This dissertation explores interactions between agrobiodiversity and livelihoods among small-scale coffee farmers in El Salvador. It finds that farmers maintain crop diversity through cultivation of maize landraces for food security against climate variability and price fluctuations. While agrobiodiversity provided cultural value, there was no strong relationship between diversity and food security; income was a better predictor. Shade tree diversity in coffee farms was stable over time despite some tree removal. Higher initial shade tree species richness correlated with greater carbon storage in those trees.

1. AGROBIODIVERSITY, CONSERVATION, AND FOOD SECURITY AMONG
SMALL-SCALE COFFEE FARMERS IN EL SALVADOR
A Dissertation Presented
by
Meryl Breton Richards
to
The Faculty of the Graduate College
of
The University of Vermont
In Partial Fulfillment of the Requirements
for the Degree of Doctor of Philosophy
Specializing in Plant and Soil Science
May, 2013

2. Accepted by the Faculty of the Graduate College, The University of Vermont, in
partial fulfillment of the requirements for the degree of Doctor of Philosophy
specializing in Plant and Soil Science.
Dissertation Examination Committee:
____________________________________ Advisor
V. Ernesto Méndez, Ph.D.
____________________________________
Donald Ross, Ph.D.
____________________________________
Sarah Taylor Lovell, Ph.D.
____________________________________ Chairperson
Marta Ceroni, Ph.D.
____________________________________ Dean, Graduate College
Domenico Grasso, Ph.D.
Date: March 5, 2013

3. ABSTRACT
Smallholder shade coffee agroecosystems have been noted for their potential for
tree, bird, and insect biodiversity conservation in the tropics. However, there is a lack of
research on other productive areas managed by small-scale coffee farmers, such as
homegardens and maize and bean plots. Additionally, little research has investigated why
farmers maintain this agrobiodiversity, how it contributes to their livelihoods, and how
stable it is over time.
In this dissertation, I explore interactions between farmers’ livelihoods and
diversity of cultivated plant species and landraces among smallholder shade coffee
farmers in El Salvador. The research was conducted using a mix of natural and social
science methods, and was driven by the following objectives:
1. understanding the factors and livelihood contexts that motivate farmers to
conserve agrobiodiversity;
2. analyzing how agrobiodiversity contributes to livelihood outcomes,
specifically food security;
3. exploring co-benefits between agrobiodiversity conservation and carbon
sequestration; and
4. examining how agrobiodiversity changes over time as a result of farmers’
management choices.
Farmers maintained agrobiodiversity on their farms as insurance against variable
growing conditions (e.g. climate) and as a buffer against fluctuating market prices for
food. Agrobiodiversity, and maize landraces in particular, held considerable cultural
value as well. There was no strong quantitative relationship between agrobiodiversity and
food security; income was a stronger predictor of food security. Shade tree biodiversity in
coffee plantations was stable over time, despite farmers’ removal of some trees. Shade
tree biodiversity may also have other ecosystem service benefits, as carbon sequestration
in shade trees was correlated with initial species richness.
The value of agrobiodiversity to livelihoods and ecosystem services has
historically been difficult to measure. This dissertation shows positive relationships in
several domains and suggests methodological improvements for measurement in others.

4. ii
CITATIONS
Material from this dissertation has been published in the following form:
Olson, M.B., Morris, K.S., Méndez, V.E., 2012. Cultivation of maize landraces by small-
scale shade coffee farmers in western El Salvador. Agricultural Systems 111, 63-
74.
Material from this dissertation has been submitted for publication to Conservation
Biology on February 11, 2013 in the following form:
Richards, M.B., Méndez, V.E., 2013. Changes in shade tree biodiversity and above-
ground carbon stocks in a smallholder coffee cooperative of El Salvador.
Conservation Biology.

5. iii
ACKNOWLEDGEMENTS
I owe the largest debt of gratitude to my advisor, Dr. Ernesto Méndez, whom I
admire greatly for his scholarship and dedication to research that furthers social justice.
He has been not only an advisor and teacher, but also a friend. I also fondly thank all the
members of the Agroecology and Rural Livelihoods Lab at the University of Vermont,
who have provided valuable support, collaboration, camaraderie, and entertainment.
I would also like to thank the members and families of ACOES, as well as the
farmers of the Las Casitas group of APICAFE for their collaboration in this research.
I received funding for this research from the University of Vermont’s College of
Agriculture and Life Science and the Annie’s Homegrown Sustainable Agriculture
Scholarship.
Finally, my parents, Constance and Douglas Richards, and my twin brother, Matt,
inspired my love of education, fostered my belief in myself, and continue to be my
biggest fans.

6. iv
TABLE OF CONTENTS
CITATIONS ....................................................................................................................... ii
LIST OF TABLES............................................................................................................ vii
LIST OF FIGURES .........................................................................................................viii
CHAPTER 1: REVIEW OF LITERATURE...................................................... 1
1.1. Introduction.................................................................................................. 1
1.2. A framework for examining biodiversity-livelihoods interactions among
coffee farming households ........................................................................... 4
1.2.1. The livelihoods concept .................................................................... 4
1.2.2. Scale in biodiversity-livelihoods analysis......................................... 6
1.2.3. Academic fields to examine relationships between biodiversity and
livelihoods in agroecosystems................................................................ 8
1.3. Causes of biodiversity loss in agroecosystems.......................................... 12
1.4. What is the importance of biodiversity in agroecosystems?...................... 15
1.5. How does biodiversity support livelihoods?.............................................. 18
1.6. Efforts to provide incentives for agrobiodiversity conservation................ 20
1.7. References.................................................................................................. 26
CHAPTER 2: CULTIVATION OF MAIZE LANDRACES BY SMALL-SCALE
SHADE COFFEE FARMERS IN WESTERN EL SALVADOR.................................... 41
2.1. Abstract...................................................................................................... 41
2.2. Introduction................................................................................................ 42
2.2.1. Maize Seed and Landraces in El Salvador...................................... 45
2.3. Methods...................................................................................................... 47
2.3.1. Study site......................................................................................... 47
2.3.2. Study plots....................................................................................... 49
2.3.3. Data collection ................................................................................ 51
2.3.4. Data analysis ................................................................................... 53
2.4. Results........................................................................................................ 55
2.4.1. Maize seed types ............................................................................. 55
2.4.2. Predictors of maize seed choice...................................................... 56
2.4.3. Interactions between management and seed type ........................... 62
2.4.4. Maize yields .................................................................................... 66

7. v
2.5. Discussion.................................................................................................. 69
2.6. Conclusion ................................................................................................. 75
2.7. References.................................................................................................. 76
CHAPTER 3: CHANGES IN SHADE TREE BIODIVERSITY AND ABOVE-
GROUND CARBON STOCKS IN A SMALLHOLDER COFFEE COOPERATIVE OF
EL SALVADOR............................................................................................................... 83
3.1. Abstract...................................................................................................... 83
3.2. Introduction................................................................................................ 83
3.3. Methods...................................................................................................... 87
3.3.1. Study site......................................................................................... 87
3.3.2. Data collection ................................................................................ 90
3.3.3. Data analysis ................................................................................... 90
3.4. Results........................................................................................................ 92
3.4.1. Vegetation structure ........................................................................ 92
3.4.2. Shade tree diversity......................................................................... 93
3.4.3. Shade tree C stocks ......................................................................... 97
3.4.4. Shade tree biodiversity and carbon sequestration........................... 99
3.5. Discussion................................................................................................ 101
3.5.1. Conservation of shade tree biodiversity........................................ 101
3.5.2. Carbon sequestration in coffee agroforestry systems ................... 103
3.5.3. Shade tree biodiversity and carbon sequestration......................... 104
3.6. References................................................................................................ 106
CHAPTER 4: IS THERE A QUANTITATIVE LINK BETWEEN SPECIES
DIVERSITY AND FOOD SECURITY IN SMALLHOLDER SHADE COFFEE
AGROECOSYSTEMS? ................................................................................................. 111
4.1. Introduction.............................................................................................. 111
4.2. Methods.................................................................................................... 114
4.2.1. Study sites ..................................................................................... 114
4.2.2. Data collection .............................................................................. 115
4.2.3. Data analysis ................................................................................. 117
4.3. Results...................................................................................................... 120
4.3.1. Community characteristics............................................................ 120

8. vi
4.3.2. Biodiversity of farm components.................................................. 121
4.3.3. Food security indices .................................................................... 124
4.4. Discussion................................................................................................ 128
4.5.References ................................................................................................ 131
COMPREHENSIVE BIBLIOGRAPHY ........................................................................ 135
APPENDICES ................................................................................................................ 150

9. vii
LIST OF TABLES
Table Page
Table 2.1. Variables included in the study........................................................................ 51
Table 2.2. Maize types in milpa plots (n = 42 plots). Some plots were planted with more
than one type of seed................................................................................................. 57
Table 2.3. Descriptive statistics and significance of continuous explanatory variables used
in the comparison of plots planted with criollo seed and plots planted with
certificado seed. Variables for which there were significant differences are in bold.
................................................................................................................................... 59
Table 2.4. Frequencies and significance of categorical variables in the comparison of
plots planted with criollo seed and plots planted with certificado seed. None were
significantly (p ≤ 0.05) associated with seed type. ................................................... 60
Table 2.5. Logistic regression model for prediction of use of certificado seed................ 62
Table 3.1. Properties and t tests (where applicable) of the shade tree canopy of a 35 ha
coffee cooperative in Tacuba, El Salvador in 2001 and 2010. ................................. 93
Table 4.1. Example species by trait matrix of nutrient values........................................ 118

10. viii
LIST OF FIGURES
Figure Page
Figure 2.1. Study site ........................................................................................................ 48
Figure 2.2. Land use in study site ..................................................................................... 49
Figure 3.1. Study site ........................................................................................................ 89
Figure 3.2. Species rarefaction curves (Mao-Tau) for 2001 and 2010, with 95%
confidence intervals, found in 14 quadrats in Tacuba, El Salvador.......................... 95
Figure 3.3. Shade tree rank-abundance curves, 2001 and 2010, found in 14 quadrats in
Tacuba, El Salvador.................................................................................................. 96
Figure 3.4. Similarity of shade tree species assemblages in 2001 and 2010 in 14 quadrats
in Tacuba, El Salvador.............................................................................................. 97
Figure 3.5. Carbon stocks stored in shade trees over 9 years in a coffee cooperative of
Tacuba, El Salvador. In general, carbon stocks increased due increases in average
tree size, despite small reductions in shade tree density. In quadrats 4, 5, and 7,
which are on the southwestern edge of the cooperative, increases in tree size were
not large enough to compensate for reductions in tree density, and carbon stocks
decreased................................................................................................................... 99
Figure 3.6. Correlation between carbon sequestration rate and initial (2001) species per
stem in 14 quadrats in Tacuba, El Salvador............................................................ 100
Figure 3.7. Correlations between carbon stocks and species per stem in each sample year
in 14 quadrats in Tacuba, El Salvador. There were no significant correlations
between carbon stocks and species density. ........................................................... 101
Figure 4.1. Nutritional functional nutritional diversity of combined homegarden and field
crop species was positively correlated with species richness................................. 123
Figure 4.2. Nutritional functional diversity was positively correlated with species richness
in coffee plantations................................................................................................ 124
Figure 4.3. Most commonly consumed food groups in 24-hour recall........................... 125
Figure 4.4. There were no significant correlations between dietary diversity and
nutritional FD measures.......................................................................................... 126

11. ix
Figure 4.5. Dietary diversity was weakly correlated with monthly household food
spending (a) and income per person (b).................................................................. 127

