible
for the content or functionality of any supplementary
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than missing material) should be directed to the corresponding author for the article.

35

Landscape design for pollination services

References
Allen-Wardell G., Bernhardt P., Bitner R. et al. (1998) The
potential consequences of pollinator declines on the
conservation of biodiversity and stability of food crop
yields. Conserv Biol 12, 817.
Armas, G.C. (2007) Mystery ailment strikes honeybees.
Washington Post February 11, 2007.
Balvanera, P., Daily G.C., Ehrlich P.R. et al. (2001) Conserving
biodiversity and ecosystem services. Science 291, 2047.
Bawa, K. (1990) Plant-pollinator interactions in tropical
rain-forests. Annu Rev Ecol System 21, 399422.
Biesmeijer, J.C., Roberts S.P.M., Reemer M. et al. (2006)
Parallel declines in pollinators and insect-pollinated plants
in Britain and the Netherlands. Science 313, 351354.
Bodin, O., Tengo M., Norman A., Lundberg J., Elmqvist T.
(2006) The value of small size: loss of forest patches and
ecological thresholds in southern Madagascar. Ecol Appl 16,
440451.
Bohart, G.E. (1972) Management of wild bees for the
pollination of crops. Annu Rev Entomol 17, 287312.
Brosi, B.J., Daily G.C., Ehrlich P.R. (2007) Bee community
shifts with landscape context in a tropical countryside. Ecol
Appl 17, 418430.
G. (In
Brosi, B.J., Daily G.C., Shih T.M., Oviedo F., Duran
press) The effects of forest fragmentation on bee
communities in tropical countryside. J Appl Ecol doi:
10.1111/j.13652664.2007.01412.x.
Cane, J.H., Minckley R.L., Kervin L.J., Roulston T.H.,
Williams N.M. (2006) Complex responses within a desert
bee guild (Hymenoptera: Apiformes) to urban habitat
fragmentation. Ecol Appl 16, 632644.
Chan, K., Shaw M., Cameron D., Underwood E., Daily G.
(2006) Conservation planning for ecosystem services. PLoS
Biol 4, 21382152.
Chichilnisky, G., Heal G. (1998) Economic returns from the
biosphere. Nature 391, 629630.
Cox-Foster, D.L., Conlan S., Holmes E.C. et al. (2007) A
metagenomic survey of microbes in honey bee colony
collapse disorder. Science 318, 283287.
Daily, G., Ehrlich P., Sanchez-Azofeifa G. (2001) Countryside
biogeography: use of human-dominated habitats by the
avifauna of southern Costa Rica. Ecol Appl 11, 113.
Daily, G.C. (1997) Natures services: societal dependence on natural
ecosystems. Island Press, Washington, D.C.
Daily, G.C., Ellison K. (2002) The new economy of nature: the
quest to make conservation profitable. Island Press: Shearwater
Books, Washington, D.C.
Danforth, B.N. (1999). Emergence dynamics and bet hedging
in a desert bee, Perdita portalis. Proc Royal Soc Biol Sci, Series B
266, 19851994.
Estrada, A., Coates-Estrada R., Meritt D. (1997)
Anthropogenic landscape changes and avian diversity at
Los Tuxtlas, Mexico. Biodivers Conserv 6, 1943.
Free, J.B. (1993) Insect pollination of crops. Academic Press,
London.

36

B. J. Brosi et al.

Gould, J. L., Gould C. G. (1988) The honey bee. Scientific

American Library. W. H. Freeman, New York.
Kareiva, P., Marvier M. (2003) Conserving biodiversity
coldspotsrecent calls to direct conservation funding to
the worlds biodiversity hotspots may be bad investment
advice. Am Sci 91, 344351.
Klein, A., Steffan-Dewenter I., Tscharntke T. (2003) Fruit
set of highland coffee increases with the diversity of
pollinating bees. Proc Royal Soc London Series B-Biol Sci 270,
955961.
Kremen, C. (2005) Managing ecosystem services: what do we
need to know about their ecology? Ecol Lett 8, 468479.
Kremen, C., Ostfeld R.S. (2005) A call to ecologists:
measuring, analyzing, and managing ecosystem services.
Front Ecol Environ 3, 540548.
Kremen, C., Williams N.M., Thorp R.W. (2002) Crop
pollination from native bees at risk from agricultural
intensification. Proc Natl Acad Sci USA 99, 1681216816.
Millennium Ecosystem Assessment (2005) Ecosystems and
human well-being. Island Press, Washington, D.C.
Morandin, L., Winston M. (2006) Pollinators provide
economic incentive to preserve natural land in
agroecosystems. Agric Ecosyst Environ 116, 289292.
National Research Council (2007) Status of pollinators in North
America. National Academies Press, Washington, D.C.
Neubert, M. G. (2003) Marine reserves and optimal
harvesting. Ecology Letters 6, 843849.
Olschewski, R., Tscharntke T., Bentez P.C., Schwarze S.,
Klein A.-M. (2006) Economic evaluation of pollination
services comparing coffee landscapes in Ecuador and
Indonesia. Ecol Soc 11(1), 7.
Perfecto, I., Vandermeer J. (2002) Quality of agroecological
matrix in a tropical montane landscape: ants in coffee
plantations in southern Mexico. Conserv Biol 16, 174182.
Ricketts, T.H. (2004). Tropical forest fragments enhance
pollinator activity in nearby coffee crops. Conserv Biol 18,
12621271.
Ricketts, T.H., Daily G.C., Ehrlich P.R., Michener C.D. (2004).
Economic value of tropical forest to coffee production. Proc
Natl Acad Sci USA 101, 1257912582.
Ricketts, T.H., Regetz J., Steffan-Dewenter I. et al. (In press)
Landscape effects on crop pollination services: are there
general patterns? Ecol Lett. doi: 10.1111/j.1461-0248.2008.
01157.x
Roubik, D.W. (1995) Pollination of cultivated plants in the tropics.
Food and Agriculture Organization of the United Nations,
Rome.
Roubik, D. W. (1988) Ecology and natural history of tropical bees.
Cambridge University Press, Cambridge, UK.
Thies, C., Tscharntke T. (1999) Landscape structure and
biological control in agroecosystems. Science 285, 893895.
Watanabe, M.E. (1994). Pollination worries rise as
honey-bees decline. Science 265, 11701170.
Editor: Dr. Corey Bradshaw

c 2008 Blackwell Publishing, Inc.

Conservation Letters 1 (2008) 2736