12. 1
CHAPTER 1: REVIEW OF LITERATURE
1.1. Introduction
Approximately 40% of the earth’s land surface is currently in agricultural use
(Chappell and LaValle, 2009; Foley et al., 2011), and it is estimated that agricultural
production will need to double in order to meet the food and energy needs of a projected
population of 9 million by 2050 (Godfray et al., 2010; Foley et al., 2011). This will place
substantial pressure on the world’s land resources, particularly in the tropics, where the
most significant expansions in cropland are occurring (Tilman et al., 2001a; Foley et al.,
2011), along with intensification of existing agricultural lands (Matson and Vitousek,
2006).
One of the most severe consequences of agricultural expansion and intensification
is loss of biodiversity. While global loss of biodiversity is difficult to measure, it is clear
that all aspects of biodiversity are currently at risk. A 2010 review of multiple biodiversity
measures found that most show biodiversity continuing to decline with no reduction in the
rate of decline, despite the commitments made at the 2002 Convention on Biological
Diversity (Butchart et al., 2010). Various estimates put the current rate of extinction at
possibly thousands of times higher than for most of evolutionary history (Chappell and
LaValle, 2009). While not all of this biodiversity loss is related to agriculture, a significant
portion (Tilman et al., 2001a) is due to agriculture-related human impacts on the
environment, primarily the conversion of complex natural systems to simplified
agroecosystems and the intensification of synthetic fertilizer and pesticide use in existing
agroecosystems (Tscharntke et al., 2005). The loss of biodiversity in agricultural

13. 2
landscapes is important not only because of the intrinsic value of plant and animal species,
but because biodiversity provides direct benefits to humans (in the form of food, fiber, and
timber), and also underpins the ecological functions that generate other ecosystem services
(Ceroni et al., 2007; Jarvis et al., 2007; Smukler et al., 2010). Ecosystem services refer to
the benefits that humans derive, directly or indirectly, from the properties and processes of
ecosystems (Costanza et al., 1997). These services include pollination, water regulation and
filtration and CO2 regulation, among others (Costanza et al., 1997; Ceroni et al., 2007).
The conversion of natural ecosystems to agroecosystems has generated debate
about the best approach to ensure the continued provision of ecosystem services while
maintaining or increasing agricultural production. This debate, characterized as “land
sharing” versus “land sparing” compares the alternatives of intensifying existing
agricultural production for high yields while protecting natural areas from agricultural
conversion (“sparing”) and combining agricultural production and conservation on the
same land through ecological farming and agroforestry (“sharing”) (Mas and Dietsch,
2004; Green et al., 2005; Robertson and Swinton, 2005; Matson and Vitousek, 2006;
Phalan et al., 2011; Farley et al., 2012). While these two approaches are often presented as
mutually exclusive, it is likely that any effective conservation strategy, particularly for
conservation of biodiversity in the tropics, will need to integrate elements of both (Dietsch
et al., 2004; Fischer et al., 2008). Tropical landscapes tended to be fragmented, with
patches of natural forest interspersed among patches of urban and agricultural land uses
(Perfecto and Vandermeer, 2008a). Ecology has long recognized that populations exist as
metapopulations, with local extinctions balanced by migrations from and to other areas.

14. 3
The quality of the agricultural “matrix” surrounding natural areas is important for allowing
the movement of organisms between patches, even if it is not ideal habitat itself (Perfecto
and Vandermeer, 2008a; Perfecto and Vandermeer, 2010). Therefore, an ideal scenario is
to “spare” some land from human impacts, while allowing agricultural production and
biodiversity conservation to “share” some land.
Conserving biodiversity within agroecosystems is inevitably more complicated than
maintaining a protected natural area. However, there is evidence that smallholder farmers
in the tropics manage substantial biodiversity on their farms, and there is great potential for
biodiversity conservation within these systems (Thrupp, 2000). Taking advantage of this
potential requires an understanding not only of the ecology of these systems, but of the
interactions between human management and ecology and interactions between farmers’
livelihoods and biodiversity conservation.
In this dissertation, I explore these interactions in agroecosystems managed by
smallholder shade coffee farmers in western El Salvador. Coffee smallholders are of
particular relevance for this research because they manage some of the world’s most
biodiverse agroecosystems while producing a major global commodity. The research was
driven by the following objectives:
1. Understand the factors and livelihood contexts that motivate farmers to conserve
agrobiodiversity
2. Analyze how agrobiodiversity contributes to livelihood outcomes, such as food
security
3. Explore co-benefits between agrobiodiversity and climate change mitigation

15. 4
4. Examine how agrobiodiversity changes over time as a result of farmers’
management choices
1.2. A framework for examining biodiversity-livelihoods interactions among coffee
farming households
1.2.1. The livelihoods concept
In this dissertation, I use the concept of livelihoods to examine the ways in which
biodiversity supports the socioeconomic goals of farmers and their potential motivations
for maintaining biodiversity on their farms. The livelihoods concept has been used to
examine biodiversity conservation in cacao (Dahlquist et al., 2007), coffee (Méndez et al.,
2007; Bacon et al., 2008; Mendez, 2008; Mendez et al., 2010a), maize (Keleman et al.,
2009), and livestock (Devendra and Chantalakhana, 2002; Paris, 2002) agriculture, among
others.
The word “livelihoods” is often preceded by “sustainable” in current discourse,
popularized by Chambers and Conway in their influential 1992 discussion paper for the
Institute of Development Studies (Chambers and Conway, 1992). The most cited definition
of sustainable livelihoods comes out of that paper and numerous adaptations by other
researchers (Carney, 1998; Scoones, 1998; Hussein, 2002):
“A livelihood comprises the capabilities, assets (including both material and social
resources), and activities for a means of living. A livelihood is sustainable when it can cope
with recover from stresses and shocks, maintain or enhance its capabilities and assets,
while not undermining the resource base” (Scoones, 2009).

16. 5
The livelihoods concept provides an alternative to the macro-level, economic- and
technology-focus that came to dominate development policy after World War II (Scoones,
2009), which has been criticized for projecting Northern, industrial concerns onto
Southern, rural people (Chambers, 1995). The livelihoods focus is distinctly micro-level,
with an emphasis on individuals and households, the activities they undertake, and the
factors that affect them. It has been described as “people-oriented” rather than “sector-
oriented” (Scoones, 2009). There is also an emphasis on examining factors other than
employment and income, as the ways in which poor rural people put together and perceive
their livelihoods are not always easily captured by such measures (Chambers and Conway,
1992).
Several authors have put forward an outline of the livelihoods framework for rural
household livelihoods analysis. Concisely, these frameworks commonly include a set of
conditions and trends that form the context of livelihoods (Ellis (2000) also includes shocks
such as drought and war) and a set of livelihood resources (Scoones, 1998) or the
livelihood “platform” (Ellis, 2000), comprised of the natural, economic, human, and social
resources possessed by a household. Access to these assets is modified by institutions,
organizations, and social relationships, resulting in the unique set of livelihood strategies
undertaken by a household. Ellis (2000) separates these livelihood strategies into natural-
resource based (e.g. cultivation of food or non-food items, livestock raising) and non-
natural resource based (e.g. rural manufacture, remittances from emigrated household
members). The portfolio of strategies chosen by a household result in a set of outcomes
which affect both livelihood security (such as income level or household capabilities) and

17. 6
environmental sustainability (such as enhancement or degradation of the resource base)
(Scoones, 1998; Ellis, 2000; Scoones, 2009). The livelihoods concept has been employed
by researchers and development professionals from a wide variety of disciplines with a
wide variety of themes, ranging from livestock cultivation (Paterson and Rojas, 2004), to
tourism (Tao and Wall, 2009), to agroforestry (Dahlquist et al., 2007) proving that it
provides a useful framework for approaching the problem of rural poverty in developing
countries. In this research, I have employed it to analyze how maintenance of biodiversity
on farms fits into households’ livelihoods portfolios, how this strategy influences
livelihood and environmental outcomes.
1.2.2. Scale in biodiversity-livelihoods analysis
I chose the household as the unit of analysis for this study, because it is the
household level at which livelihoods decisions are made that determine the diversity of
plots and farms. It is also at the household level where biodiversity-livelihood interactions
are most apparent (Méndez et al., 2010a). While a large body of literature has examined the
biodiversity on smallholder coffee plantations (Perfecto et al., 1996; Moguel and Toledo,
1999; Perfecto and Vandermeer, 2002; Perfecto et al., 2003; Gordon et al., 2007; Philpott
et al., 2008), little has documented all of the biodiversity managed by coffee farming
households, including homegardens, subsistence crops, and live fences (Méndez et al.,
2010a). A household focus is necessary in order to understand the full range of biodiversity
managed by coffee farmers.

18. 7
The farm was the logical spatial scale to use for this research, because it is at the
plot and farm scales at which most decisions are made in agroecosystems. However,
activities at the plot and farm scales interact ecologically with influences at the landscape
scale, particularly in terms of interactions between biodiversity and ecosystem services.
Many of the ways in which biodiversity benefits livelihoods are via emergent qualities at
the landscape scale that support production at the farm scale. For example, ecologists have
hypothesized that greater species diversity increases the resilience of agroecosystems in
terms of recovering from disturbance (the “insurance hypothesis”), and that this
interactions is most observable at the landscape level (Jackson et al., 2007). Landscape-
level species diversity is determined, in part, by decisions made by land managers (e.g.
farmers) at the plot and farm scales. Landscape-level resilience, in turn, affects
agroecosystem functioning at the farm scale.
Likewise, livelihood decisions at the household scale interact socio-politically with
influences at the regional, national, and global scales via markets, government legislation
and NGO activities, and social networks. For example, price instabilities in global
markets—such as the global crash in coffee prices in 1992 and high corn prices beginning
in 2008—led some farmers to remove coffee plantations in favor of subsistence crops
(Jaffee, 2007; Trujillo, 2008). As another example, the seed supply networks utilized by
small-scale farmers are incredibly complex and operate at many spatial scales (Zimmerer,
2003; Hodgkin et al., 2007), and the genetic diversity maintained by the entire network is
much greater than any one farmer would be able to maintain on his or her farm. Thus,

19. 8
farmers benefit from the genetic diversity maintained over the entire seed supply network,
which may occur at levels from inter-farm to inter-community (Zimmerer, 2003).
1.2.3. Academic fields to examine relationships between biodiversity and livelihoods
in agroecosystems
Analyzing relationships between ecological characteristics and human well-
being is inherently a multidisciplinary undertaking. As a transdisciplinary field,
agroecology (Francis et al., 2008) is well-suited for this, and therefore formed the
backbone for much of the analysis in this thesis. However, I also integrated elements from
landscape ecology in order to address some of the larger-scale elements in this research.
Agroecology
Perhaps the strongest agroecological support for analysis of
agrobiodiversity/livelihood interactions is the concept of using natural ecosystems as a
model and maintaining ecological processes on farms. This concept is at the very heart of
agroecology and the focus of a great deal of agroecological research (Nicholls and Altieri,
2001; Bunch, 2002; Altieri, 2004; Cox et al., 2004; Fujiyoshi et al., 2007; Gliessman,
2007; Moonen and Bàrberi, 2008; Méndez, 2010). Gliessman (2007) identifies population
regulation (particularly with regard to pests), nutrient cycling, and resilience and stability,
and energy flows as the core ecosystem processes to be maintained and monitored in
agroecosystems. The primary “anthropocentric” argument for conservation
agrobiodiversity is that biodiversity—whether in agricultural or natural systems—supports
these ecosystem processes, which provide critical and irreplaceable services to humankind

20. 9
(Ceroni et al., 2007; Jarvis et al., 2007). Thus, agrobiodiversity supports the natural
resource base upon which rural farmers depend for their livelihoods.
This focus on natural ecosystems as a model for agriculture makes agroecology
well-suited to analyze the contribution of plant agrobiodiversity to food security.
Agroecological research has shown that small-scale, biodiverse agriculture can support
high yields and food security, not only at the farm scale but at the global scale. Chappell &
LaValle (2009) have made a particularly compelling argument that not only can biodiverse,
alternative agriculture provide sufficient food for a growing global population, it may be
able to do so more efficiently and with more side benefits for farmers, such as stronger
food sovereignty.
Agroecologists tend to distinguish between two types of biodiversity in agricultural
systems; planned (deliberately included by the land manager) and associated (living or
spending time in the agroecosystem but not deliberately included by the farmer) (Perfecto
and Vandermeer, 2008a). This distinction is useful because each type of biodiversity has
different contributions to food security and may also be influenced differently by farmers’
livelihoods. The contribution of planned biodiversity to food security is more obvious than
that of associated biodiversity. Numerous agroecologists have argued that polycultures,
genetic crop diversity, diverse agroforestry, and inclusion of perennial plants increase the
stability and resilience of agroecosystems (Thrupp, 2000; Holt-Gimenez, 2002), provide a
more varied, nutritious diet (Burlingame and Toledo, 2006; Johns, 2007) and ensure access
to culturally preferred foods (Abbott, 2005; Jaffee, 2007). Evidence for the contribution of

21. 10
associated biodiversity to food security tends to rest more on the fact that low input,
agroecological agricultural systems can provide high yields by taking advantage of
ecological processes (which depend on associated biodiversity) instead of using chemical
fertilizers and pesticides. Empirical studies include those by Bunch (1999, 2002) showing
high maize and bean productivity with agroecological methods in Central America and
Altieri et al. (2005) examining how agroecological methods can support natural pest
predators.
Another of the salient features of agroecology is its focus on traditional knowledge
and its consideration of small-scale, resource-limited farmers as important actors in
agroecological processes. This focus makes agroecology well-suited for examining the
influence of livelihood factors on agrobiodiversity. Agroecological research tends to focus
on small farmers as important agents of agrobiodiversity conservation, examining the
rationale for their management decisions and how these determine the agrobiodiversity of
their farming systems (Altieri and Merrick, 1987; Merrick, 1990; Zimmerer, 2003; Altieri,
2004). Soto-Pinto et al. (2007) examined how farmers’ knowledge and livelihood strategies
influence biodiversity of coffee shade tree canopies, finding that farmers maintained some
trees less suitable as shade trees because they provided other important resources such as
fruit and timber. Méndez et al. (Méndez et al., 2001) investigated similar processes in
homegardens, finding that homegardens varied with the livelihood strategies and needs of
the households that managed them, as well as influenced external forces. Finally, Caceres
(2006) examined how farmers’ diversification strategies impact biodiversity in agricultural
systems.

22. 11
Landscape ecology
Agroecological research tends to focus at the field or plot scale (Dalgaard et al.,
2003). While this made the field well-suited for this research, I also integrated concepts
from landscape ecology when necessary. Landscape ecology is well-equipped to consider
biodiversity conservation at larger spatial scales (Otte et al., 2007). In particular, landscape
ecology presents a potential solution for measuring biodiversity at large scales by making
use of metapopulation dynamics concepts such as the patch-corridor-matrix model, a
concept that has also been adopted by agroecologists (Perfecto and Vandermeer, 2002; Otte
et al., 2007). Landscape ecology also proposes a potential solution for the measurement of
biodiversity across scales far too large for species inventory in the form of landscape
heterogeneity. Benton et al. (2003), for example, describe agricultural intensification and
its impacts on biodiversity at multiple scales, and propose the enhancing habitat
heterogeneity as a key goal for reversing declines in agrobiodiversity.
Landscape ecology is also able to analyze the influence of larger-scale social
processes on agrobiodiversity via examinations of land use and land cover change.
Gottschalk et al. (2007), for example, combined plot-level measures of biodiversity with
landscape-level measures of potential land-use change to estimate the impact of agricultural
subsidies on land-scale level biodiversity. These types of methods are useful for examining
the influence of larger socio-economic processes on agrobiodiversity-food security
interactions.

23. 12
1.3. Causes of biodiversity loss in agroecosystems
Agricultural biodiversity, also called “agrobiodiversity,” refers to “all crops and
livestock, their wild relatives, and the species that interact with and support these species”
(Qualset et al., 1995). Perfecto and Vandeermeer (2008a) separate the biodiversity of
agricultural systems into planned biodiversity (the diversity of plant and animal species
deliberately included by the farmer) and associated biodiversity (the diversity of wild plant
and animal species that colonize the agroecosystem. Most definitions of agrobiodiversity
also include the genetic diversity of crop populations along with the species richness of
crop assemblages (Thrupp, 2000). This thesis focuses on the species and genetic diversity
of cultivated plants (planned plant biodiversity), while recognizing that diverse plant
assemblages can support faunal diversity as well. I use the terms “agricultural biodiversity”
and “agrobiodiversity” interchangeably.
The causes of biodiversity loss in agricultural systems are complex, with interacting
socio-economic and natural factors at different scales. (Perfecto and Vandermeer, 2008a).
At the landscape scale, one of the primary causes of biodiversity loss is increased
homogeneity of farming landscapes (Benton et al., 2003) resulting from the intensification
of agriculture (Jackson et al., 2007). This intensification takes several forms. One is the
physical expansion of cropped areas. The area of the globe devoted to grains, for example,
increased 20 percent from 1950 to 1990 (Ehrlich et al., 1993). Fields and farms have also
become larger, as agricultural industrialization has favored agricultural systems that are
easier to manage with machines than human labor (Jackson et al., 2007). As fields and
farms become larger, the structural composition of the landscape changes as hedgerows,

24. 13
windbreaks, and other field boundaries and non-cropped areas are removed (Benton et al.,
2003; Klimek et al., 2008), which affects species richness (Weibull et al., 2003). This loss
of natural habitat is one of the most direct impacts of agricultural intensification on
biodiversity (McLaughlin and Mineau, 1995).
Intensification also takes the form of biological simplification of agroecosystems.
Industrial agriculture has introduced high-yielding crop varieties that are designed for
simplified management at large scales (Jackson et al., 2007). These varieties have,
however, displaced genetically diverse crop varieties and fostered the large-scale
cultivation of monocultures, causing substantial loss of crop and livestock biodiversity
(Jarvis et al., 2007). It is estimated that 75% of genetic diversity in agricultural crops has
been lost in the past 100 years (Brookfield et al., 2002). Furthermore, these high-yielding
crop varieties generally rely on intensive input application (e.g. synthetic fertilizers and
pesticides), tillage, and irrigation to achieve their high yields (Jackson et al., 2007). The use
of these cultivation methods has consequences for biodiversity, particularly associated
biodiversity such as soil microorganisms and beneficial insects. For example, the long-term
use of fertilizers and tillage has been linked to soil degradation (Aduayi, 1984) and erosion,
with impacts on diversity of soil microorganisms. Fertilizer use also has the indirect effect
of replacing other fertility management strategies such as cover cropping and crop
rotations, the use of which both directly supplies crop species diversity and harbors
associated biodiversity such as beneficial insects (Altieri, 1999). Pesticides and herbicides
have also been shown in some cases to have direct impacts on non-target organisms, such
as killing songbirds on field margins (McLaughlin and Mineau, 1995).

25. 14
The broader socio-economic causes for these changes are complex, however
researchers have attempted to uncover these mechanisms, particularly in areas such as the
tropics where biodiversity loss is acute and particularly important to global sustainability
(Perfecto and Vandermeer, 2008a). Shade coffee agroecosystems have been found to
harbor nearly as much biodiversity as natural forests (Perfecto et al., 1996), and often
comprise a large part of the remaining forest in tropical countries in Central America. In El
Salvador, for example, shade coffee comprises 60% of the remaining forest (Rice and
Ward, 1996). However, much of the coffee grown in Central America has been converted
to sun production, which eliminates the shade canopy and thus much of the biodiversity,
due to production “modernization” programs promoted by Central American governments
during the 1970s and 1980s (Westphal, 2008). The market volatility faced by coffee
farmers also drives the conversion of biodiversity coffee agroforestry systems to other land
uses. During the 1990s, for example, coffee prices plummeted, which forced many coffee
farmers to cut down their coffee plantations and convert them to maize and other field
crops (Jaffee, 2007).
National agricultural policy, particularly in the form of subsidies, can also be an
inadvertent driver of biodiversity loss. Production-based subsidies can be particularly
detrimental to biodiversity as they incentivize high production over other farm-based goods
such as conservation (Boody et al., 2005; Gottschalk et al., 2007; Pascual and Perrings,
2007). Though such subsidies are less common in Central American countries, other types
of subsidies or extension efforts, such as the free distribution of improved seeds, may

26. 15
inadvertently discourage conservation of genetically diverse native seed landraces (Abbott,
2005).
Land tenure critically impacts agrobiodiversity conservation as well, particularly
among small-scale, indigenous, resource-limited farmers who have historically been
excluded from land ownership in Central America. The feudal concentration of land among
a few wealthy families played a large role in the erosion of agrobiodiversity in colonized
countries, as these large estates generally cultivated only a few crops, generally in
monoculture (Faizi and Ravichandran, 2008). In recent years, however, this trend
fortunately seems to be reversing, as land reforms divide up large plantations into small
parcels to be managed by the formerly landless. The Salvadoran coffee cooperative with
whom field work was conducted for this research is an example of this; the farmers
acquired their land in the 1980s and 1990s as part of El Salvador’s land reforms, and now
maintain most of it in coffee under a diverse shade canopy.
1.4. What is the importance of biodiversity in agroecosystems?
Ecologists have hypothesized that biodiversity fosters ecosystem function via two
primary mechanisms. First, species may add to ecosystem function by occupying a unique
niche, complementary to other species in the ecosystem, thus allowing for resource
partitioning or positive intraguild interactions (Hooper et al., 2005; Tscharntke et al.,
2005). This is called species complementarity. The beneficial effects of intercropping are
likely due to complementarity (Vandermeer et al., 2002; Tscharntke et al., 2005), and
researchers have also found that the complementary roles of multiple species may be
responsible for effective biological control of pest insect species (Tscharntke et al., 2005).

27. 16
Second, greater species diversity increases the likelihood that certain individual species
will be present that are particularly important to ecosystem functioning, due either to high
productivity (for plant species) or a particular role such as a dominant natural pest enemy
(Tscharntke et al., 2005). This is referred to as sampling effect. These two mechanisms are
not mutually exclusive, but can be thought of as poles of a continuum of species
interactions (Loreau et al., 2001). Species diversity also increases functional redundancy on
local or landscape scales, which is key to recovery of agroecosystems following
disturbance (Loreau et al., 2001; Tscharntke et al., 2005)
When ecosystems lose biological complexity, energy and nutrient pathways are
lost, which alters ecosystem functioning. In agroecosystems, this translates into the need to
replace natural ecosystem functions with large inputs of energy in the form of fertilizers,
pesticides, and herbicides (Altieri, 1999; Ceroni et al., 2007). For example, biodiversity is
the basis for the population-regulating mechanisms that keep pests in check. When
agroecosystems are biologically simplified, they become more susceptible to pest outbreaks
(Altieri, 1999). Tillage has also been shown to reduce the diversity of soil organisms
(Altieri, 1999), which may reduce efficiency of nutrient cycling, as some studies have
shown that higher diversity of microbial communities increases efficiency of resource use
by microbes (Ceroni et al., 2007). Biologically simplified agroecosystems may also be less
resilient to environmental stresses and shocks such as extreme weather events (Pascual and
Perrings, 2007). Holt-Gimenez (2002), for example, found that farmers making use of more
“sustainable” methods such as use of windbreaks, green manure, and crop rotations (which
also contribute to agrobiodiversity), had higher agroecological resistance to the major

28. 17
disturbance of Hurricane Mitch. Additionally, the highly genetically diverse crop
populations could have advantage over genetically uniform ones in adapting to the effects
of climate change, albeit with variation in adaptation capacity between specific populations
(Mercer & Perales, 2010). Vigoroux et al. (2011), for example, found that phenotypic
adaptation of genetically diverse landraces via human and natural selection have played a
significant part in crop adaptation to climate change in the Sahel.
Agrobiodiversity decline at the farm level also has effects on ecosystem services at
the landscape level. For example, not only does the use of diverse seed varieties benefit
farmers by increasing yield stability, it also maintains the genetic diversity that underpins
all agricultural improvement, which provides public benefit (Jarvis et al., 2007). The
erosion of that genetic diversity has implications for future generations. Agrobiodiversity
also provides habitat for pollinators. The value of pollination as an ecosystem service has
recently become more obvious as bumble bee populations have declined in the United
States and Europe (Goulson et al., 2008). As another example, agrobiodiversity loss may
have consequences for ecosystem services such as water filtration and water regulation.
When forests are removed, stream flows, runoff, and erosion increase, while water
infiltration into the soil decreases (Pimentel et al., 1997). This increases water pollution,
prevents the recharge of aquifers, and causes flooding.
Scientists are also increasingly interested in connections between plant biodiversity
and carbon sequestration in agroecosystems, in the interest of agricultural landscapes
contributing to climate change mitigation. Thus far, research results have been mixed, with
some studies finding a link between species diversity and carbon storage and others finding

29. 18
no relationship. In grasslands, plant species diversity appears to increase primary
productivity, and thus carbon storage in aboveground biomass (Tilman et al., 2001b). This
may (De Deyn et al., 2011) or may not (Reid et al., 2012) translate to higher soil carbon
storage.
The relationship between biodiversity and carbon in agroforestry systems is also
complex. Studies in Panama (Kirby and Potvin, 2007), Kenya (Henry et al., 2009), Mexico
(Soto-Pinto et al., 2010), El Salvador (Méndez et al., 2009), and Indonesia (Kessler et al.,
2012) have found no significant positive relationship between tree species richness and
aboveground or belowground carbon storage. Two studies, however, did find a positive
correlation between species richness and carbon. Woody plant species richness was
associated with higher total C (aboveground and soil) storage in a recent study of Costa
Rican coffee agroforestry systems by Häger (2012). Saha et al. (2009) found a positive
correlation between soil organic C and plant diversity in homegardens of India.
1.5. How does biodiversity support livelihoods?
Many of the ecosystem services supported by agrobiodiversity are of direct
economic, social, or cultural benefit to farmers. Diversity of cultivated crop species, and
diversity of germplasm within those species populations, provides farmers with multiple
sources of food, income, and other products such as firewood and timber, with harvests
spread out over the year (Méndez et al., 2010a). This functions as a form of “insurance”
against total loss of livelihood in the case of market fluctuations or crop failures due to pest
invasions or climatic events (Clawson, 1985; Ceroni et al., 2007). For example, farmers

30. 19
dependent solely on coffee for their livelihood suffered worse during the coffee price crisis
of the 1990s than farmers who also maintained field crops (Jaffee, 2007).
On average, productivity and nutrient use efficiency in ecosystems increases with
species richness (Lepš, 2005). This relationship is highly dependent on environmental
conditions and species composition, but has been demonstrated in agroecosystems as well,
particularly in hayfield systems (Bullock et al., 2001). While polycultures of annual crops
are more complicated than monocultures to manage in intensive, mechanized farming
systems, smallholder farmers often manage highly biodiverse polycultures that yield more
crop per hectare than they would in monoculture. Furthermore, the complementary nutrient
uses in such systems means that farmers may not need to apply as much fertilizer as they
would in monocultures.
Researchers have also hypothesized that agrobiodiversity benefits farming
households in terms of food security, particularly when households depend on their farms
for a substantial portion of their food intake. Some have argued that measures of
agricultural progress must go beyond calories produced per capita and include the
nutritional diversity of cultivated crops (Remans et al., 2011), not only in areas devoted to
subsistence agriculture but in industrializing countries where agricultural and dietary
simplification has led to calorie-rich but nutrient-poor diets (Frison et al., 2011). Research
has tended to focus on dietary diversity as the outcome of interest, because it is the aspect
of food security most likely to be related to agrobiodiversity and also a good proxy measure
of food security (Hoddinott and Yohannes, 2002), especially among children. Dietary
diversity has been shown to correlate with nutrient adequacy (Spigelski, 2004), and child

31. 20
growth (Arimond and Ruel, 2004; Ruel, 2006) in developing countries. So far, no studies
have conclusively demonstrated an impact of agrobiodiversity on dietary diversity or
health, though scientists have shown a relationship between species diversity and
nutritional diversity of cultivated plants (Remans et al., 2011).
Agrobiodiversity provides social and cultural values that are important to farmers
and, at larger scales, to those who live in agricultural landscapes. For example, diverse
landscapes have aesthetic value that provides opportunities for tourism (Ceroni et al., 2007)
and recreational activities. The cultivation of genetically diverse native maize varieties in
Mexico allows farming households to continue cultural traditions by using types of maize
that they prefer for tortilla making, rather than the maize meal of perceived lower quality
that is available through government food support programs (Jaffee, 2007).
Agrobiodiversity also fosters the maintenance of medicinal plants and the knowledge
associated with their use. Research in El Salvador identified 119 species of medicinal
plants cultivated by just 13 households (Méndez et al., 2010a). Medicinal plants are often
the preferred remedies for common ailments in areas where clinics are expensive, difficult
to access, or poorly staffed.
1.6. Efforts to provide incentives for agrobiodiversity conservation
While agrobiodiversity provides ecosystem services critical to agroecosystem
functioning and supports farmer livelihoods, these benefits do not guarantee the
conservation of biodiversity on farms and in agricultural landscapes. There are trade-offs as
well as benefits associated with the maintenance of biodiversity in agroecosystems, and
conserving attributes that are not of direct human value is understandably not always a high

32. 21
priority for farmers, especially those with limited resources. In economic terms, most
ecosystem services resulting from agrobiodiversity—such as pollination and nutrient
cycling—are not excludable (Ceroni and Farley, 2009). In other words, no person can
prevent another from using them. Many ecosystem services—such as hydrologic
regulation—are also non-rival, meaning that use by one person does not leave less of the
service for others to use (Ceroni and Farley, 2009). Markets are only possible for
excludable goods and most efficient for rival goods. Thus, the public (non-excludable, non-
rival) goods tend to be undervalued in favor of market products (Ceroni and Farley, 2009).
Agricultural production is a market product, which explains the destruction of
agrobiodiversity and the non-rival, non-excludable goods it produces in favor of higher on-
farm production.
Because markets cannot provide most ecosystem services resulting from
agrobiodiversity, many researchers have attempted to quantify their value in order to justify
their conservation. This is, however, a complex undertaking due to a lack of adequate
knowledge about how the ecological functions provided by agrobiodiversity translate into
human benefits, particularly with regards to separating the value of biological resources
(e.g. trees) and biological diversity (e.g. the diversity of the tree species) (Jackson et al.,
2007). It is much simpler to assign economic values to biological resources because they
have direct use values; i.e. they are of direct interest to our existing economic system
(Ceroni and Farley, 2009). It is more difficult to assign values to ecosystem services that
have indirect use values, such as ecological stability. This is the case with most ecosystem
services resulting from biodiversity. As a result, only a few studies (e.g. Costanza et al.,

33. 22
1997) have attempted to assign value to these ecosystem services. Non-use values (such as
cultural or social values) are even more difficult to quantify. Interview methods assessing
willingness to pay have been used to assess these values, but people’s willingness to pay
for ecosystem services depends on the existing distribution of income, which might lead to
skewed relative values for rich versus poor people (MartinezAlier, 1995).
Even once the value of agrobiodiversity and its resultant ecosystem services have
been recognized and quantified, the obstacle remains of determining who should pay for
the maintenance of agrobiodiversity. Mechanisms for maintaining agrobiodiversity must
identify the producer and the consumer of the ecosystem services provided by it (Herrador
and Dimas, 2000). Some of the benefits of biodiversity (e.g. higher soil fertility) accrue to
the farmer in the form of increased yields, and are thus often voluntarily maintained by the
farmer, but others (e.g. the maintenance of habitat for a variety of fauna) do not.
Additionally, researchers have noted that threatened species—those most critical to
biodiversity conservation—are rarely those that are most useful to farmers (Gordon et al.,
2003; Méndez et al., 2007).
In an effort to support conservation of ecosystem goods and services not directly
valuable to farmers (including biodiversity), researchers, conservationists, businesses, and
governments have attempted a number of schemes to compensate farmers for these goods
and services. One such scheme is payment for ecosystem services (PES), in which the
beneficiaries of biodiversity conservation directly compensate the providers of this service
(Pascual and Perrings, 2007), thus “internalizing” some of the externalities associated with
agriculture (Pattanayak et al., 2010). These schemes may be public or private. A public

34. 23
scheme has been attempted in El Salvador with water provision from conserved areas, with
some success (Herrador and Dimas, 2000), but required considerable social capital,
willingness to pay, and secure property rights (Pascual and Perrings, 2007; Pattanayak et
al., 2010). PES schemes for biodiversity conservation are made more complicated because
the beneficiaries of biodiversity conservation are often global, not local as is the case with
water provision (Wünscher and Engel, 2012). Furthermore, the relationship between
biodiversity and the services it provides is location-specific, variable, difficult to measure,
and difficult to standardize (Wünscher and Engel, 2012). However, this does not mean that
a PES scheme for agrobiodiversity is impossible. Researchers in Germany successfully
tested a PES scheme in at the regional scale in which farmers bid on contracts for grassland
species diversity conservation in an open auction (Klimek et al., 2008). The bidding
process allowed for revealing the farmers’ true costs of such conservation, removing the
information asymmetries between environmental professionals and farmers that are often
one reason for failure of agrobiodiversity markets (Pascual and Perrings, 2007).
International carbon markets for climate change mitigation such as REDD+ and the
UN Clean Development Mechanism could also provide templates for biodiversity
conservation markets. Though carbon markets for smallholder agriculture are still in their
infancy, researchers and practitioners have already learned important lessons such as the
importance of clear and secure land tenure (Semroc et al., 2012), the need to demonstrate
economic benefits to farmers early in the project, the importance of simple reporting
requirements and capacity building at the regional and national levels (Negra and
Wollenberg, 2012). While it seems unlikely that conservation of agrobiodiversity will be

35. 24
able to garner the political will and financial investment that climate mitigation has, there is
an opportunity to incorporate biodiversity benefits into carbon finance projects for
smallholders, particularly if there are synergies between the two, as some researchers have
suggested (Häger, 2012).
There is a longer history with sustainable coffee certifications for agrobiodiversity
conservation; several such schemes have been developed in the past several decades in
response to growing consumer awareness of environmental and social justice issues
associated with crop production (Méndez et al., 2010b). Certified products are those that
receive a label or certificate from an independent agency, attesting that the product was
grown and/or processed according to a set of standards (Méndez et al., 2010b). Generally,
the price for such products is higher, thus passing along the price of the ecosystem service
to the consumer. Two such certifications—Smithsonian Migratory Bird Center “Bird
Friendly” (which only certifies coffee) and Rainforest Alliance (which certifies coffee,
cocoa, tea, fruit, vegetables, and flowers)—explicitly focus on biodiversity conservation,
requiring farmers to preserve habitat and wildlife. Smithsonian certification requires that
farmers meet organic standards as well, and is more rigorous than Rainforest Alliance
(Méndez et al., 2010b). Both certifications also include social criteria such as safe
conditions for workers and the enforcement of national labor laws (Méndez et al., 2010b).
A third certification, UTZ (formerly UTZ Kapeh), certifies coffee, cocoa, and tea and
integrates environmental and social standards (Méndez et al., 2010b).
Smithsonian, Rainforest Alliance, and UTZ account for a smaller share of the
market than the two more widely known certifications: organic and Fair Trade. Organic

36. 25
certification is regulated by national governments, and standards vary from country to
country. Organic standards generally only regulate the inputs used in crop production, and
research is mixed on whether the practices required for organic certification necessarily
support biodiversity conservation. A meta-analysis published in 2005 showed that, in
general, organic farming practices increased species richness and abundance of most
groups of organisms (e.g. plants, birds, insects, and soil organisms), but effects depended
heavily on the heterogeneity of the surrounding landscape (Bengtsson et al., 2005). Fair
Trade certification is associated with two major certifying organizations: Fairtrade
International (FLO) and Fair Trade USA (formerly part of FLO). Fair Trade focuses
primarily on the trade process and is less concerned with the production process than
Rainforest Alliance, Smithsonian, UTZ, or organic certifications, and thus cannot be
categorized as a true conservation incentive. However, products sold through Fair Trade
channels guarantee a minimum premium to the farmer, along with other benefits such as
capacity-building and a premium to be used for business and community development
(Jaffee, 2007).
Certification schemes have been criticized for failing to provide measureable
benefits to farmers’ livelihoods. Some, such as Rainforest Alliance, do not guarantee a
price premium and are more intended as a marketing tool, which may have little value to
farmers, hence limiting their adoption. Others, such as organic certification, guarantee a
price premium only when there is sufficient demand for the organic product, leaving some
farmers to sell certified organic coffee at conventional prices (Méndez et al., 2010b). It is
also difficult to determine if such certifications result in measurable ecosystem service

37. 26
provision. Certifications establish standards for farmers’ practices, not necessarily
ecosystem services outcomes. Even Smithsonian Bird Friendly certification, which requires
that coffee farmers maintain levels of shade and tree species richness on their farms,
acknowledges that more research is needed to link farming practices to biodiversity levels
(Greenberg and Rice). Partnerships between farmers, researchers, and the coffee industry
can contribute to the conservation value of certification schemes by verifying biodiversity
(Mas and Dietsch, 2004) and livelihood benefits (Méndez et al., 2010b).
1.7. References
Abbott, J.A., 2005. Counting beans: Agrobiodiversity, indigeneity, and agrarian reform.
Prof. Geogr. 57, 198-212.
Aduayi, E.A., 1984. Effects of long-term ammonium-sulfate fertilization on the pH and
elemental distribution at 3 soil depths of southwestern Nigeria soil. Commun. Soil
Sci. Plant Anal. 15, 461-475.
Albrecht, A., Kandji, S.T., 2003. Carbon sequestration in tropical agroforestry systems.
Agriculture Ecosystems & Environment 99, 15-27.
Altieri, M.A., 1999. The ecological role of biodiversity in agroecosystems. Agriculture
Ecosystems & Environment 74, 19-31.
Altieri, M.A., 2004. Linking ecologists and traditional farmers in the search for sustainable
agriculture. Frontiers in Ecology and the Environment 2, 35-42.
Altieri, M.A., Merrick, L.C., 1987. In situ conservation of crop genetic resources through
maintenance of traditional farming systems Econ. Bot. 41, 86-96.
Altieri, M.A., Ponti, L., Nicholls, C.I., 2005. Manipulating vineyard biodiversity for
improved insect pest management: case studies from northern California.
International Journal of Biodiversity Science & Management 1, 191-203.
Arimond, M., Ruel, M.T., 2004. Dietary diversity is associated with child nutritional status:
Evidence from 11 Demographic and Health Surveys. Journal of Nutrition 134,
2579-2585.

38. 27
Azadbakht, L., Esmaillzadeh, A., 2012. Dietary energy density is favorably associated with
dietary diversity score among female university students in Isfahan. Nutrition 28,
991-995.
Bacon, C., Mendez, V.E., Gliessman, S.R., Goodman, D., Fox, J.A. (Eds.), 2008.
Confronting the coffee crisis: Fair trade, sustainable livelihoods and ecosystems in
Mexico and Central America. MIT Press, Cambridge, MA.
Bebbington, A., 2001. Globalized Andes? Livelihoods, Landscapes and Development.
Cultural Geographies 8, 414-436.
Bebbington, A.J., Batterbury, S.P.J., 2001. Transnational livelihoods and landscapes:
Political ecologies of globalization. Ecumene 8, 369-380.
Becquey, E., Martin-Prevel, Y., Traissac, P., Dembele, B., Bambara, A., Delpeuch, F.,
2010. The household food insecurity access scale and an index-member dietary
diversity score contribute valid and complementary information on household food
insecurity in an urban West-African setting. The Journal of Nutrition 140, 2233-
2240.
Belanger, J., Johns, T., 2008. Biological Diversity, Dietary Diversity, and Eye Health in
Developing Country Populations: Establishing the Evidence-base. EcoHealth 5,
244-256.
Bengtsson, J., Ahnstrom, J., Weibull, A.C., 2005. The effects of organic agriculture on
biodiversity and abundance: a meta-analysis. J. Appl. Ecol. 42, 261-269.
Benton, T.G., Vickery, J.A., Wilson, J.D., 2003. Farmland biodiversity: is habitat
heterogeneity the key? Trends in Ecology & Evolution 18, 182-188.
Blackman, A., Albers, H.J., Avalos-Sartorio, B., Murphy, L.C., 2008. Land cover in a
managed forest ecosystem: Mexican shade coffee. Am. J. Agr. Econ. 90, 216-231.
Blackman, A., Avalos-Sartorio, B., Chow, J., 2007. Shade coffee & tree cover loss -
Lessons from El Salvador. Environment 49, 22-32.
Boody, G., Vondracek, B., Andow, D.A., Krinke, M., Westra, J., Zimmerman, J., Welle, P.,
2005. Multifunctional agriculture in the United States. Bioscience 55, 27-38.
Brookfield, H., Padoch, C., Parsons, H., Stocking, M., 2002. Cultivating biodiversity:
setting the scene. In: Brookfield, H., Padoch, C., Parsons, H., Stocking, M. (Eds.),
Cultivating Biodiversity: Understanding, Analysing, & Using Agricultural
Diversity. ITDG Publishing, London, pp. 1-8.
Bullock, J.M., Pywell, R.F., Burke, M.J.W., Walker, K.J., 2001. Restoration of biodiversity
enhances agricultural production. Ecol. Lett. 4, 185-189.

39. 28
Bunch, R., 1999. More productivity with fewer external inputs: Central American case
studies of agroecological development and their broader implications. Environment,
Development, and Sustainability 1, 219-233.
Bunch, R., 2002. Increasing productivity through agroecological approaches in Central
America: experiences from hillside agriculture. In: Uphoff, N. (Ed.),
Agroecological innovations: Increasing food production with participatory
development. Earthscan, London, UK.
Burlingame, B., Toledo, A., 2006. Biodiversity and nutrition: A common path toward
global food security and sustainable development. Journal of Food Composition and
Analysis 19, 477-483.
Butchart, S.H.M., Walpole, M., Collen, B., van Strien, A., Scharlemann, J.P.W., Almond,
R.E.A., Baillie, J.E.M., Bomhard, B., Brown, C., Bruno, J., Carpenter, K.E., Carr,
G.M., Chanson, J., Chenery, A.M., Csirke, J., Davidson, N.C., Dentener, F., Foster,
M., Galli, A., Galloway, J.N., Genovesi, P., Gregory, R.D., Hockings, M., Kapos,
V., Lamarque, J.-F., Leverington, F., Loh, J., McGeoch, M.A., McRae, L.,
Minasyan, A., Morcillo, M.H., Oldfield, T.E.E., Pauly, D., Quader, S., Revenga, C.,
Sauer, J.R., Skolnik, B., Spear, D., Stanwell-Smith, D., Stuart, S.N., Symes, A.,
Tierney, M., Tyrrell, T.D., Vie, J.-C., Watson, R., 2010. Global Biodiversity:
Indicators of Recent Declines. Science 328, 1164-1168.
Caceres, D.M., 2006. Agrobiodiversity and technology in resource-poor farms. Interciencia
31, 403-410.
Canadell, J.G., Raupach, M.R., 2008. Managing forests for climate change mitigation.
Science 320, 1456-1457.
Carney, D. (Ed), 1998. Sustainable Rural Livelihoods: What Contribution can we Make?
Department for International Development, London, UK.
Ceroni, M., Farley, J., 2009. Ecological economics of biodiversity: Synthesis document. In:
Gund Institute for Ecological Economics, U.o.V. (Ed.), Burlington, VT.
Ceroni, M., Liu, S., Costanza, R., 2007. Ecological and economic roles of biodiversity in
agroecosystems. In: Jarvis, D.I., Padoch, C., Cooper, H.D. (Eds.), Managing
Biodiversity in Agricultural Systems. Columbia University Press, New York, pp.
338-361.
Chambers, R., 1995. Poverty and livelihoods: Whose reality counts? Institute of
Development Studies, Brighton, England.
Chambers, R., Conway, G.R., 1992. Sustainable rural livelihoods: practical concepts for the
21st century. IDS Discussion Paper. Institute of Development Studies.

40. 29
Chappell, M.J., LaValle, L.A., 2009. Food security and biodiversity: can we have both? An
agroecological analysis. Agric. Human Values 28, 3-26.
Chave, J., Condit, R., Lao, S., Caspersen, J.P., Foster, R.B., Hubbell, S., 2003. Spatial and
temporal variation of biomass in a tropical forest: Results from a large census plot
in Panama. Journal of Ecology 91, 240-252.
Clawson, D.L., 1985. Harvest security and intraspecific diversity in traditional tropical
agriculture. Econ. Bot. 39, 56-67.
Costanza, R., dArge, R., deGroot, R., Farber, S., Grasso, M., Hannon, B., Limburg, K.,
Naeem, S., Oneill, R.V., Paruelo, J., Raskin, R.G., Sutton, P., vandenBelt, M., 1997.
The value of the world's ecosystem services and natural capital. Nature 387, 253-
260.
Cox, T.S., Picone, C., Jackson, W., 2004. Research Priorities in Natural Systems
Agriculture. Journal of Crop Improvement 12, 511.
Dahlquist, R.M., Whelan, M.P., Winowiecki, L., Polidoro, B., Candela, S., Harvey, C.A.,
Wulfhorst, J.D., McDaniel, P.A., Bosque-Perez, N.A., 2007. Incorporating
livelihoods in biodiversity conservation: a case study of cacao agroforestry systems
in Talamanca, Costa Rica. Biodiversity and Conservation 16, 2311-2333.
Dalgaard, T., Hutchings, N.J., Porter, J.R., 2003. Agroecology, scaling and
interdisciplinarity. Agriculture Ecosystems & Environment 100, 39-51.
Davis, A., Méndez, V.E., 2011. Prioritizing food security and livelihoods in climate change
mitigation mechanisms: experiences and opportunities for smallholder coffee
agroforestry, forest communities and REDD+. Salvadoran Research Program on
Development and Environment (PRISMA), San Salvador, El Salvador.
De Deyn, G.B., Shiel, R.S., Ostle, N.J., McNamara, N.P., Oakley, S., Young, I., Freeman,
C., Fenner, N., Quirk, H., Bardgett, R.D., 2011. Additional carbon sequestration
benefits of grassland diversity restoration. J. Appl. Ecol. 48, 600-608.
Devendra, C., Chantalakhana, C., 2002. Animals, poor people and food insecurity:
opportunities for improved livelihoods through efficient natural resource
management. Outlook on Agriculture 31, 161-175.
Dietsch, T.V., Philpott, S.M., Rice, R.A., Greenberg, R., Bichier, P., 2004. Conservation
policy in coffee landscapes. Science 303, 625-625.
Ehrlich, P.R., Ehrlich, A.H., Daily, G.C., 1993. Food Security, Population and
Environment. Population and Development Review 19, 1-32.
Ellis, F., 2000. Rural livelihoods and diversity in developing countries. Oxford University
Press, Oxford, UK.

41. 30
Faizi, S., Ravichandran, M., 2008. Biodiversity and land reforms: a neglected linkage.
Biodiversity and Conservation 17, 2817-2819.
Farley, J., Schmitt F., A., Alvez, J., Ribeiro de Freitas Jr., N., 2012. How Valuing Nature
Can Transform Agriculture. Solutions 2, 64-73.
Fischer, J., Brosi, B., Daily, G.C., Ehrlich, P.R., Goldman, R., Goldstein, J., Lindenmayer,
D.B., Manning, A.D., Mooney, H.A., Pejchar, L., Ranganathan, J., Tallis, H., 2008.
Should agricultural policies encourage land sparing or wildlife-friendly farming?
Frontiers in Ecology and the Environment 6, 380-385.
Foley, J.A., Ramankutty, N., Brauman, K.A., Cassidy, E.S., Gerber, J.S., Johnston, M.,
Mueller, N.D., O/'Connell, C., Ray, D.K., West, P.C., Balzer, C., Bennett, E.M.,
Carpenter, S.R., Hill, J., Monfreda, C., Polasky, S., Rockstrom, J., Sheehan, J.,
Siebert, S., Tilman, D., Zaks, D.P.M., 2011. Solutions for a cultivated planet.
Nature 478, 337-342.
Francis, C.A., Lieblein, G., Breland, T.A., Salomonsson, L., Geber, U., Sriskandarajah, N.,
Langer, V., 2008. Transdisciplinary research for a sustainable agriculture and food
sector. Agronomy Journal 100, 771-776.
Frison, E.A., Cherfas, J., Hodgkin, T., 2011. Agricultural Biodiversity Is Essential for a
Sustainable Improvement in Food and Nutrition Security. Sustain. 3, 238-253.
Frison, E.A., Smith, I.F., Johns, T., Cherfas, J., Eyzaguirre, P.B., 2006. Agricultural
biodiversity, nutrition, and health: Making a difference to hunger and nutrition in
the developing world. Food Nutr. Bull. 27, 167-179.
Fujita, M., Lo, Y.J., Baranski, J.R., 2012. Dietary diversity score is a useful indicator of
vitamin A status of adult women in Northern Kenya. American Journal of Human
Biology 24, 829-834.
Fujiyoshi, P.T., Gliessman, S.R., Langenheim, J.H., 2007. Factors in the suppression of
weeds by squash interplanted in corn. Weed Biology and Management 7, 105-114.
Gliessman, S.R., 2007. Agroecology: The Ecology of Sustainable Food Systems. CRC
Press, New York.
Godfray, H.C.J., Beddington, J.R., Crute, I.R., Haddad, L., Lawrence, D., Muir, J.F.,
Pretty, J., Robinson, S., Thomas, S.M., Toulmin, C., 2010. Food Security: The
Challenge of Feeding 9 Billion People. Science 327, 812-818.
Gordon, C., Manson, R., Sundberg, J., Cruz-Angon, A., 2007. Biodiversity, profitability,
and vegetation structure in a Mexican coffee agroecosystern. Agriculture
Ecosystems & Environment 118, 256-266.

42. 31
Gordon, J.E., Barrance, A.J., Schreckenberg, K., 2003. Are rare species useful species?
Obstacles to the conservation of tree diversity in the dry forest zone agro-
ecosystems of Mesoamerica. Global Ecology and Biogeography 12, 13-19.
Gottschalk, T.K., Diekotter, T., Ekschmitt, K., Weinmann, B., Kuhlmann, F., Purtauf, T.,
Dauber, J., Wolters, V., 2007. Impact of agricultural subsidies on biodiversity at the
landscape level. Landscape Ecology 22, 643-656.
Goulson, D., Lye, G.C., Darvill, B., 2008. Decline and conservation of bumble bees. Annu.
Rev. Entomol. 53, 191-208.
Green, R.E., Cornell, S.J., Scharlemann, J.P.W., Balmford, A., 2005. Farming and the fate
of wild nature. Science 307, 550-555.
Greenberg, R., Rice, R., Shade management criteria for Bird Friendly coffee. Smithsonian
Migratory Bird Center.
Haeger, A., 2012. The effects of management and plant diversity on carbon storage in
coffee agroforestry systems in Costa Rica. Agroforestry Systems 86, 159-174.
Häger, A., 2012. The effects of management and plant diversity on carbon storage in coffee
agroforestry systems in Costa Rica. Agroforestry Systems 86, 159-174.
Hecht, S.B., Kandel, S., Gomes, I., Cuellar, N., Rosa, H., 2006. Globalization, forest
resurgence, and environmental politics in El Salvador. World Development 34, 308-
323.
Henry, M., Tittonell, P., Manlay, R.J., Bernoux, M., Albrecht, A., Vanlauwe, B., 2009.
Biodiversity, carbon stocks and sequestration potential in aboveground biomass in
smallholder farming systems of western Kenya. Agriculture, Ecosystems &
Environment 129, 238-252.
Herrador, D., Dimas, L., 2000. Payment for environmental services in El Salvador. Mt.
Res. Dev. 20, 306-309.
Hoddinott, J., Yohannes, Y., 2002. Dietary diversity as a food security indicator. FCND
Discussion Papers. International Food Policy Research Institute (IFPRI),
Washington, D.C.
Hodgkin, T., Rana, R., Tuxill, J., Balma, D., Subedi, A., Mar, I., Karamura, D., Valdivia,
R., Collado, L., Latournerie, L., Sadiki, M., Sawadogo, M., Brown, A.H.D., Jarvis,
D.I., 2007. Seed systems and crop genetic diversity in agroecosystems. In: Jarvis,
D.I., Padoch, C., Cooper, H.D. (Eds.), Managing Biodiversity in Agricultural
Systems. Columbia University Press, New York, pp. 77-116.
Holdridge, L.R., 1987. Ecologia basada en zonas de vida (Ecology based on life zones).
Interamerican Institute for Cooperation on Agriculture, San Jose, Costa Rica.

43. 32
Holt-Gimenez, E., 2002. Measuring farmers' agroecological resistance after Hurricane
Mitch in Nicaragua: a case study in participatory, sustainable land management
impact monitoring. Agriculture Ecosystems & Environment 93, 87-105.
Hooper, D.U., Chapin, F.S., Ewel, J.J., Hector, A., Inchausti, P., Lavorel, S., Lawton, J.H.,
Lodge, D.M., Loreau, M., Naeem, S., Schmid, B., Setala, H., Symstad, A.J.,
Vandermeer, J., Wardle, D.A., 2005. Effects of biodiversity on ecosystem
functioning: A consensus of current knowledge. Ecol. Monogr. 75, 3-35.
Hussein, K., 2002. Livelihoods Approaches Compared: A Multi-Agency Review of
Current Practice. In: Sattaur, O. (Ed.). Department for International Development,
London.
INCAP, 2012. Tabla de composicion de alimentos para Centroamerica del INCAP (INCAP
Table of food composition for Central America). Instituto de Nutrición de Centro
América y Panamá (Nutrition Institute of Central America and Panama), Guatemala
City, Guatemala.
Jackson, L.E., Pascual, U., Hodgkin, T., 2007. Utilizing and conserving agrobiodiversity in
agricultural landscapes. Agriculture, Ecosystems & Environment 121, 196-210.
Jaffee, D., 2007. Brewing justice: Fair trade coffee, sustainability, and survival. University
of California Press, Berkeley, CA.
Jarvis, D.I., Padoch, C., Cooper, H.D., 2007. Biodiversity, agriculture, and ecosystem
services. In: Jarvis, D.I., Padoch, C., Cooper, H.D. (Eds.), Managing Biodiversity in
Agricultural Systems. Columbia University Press, New York, pp. 338-361.
Jha, S., Bacon, C.M., Philpott, S.M., Rice, R.A., Méndez, V.E., Läderach, P., 2011a. A
review of ecosystem services, farmer livelihoods, and value chains in shade coffee
agroecosystems. In: Campbell, B.W., Lopez-Ortiz, S. (Eds.), Integrating
Agriculture, Conservation, an Ecotourism: Examples from the Field. Springer
Academic Publishers, New York and Berlin, Germany.
Jha, S., Bacon, C.M., Philpott, S.M., Rice, R.A., Mendez, V.E., Laederach, P., 2011b. A
Review of Ecosystem Services, Farmer Livelihoods, and Value Chains in Shade
Coffee Agroecosystems. In: Campbell, W.B., Ortiz, S.L. (Eds.), Integrating
Agriculture, Conservation and Ecotourism: Examples from the Field, pp. 141-208.
Johns, T., 2007. Agrobiodiversity, diet, and human health. In: Jarvis, D.I., Padoch, C.,
Cooper, H.D. (Eds.), Managing Biodiversity in Agricultural Systems. Biodiversity
International, New York, pp. 382-406.
Keleman, A., Hellin, J., Bellon, M.R., 2009. Maize diversity, rural development policy, and
farmers' practices: lessons from Chiapas, Mexico. Geogr. J. 175, 52-70.

44. 33
Kessler, M., Hertel, D., Jungkunst, H.F., Kluge, J., Abrahamczyk, S., Bos, M., Buchori, D.,
Gerold, G., Gradstein, S.R., Koehler, S., Leuschner, C., Moser, G., Pitopang, R.,
Saleh, S., Schulze, C.H., Sporn, S.G., Steffan-Dewenter, I., Tjitrosoedirdjo, S.S.,
Tscharntke, T., 2012. Can Joint Carbon and Biodiversity Management in Tropical
Agroforestry Landscapes Be Optimized? PLoS ONE 7.
Kirby, K.R., Potvin, C., 2007. Variation in carbon storage among tree species: Implications
for the management of a small-scale carbon sink project. For. Ecol. Manage. 246,
208-221.
Klimek, S., Kemmermann, A.R., Steinmann, H.H., Freese, J., Isselstein, J., 2008.
Rewarding farmers for delivering vascular plant diversity in managed grasslands: A
transdisciplinary case-study approach. Biological Conservation 141, 2888-2897.
Komar, O., 2006. Ecology and conservation of birds in coffee plantations: a critical review.
Bird Conservation International 16, 1-23.
Lepš, J., 2005. Diversity and ecosystem function. In: Maarel, E.v.d. (Ed.), Vegetation
Ecology. Blackwell Publishing, Oxford, UK.
Lin, B.B., 2010. The role of agroforestry in reducing water loss through soil evaporation
and crop transpiration in coffee agroecosystems. Agricultural and Forest
Meteorology 150, 510-518.
Lo, Y.T., Chang, Y.H., Lee, M.S., Wahlqvist, M.L., 2012. Dietary diversity and food
expenditure as indicators of food security in older Taiwanese. Appetite 58, 180-187.
Loreau, M., Naeem, S., Inchausti, P., Bengtsson, J., Grime, J.P., Hector, A., Hooper, D.U.,
Huston, M.A., Raffaelli, D., Schmid, B., Tilman, D., Wardle, D.A., 2001. Ecology -
Biodiversity and ecosystem functioning: Current knowledge and future challenges.
Science 294, 804-808.
Maas, B., Putra, D.D., Waltert, M., Clough, Y., Tscharntke, T., Schulze, C.H., 2009. Six
years of habitat modification in a tropical rainforest margin of Indonesia do not
affect bird diversity but endemic forest species. Biological Conservation 142, 2665-
2671.
Magurran, A.E., 2006. Measuring Biological Diversity. Blackwell Publishing, Malden, Ma.
MARN, 2013. Mapas del Sistema de Información Ambiental.
MartinezAlier, J., 1995. Distributional issues in ecological economics. Review of Social
Economy 53, 511-528.
Mas, A.H., Dietsch, T.V., 2004. Linking shade coffee certification to biodiversity
conservation: butterflies and birds in Chiapas, Mexico. Ecol. Appl. 14, 642-654.

45. 34
Matson, P.A., Vitousek, P.M., 2006. Agricultural intensification: Will land spared from
farming be land spared for nature? Conservation Biology 20, 709-710.
Maxwell, D., Ahiadeke, C., Levin, C., Armar-Klemesu, M., Zakariah, S., Lamptey, G.M.,
1999. Alternative food-security indicators: revisiting the frequency and severity of
'coping strategies'. Food Policy 24, 411-429.
Maxwell, D., Caldwell, R., Langworthy, M., 2008. Measuring food insecurity: Can an
indicator based on localized coping behaviors be used to compare across contexts?
Food Policy 33, 533-540.
Maxwell, D., Watkins, B., Wheeler, R., Collins, G., 2003. The coping strategies index: A
tool for rapidly measuring food security and the impact of food aid programs in
emergencies. FAO international workshop on "Food security in complex
emergencies: building policy frameworks to address longer-term programming
challenges". Food and Agriculture Organization of the United Nations, Tivoli, Italy.
Maxwell, S., 1996. Food security: A post-modern perspective. Food Policy 21, 155-170.
McLaughlin, A., Mineau, P., 1995. The impact of agricultural practices on biodiversity.
Agriculture Ecosystems & Environment 55, 201-212.
Méndez, V.E., 2008. Farmer livelihoods and biodiversity conservation in a coffee
landscape of El Salvador. In: Bacon, C., Mendez, V.E., Gliessman, S.R., Goodman,
D., Fox, J.A. (Eds.), Confronting the coffee crisis: sustaining livelihoods and
ecosystems in Mexico and Central America. MIT Press, Cambridge, MA, U.S.A.,
pp. 207-234.
Méndez, V.E., 2010. Agroecology. In: Warf, B. (Ed.), Encyclopedia of Geography. Sage
Publishers, Thousand Oaks, CA.
Méndez, V.E., 2004. Traditional shade, rural livelihoods and conservation in small coffee
farms and cooperatives of western El Salvador., Department of Environmental
Studies. University of California Santa Cruz, Santa Cruz, CA.
Méndez, V.E., Bacon, C.M., Olson, M., Morris, K.S., Shattuck, A., 2010a.
Agrobiodiversity and Shade Coffee Smallholder Livelihoods: A Review and
Synthesis of Ten Years of Research in Central America. Prof. Geogr. 62, 357-376.
Méndez, V.E., Bacon, C.M., Olson, M., Petchers, S., Herrador, D., Carranza, C., Trujillo,
L., Guadarrama-Zugasti, C., Cordon, A., Mendoza, A., 2010b. Effects of Fair Trade
and organic certifications on small-scale coffee farmer households in Central
America and Mexico. Renewable Agriculture and Food Systems 25, 236-251.
Méndez, V.E., Castro-Tanzi, S., Goodall, K., Morris, K.S., Bacon, C.M., Läderach, P.,
Morris, W.B., Georgeoglou-Laxalde, M.U., 2012. Livelihood and environmental
trade-offs of climate mitigation in smallholder coffee agroforestry systems. In:

46. 35
Wollenberg, E.K., Nihart, A., Grieg-Gran, M., Tapio-Biström, M.L. (Eds.), Climate
Mitigation and Agriculture. Earthscan, London, U.K.
Méndez, V.E., Gliessman, S.R., Gilbert, G.S., 2007. Tree biodiversity in farmer
cooperatives of a shade coffee landscape in western El Salvador. Agriculture
Ecosystems & Environment 119, 145-159.
Méndez, V.E., Lok, R., Somarriba, E., 2001. Interdisciplinary analysis of homegardens in
Nicaragua: micro-zonation, plant use and socioeconomic importance. Agroforestry
Systems 51, 85-96.
Méndez, V.E., Shapiro, E.N., Gilbert, G.S., 2009. Differentiated Cooperative Management
and its Effects on Shade Tree Diversity and Soil Properties of Coffee Plantations in
western El Salvador. Agroforestry Systems 76, 111-126.
Merrick, L.C., 1990. Crop genetic diversity and its conservation in traditional
agroecosystems. In: Altieri, M. (Ed.), Agroecology and Small Farm Development.
CRC Press, Boston, pp. 3-17.
Moguel, P., Toledo, V.M., 1999. Biodiversity conservation in traditional coffee systems of
Mexico. Conservation Biology 13, 11-21.
Montagnini, F., Nair, P.K.R., 2004. Carbon sequestration: An underexploited
environmental benefit of agroforestry systems. Agroforestry Systems 61-2, 281-
295.
Moonen, A.-C., Bàrberi, P., 2008. Functional biodiversity: An agroecosystem approach.
Agriculture, Ecosystems & Environment 127, 7-21.
Morris, K.S., Olson, M.B., Mendez, V.E., 2013. 'Los meses flacos': seasonal food
insecurity in a Salvadoran organic coffee cooperative. Journal of Peasant Studies In
press.
Nakakaawa, C., Aune, J., Vedeld, P., 2010. Changes in carbon stocks and tree diversity in
agro-ecosystems in south western Uganda: what role for carbon sequestration
payments? New For. 40, 19-44.
Negra, C., Wollenberg, E., 2012. Lessons learned from REDD for agriculture. In:
Wollenberg, E.K., Nihart, A., Grieg-Gran, M., Tapio-Biström, M.L. (Eds.), Climate
Mitigation and Agriculture. Earthscan, London, U.K.
Nicholls, C.A., Altieri, M., 2001. Manipulating plant biodiversity to enhance biological
control of insect pests: A case study of a northern California organic vineyard. In:
Gliessman, S. (Ed.), Agroecosystem Sustainability: Developing Practical Strategies.
CRC Press, New York, pp. 29-50.

47. 36
Nonato de Souza, H., Graaff, J.d., Pulleman, M.M., 2011. Strategies and economics of
farming systems with coffee in the Atlantic Rainforest Biome (Online first).
Agroforestry Systems.
Otte, A., Simmering, D., Wolters, V., 2007. Biodiversity at the landscape level: recent
concepts and perspectives for multifunctional land use. Landscape Ecology 22, 639-
642.
Paris, T.R., 2002. Crop-animal systems in Asia: socio-economic benefits and impacts on
rural livelihoods. Agricultural Systems 71, 147-168.
Pascual, U., Perrings, C., 2007. Developing incentives and economic mechanisms for in
situ biodiversity conservation in agricultural landscapes. Agriculture Ecosystems &
Environment 121, 256-268.
Paterson, R.T., Rojas, F., 2004. Poultry, pigs, hair sheep and guinea pigs in the livelihoods
of small-scale, subsistence farmers in tropical Bolivia. Small stock in development
workshop: Enhancing the contribution of small livestock to the livelihoods of
resource-poor communities. Natural Resources International Ltd., Hotel Brovad,
Masaka, Uganda.
Pattanayak, S.K., Wunder, S., Ferraro, P.J., 2010. Show Me the Money: Do Payments
Supply Environmental Services in Developing Countries? Review of
Environmental Economics and Policy 4, 254-274.
Perfecto, I., Mas, A., Dietsch, T., Vandermeer, J., 2003. Conservation of biodiversity in
coffee agroecosystems: a tri-taxa comparison in southern Mexico. Biodiversity and
Conservation 12, 1239-1252.
Perfecto, I., Rice, R.A., Greenberg, R., VanderVoort, M.E., 1996. Shade coffee: A
disappearing refuge for biodiversity. Bioscience 46, 598-608.
Perfecto, I., Vandermeer, J., 2002. Quality of agroecological matrix in a tropical montane
landscape: Ants in coffee plantations in southern Mexico. Conservation Biology 16,
174-182.
Perfecto, I., Vandermeer, J., 2008a. Biodiversity conservation in tropical agroecosystems -
A new conservation paradigm. Year in Ecology and Conservation Biology 2008.
Blackwell Publishing, Oxford, pp. 173-200.
Perfecto, I., Vandermeer, J., 2008b. Spatial pattern and ecological process in the coffee
agroforestry system. Ecology 89, 915-920.
Perfecto, I., Vandermeer, J., 2010. The agroecological matrix as alternative to the land-
sparing/agriculture intensification model. Proceedings of the National Academy of
Sciences 107, 5786-5791.

48. 37
Petchey, O.L., Gaston, K.J., 2002. Functional diversity (FD), species richness and
community composition. Ecol. Lett. 5, 402-411.
Phalan, B., Onial, M., Balmford, A., Green, R.E., 2011. Reconciling Food Production and
Biodiversity Conservation: Land Sharing and Land Sparing Compared. Science
333, 1289-1291.
Philpott, S.M., Arendt, W.J., Armbrecht, I., Bichier, P., Diestch, T.V., Gordon, C.,
Greenberg, R., Perfecto, I., Reynoso-Santos, R., Soto-Pinto, L., Tejeda-Cruz, C.,
Williams-Linera, G., Valenzuela, J., Zolotoff, J.M., 2008. Biodiversity Loss in
Latin American Coffee Landscapes: Review of the Evidence on Ants, Birds, and
Trees. Conservation Biology 22, 1093-1105.
Pimentel, D., McNair, M., Duck, L., Pimentel, M., Kamil, J., 1997. The value of forests to
world food security. Human Ecology 25, 91-120.
Potvin, C., Gotelli, N.J., 2008. Biodiversity enhances individual performance but does not
affect survivorship in tropical trees. Ecol. Lett. 11, 217-223.
Purdue, 2012. New Crops Resource Online Program. Purdue University Center for New
Crops and Plant Products, West Lafayette, Indiana.
Qualset, C., McGuire, P., Warburton, M., 1995. In California: ‘Agrobiodiversity’ key to
agricultural productivity. California Agriculture 49, 45-49.
Rapidel, B., DeClerck, F., Coq, J.F.L., Beer, J. (Eds.), 2011. Ecosystem Services from
Agriculture an Agroforestry. Earthscan, London, U.K.
Washington, D.C.
Reid, J.P., Adair, E.C., Hobbie, S.E., Reich, P.B., 2012. Biodiversity, Nitrogen Deposition,
and CO2 Affect Grassland Soil Carbon Cycling but not Storage. Ecosystems 15,
580-590.
Remans, R., Flynn, D.F.B., DeClerck, F., Diru, W., Fanzo, J., Gaynor, K., Lambrecht, I.,
Mudiope, J., Mutuo, P.K., Nkhoma, P., Siriri, D., Sullivan, C., Palm, C.A., 2011.
Assessing Nutritional Diversity of Cropping Systems in African Villages. PLoS
ONE 6.
Rice, R.A., Ward, J.R., 1996. Coffee, conservation, and commerce in the western
hemisphere. Smithsonian Migratory Bird Center/Natural Resources Defence
Council, Washington, D.C.
Robertson, G.P., Swinton, S.M., 2005. Reconciling agricultural productivity and
environmental integrity: a grand challenge for agriculture. Frontiers in Ecology and
the Environment 3, 38-46.

49. 38
Ruel, M.T., 2003. Is dietary diversity an indicator of food security or dietary quality? A
review of measurement issues and research needs. Food Nutr. Bull. 24, 231-232.
Ruel, M.T., 2006. Operationalizing dietary diversity: A review of measurement issues and
research priorities. Journal of Nutrition 133, 3911S-3926S.
Saha, S.K., Nair, P.K.R., Nair, V.D., Kumar, B.M., 2009. Soil carbon stock in relation to
plant diversity of homegardens in Kerala, India. Agroforestry Systems 76, 53-65.
Schmitt-Harsh, M., Evans, T.P., Castellanos, E., Randolph, J.C., 2012. Carbon stocks in
coffee agroforests and mixed dry tropical forests in the western highlands of
Guatemala. Agroforestry Systems 86, 141-157.
Scoones, I., 1998. Sustainable Rural Livelihoods: A Framework for Analysis. Institute of
Development Studies, Brighton, UK.
Scoones, I., 2009. Livelihoods perspectives and rural development. Journal of Peasant
Studies 36, 171-196.
Semroc, B.L., Schroth, G., Harvey, C.A., Zepeda, Y., Hills, T., Lubis, S., Arief, C.W.,
Zerbock, O., Boltz, F., 2012. Climate change mitigation in agroforestry systems:
linking smallholders to forest carbon markets. In: Wollenberg, E.K., Nihart, A.,
Grieg-Gran, M., Tapio-Biström, M.L. (Eds.), Climate Mitigation and Agriculture.
Earthscan, London, U.K.
Smith, P., Gregory, P.J., 2013. Climate change and sustainable food production. The
Proceedings of the Nutrition Society 72, 21-28.
Smukler, S.M., Sanchez-Moreno, S., Fonte, S.J., Ferris, H., Klonsky, K., O'Geen, A.T.,
Scow, K.M., Steenwerth, K.L., Jackson, L.E., 2010. Biodiversity and multiple
ecosystem functions in an organic farmscape. Agriculture Ecosystems &
Environment 139, 80-97.
Somarriba, E., Harvey, C., Samper, M., Anthony, F., Gonzalez, J., Staver, C., Rice, R.,
2004. Biodiversity conservation in neotropical coffee (Coffea arabica) plantations.
In: Schroth, G., Foseca, G., Harvey, C.A., Gascon, C., Vasconcelos, H., Izac,
A.M.N. (Eds.), Agroforestry and Biodiversity Conservation in Tropical Landscapes.
Island Press, Washington, D.C.
Soto-Pinto, L., Anzueto, M., Mendoza, J., Ferrer, G.J., de Jong, B., 2010. Carbon
sequestration through agroforestry in indigenous communities of Chiapas, Mexico.
Agroforestry Systems 78, 39-51.
Soto-Pinto, L., Villalvazo-Lopez, V., Jimenez-Ferrer, G., Ramirez-Marcial, N., Montoya,
G., Sinclair, F.L., 2007. The role of local knowledge in determining shade
composition of multistrata coffee systems in Chiapas, Mexico. Biodiversity and
Conservation 16, 419-436.

50. 39
Spigelski, D., 2004. Dietary diversity and nutrient adequacy in women of childbearing age
ina Senegalese peri-urban community. School of Dietetics and Human Nutrition.
McGill University, Montreal, CA.
Swindale, A., Bilinsky, P., 2006. Household dietary diversity score for measurement of
household food access: Indicator guide (v. 2). Food and Nutrition Technical
Assistance Project, Academy for Educational Development, Washington D.C.
Tao, T.C.H., Wall, G., 2009. Tourism as a sustainable livelihood strategy. Tourism
Manage. 30, 90-98.
Thrupp, L.A., 2000. Linking Agricultural Biodiversity and Food Security: The Valuable
Role of Sustainable Agriculture. International Affairs (Royal Institute of
International Affairs 1944-) 76, 265-281.
Tilman, D., Fargione, J., Wolff, B., D'Antonio, C., Dobson, A., Howarth, R., Schindler, D.,
Schlesinger, W.H., Simberloff, D., Swackhamer, D., 2001a. Forecasting
agriculturally driven global environmental change. Science 292, 281-284.
Tilman, D., Reich, P.B., Knops, J., Wedin, D., Mielke, T., Lehman, C., 2001b. Diversity
and productivity in a long-term grassland experiment. Science 294, 843-845.
Tosi, J., Hartshorn, G., 1978. Mapa Ecologico de El Salvador: Sistema de Zonas de Vida
del Dr. L.R. Holdridge (Ecological Map of El Salvador: Life Zone System of Dr.
L.R. Holdridge). Minsterio de Agricultura y Ganaderia de El Salvador (Agriculture
and Livestock Ministry of El Salvador), San Salvador.
Trujillo, L., 2008. Coffee production strategies in a changing rural landscape: A case study
in central Veracruz, Mexico. In: Bacon, C., Mendez, V.E., Gliessman, S.R.,
Goodman, D., Fox, J.A. (Eds.), Confronting the coffee crisis: sustaining livelihoods
and ecosystems in Mexico and Central America. MIT Press, Cambridge, MA,
U.S.A., pp. 69-98.
Tscharntke, T., Clough, Y., Bhagwat, S.A., Buchori, D., Faust, H., Hertel, D., Holscher, D.,
Juhrbandt, J., Kessler, M., Perfecto, I., Scherber, C., Schroth, G., Veldkamp, E.,
Wanger, T.C., 2011. Multifunctional shade-tree management in tropical
agroforestry landscapes - a review. J. Appl. Ecol. 48, 619-629.
Tscharntke, T., Klein, A.M., Kruess, A., Steffan-Dewenter, I., Thies, C., 2005. Landscape
perspectives on agricultural intensification and biodiversity - ecosystem service
management. Ecol. Lett. 8, 857-874.
USDA, 2012. Dietary reference intake tables. USDA National Agricultural Library.
Vandermeer, J.D., Lawrence, A., Symstad, A., Hobbie, S., 2002. Effect of biodiversity on
ecosystem function in managed ecosystems. In: Loreau, M., Naeem, S., Inchausti,

51. 40
P. (Eds.), Biodiversity and ecosystem functioning: synthesis and perspectives.
Oxford University Press, Oxford, UK, pp. 221-235.
Weibull, A.C., Ostman, O., Granqvist, A., 2003. Species richness in agroecosystems: the
effect of landscape, habitat and farm management. Biodiversity and Conservation
12, 1335-1355.
Westphal, S.M., 2008. Coffee agroforestry in the aftermath of modernization: Diversified
production and livelihood strategies in post-reform Nicaragua. In: Bacon, C.,
Mendez, V.E., Gliessman, S.R., Goodman, D., Fox, J.A. (Eds.), Confronting the
coffee crisis: sustaining livelihoods and ecosystems in Mexico and Central
America. MIT Press, Cambridge, MA, U.S.A., pp. 179-206.
Wunder, S., Börner, J., 2012. Payments for environmental services to mitigate climate
change: Agriculture and forestry compared. In: Wollenberg, E.K., Nihart, A.,
Grieg-Gran, M., Tapio-Biström, M.L. (Eds.), Climate Mitigation and Agriculture.
Earthscan, London, U.K.
Wünscher, T., Engel, S., 2012. International payments for biodiversity services: Review
and evaluation of conservation targeting approaches. Biological Conservation 152,
222-230.
Zimmerer, K.S., 2003. Geographies of seed networks for food plants (potato, ulluco) and
approaches to agrobiodiversity conservation in the Andean countries. Taylor &
Francis Inc, pp. 583-601.
Zimmerer, K.S., 2007. Agriculture, livelihoods, and globalization: The analysis of new
trajectories (and avoidance of just-so stories) of human-environment change and
conservation. Agric. Human Values 24, 9-16.

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CHAPTER 2: CULTIVATION OF MAIZE LANDRACES BY SMALL-SCALE
SHADE COFFEE FARMERS IN WESTERN EL SALVADOR
2.1. Abstract
Small-scale shade coffee agroecosystems have been noted for their potential for
tree, bird, and insect biodiversity conservation in the tropics. However, there is a lack of
research on other productive areas managed by small-scale coffee farmers such as
subsistence maize and bean (milpa) plots, which may be sites of important crop
biodiversity conservation, particularly through the on-farm cultivation of native landraces.
This study empirically examined the factors that influence farmers’ choices between
landraces and improved varieties of maize, how seed type interacts with management
decisions, and how yields of local maize landraces compare with improved varieties on the
farms of small-scale shade coffee farmers in western El Salvador. We conducted household
interviews and focus groups with the membership of a 29-household coffee cooperative
and tracked management and maize yields in the 42 milpa plots managed by these
households. Farmers planted both a hybrid improved variety and five local maize
landraces. ANOVA and Pearson’s chi-square test were used to compare household
characteristics, management, agroecological variables, and yields between plots planted
with landraces and plots planted with the improved variety. Logistic regression was used to
evaluate the strongest drivers of farmers’ choice between landrace seed and improved seed.
Analyses indicated that use of maize landraces was associated with higher household
income and steeper plot slope. Landrace maize and improved maize were not managed
differently, with the exception of synthetic insecticide use. There was no yield advantage

53. 42
for improved varieties over landraces in the 2009 growing season. Farmers appear to prefer
local maize landraces for milpa plots on more marginal land, and continue to cultivate
landraces despite the availability of improved seed. The farms of small-scale shade coffee
farmers could have substantial conservation potential for crop genetic diversity, and the
seed-saving and exchange activities among such farmers should be supported.
2.2. Introduction
Maize (Zea mays) is a critically important food crop in Latin America and much of
Africa and Asia (Smale et al., 2001). Along with beans (Phaseolus vulgaris), it provides
the sustenance for millions of people, particularly in rural areas, and is intricately tied to
social and cultural traditions (Keleman et al., 2009; Staller, 2010). The ecology and genetic
diversity of maize, the diversity and dynamics of maize populations, and the maintenance
of maize landraces have been well-studied, particularly among subsistence farmers in
Mexico, maize’s center of origin (Bellon, 1991; Bellon & Brush, 1994; Bellon et al.,
2003a; Bellon & Berthaud, 2004; Birol et al., 2009; Brush, et al., 2003; Brush & Perales,
2007; Keleman et al., 2009). In particular, the diversity of maize populations has been
subjected to intensive research due to a concern that widespread adoption of improved
varieties is causing the loss of maize diversity present in local landraces (van Heerwarden
et al., 2009). We use the term “improved” to refer to those varieties or cultivars of maize
that have been scientifically bred to be uniform and stable, as distinct from landraces
(Badstue, 2006). A landrace is defined as a population of a cultivated plant having
historical origin, distinct identity, and lacking formal crop improvement (Camacho Villa et
al., 2005). Landraces also tend to be genetically diverse, locally adapted, and associated